Poecilimon bosphoricus group (Orthoptera, Phaneropterinae): iteration of morpho-taxonomy by song...

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ZOOTAXAISSN 1175-5326 (print edition)

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Zootaxa 3225: 1–71 (2012) www.mapress.com/zootaxa/ Monograph

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Poecilimon bosphoricus group (Orthoptera, Phaneropterinae): iteration of morpho-taxonomy by song characteristics

SARP KAYA1, BATTAL ÇIPLAK2, 5 , DRAGAN CHOBANOV3 , KLAUS-GERHARD HELLER4

1 Department of Biology, Graduate School of Applied and Natural Sciences, Antalya, Turkey, E-mail: [email protected] 2 Department of Biology, Faculty of Science, Akdeniz University 07058 Antalya, Turkey, Tel: +90 242 310 23 56, Fax: +90 242 227 89

11. E-mail: [email protected] Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Tsar Osvoboditel boulevard 1, 1000 Sofia, Bul-

garia, E-mail: [email protected] Grillenstieg 18, 39120 Magdeburg, Germany, E-mail: [email protected]

5 Corresponding Author

Magnolia PressAuckland, New Zealand

3225

Accepted by D. Rentz: 21 Dec. 2011; published: 7 Mar. 2012

mailto:kaya_sarp@hotm

mailto:[email protected]


SARP KAYA, BATTAL ÇIPLAK, DRAGAN CHOBANOV, KLAUS-GERHARD HELLERPoecilimon bosphoricus group (Orthoptera, Phaneropterinae): iteration of morpho-taxonomy by song characteristics(Zootaxa 3225)

71 pp.; 30 cm.

7 Mar. 2012

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ISBN 978-1-86977-882-8 (Online edition)

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KAYA ET AL.2 · Zootaxa 3225 © 2012 Magnolia Press


Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Song . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Song . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Key to species (males only) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Subgroup 1: Species with syllable Type-1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Poecilimon sureyanus Uvarov, 1930 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Poecilimon kocaki Ünal, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Poecilimon athos Tilmans, F. Willemse & L. Willemse, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12Subgroup 2: Species with syllable Type-2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Poecilimon turcicus Karabağ, 1950 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Poecilimon turciae (Ramme, 1951) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Subgroup 3: Species with syllable Type-3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Poecilimon heinrichi (Ramme, 1951) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Poecilimon proximus Ünal, 2010 stat. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Poecilimon cervus Karabağ, 1950 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Poecilimon demirsoyi Sevgili, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Poecilimon bosphoricus Brunner von Wattenwyl, 1878 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Poecilimon bidens Retowski, 1889 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Poecilimon istanbul Ünal, 2010 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Poecilimon miramae Ramme, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Poecilimon roseoviridis Chobanov & Kaya sp.n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Poecilimon similis Retowski, 1889 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Poecilimon geoktschajcus Stshelkanovtzev, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Poecilimon djakonovi Miram, 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Poecilimon bischoffi Ramme, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Poecilimon scythicus Stshelkanovtzev, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Poecilimon tauricus Retowski, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Poecilimon pliginskii Miram, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25Morphology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Song. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Taxonomy and species distributions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

Abstract

More than 20 species were reported under the circum Black Sea lineage Poecilimon bosphoricus group (Orthoptera, Tet-tigonioidea, Phaneropterinae). The taxonomy of the group has for a long time been controversial; once these species were transferred to Eupoecilimon and many new species have been described since the revision by Ramme (1933) or synonyms have been suggested/re-established. This study aims to test the classification of the group presently based on morpholog-ical characters by bioacoustic data. The following results were obtained or conclusions arrived. First, several qualitative morphological characters previously used in descriptions/diagnoses of the species are variable and overlap between spe-cies. Those are the elevation and widening of pronotum in metazona, the emargination of caudal margin of pronotal disc and the structure of male subgenital plate at caudal margin. Thus, still the male cercus, especially the orientation of den-ticles, is the most productive structure may allow more objective delimitation of species. As in qualitative morphology the general morphometry seems uninformative for the taxonomy of the group. Second, male calling song and partly the num-ber of stridulatory pegs are more useful characters both for delimitation of species and describing their relationships. Es-pecially, the pattern of the syllable, the number of impulses per syllable and the duration of early part of syllable in species group allow us a more objective delimitation of the species and definition of relationships. Third, from the distribution and relationships of species, we suggested three radiation centres for the lineage: (1) Northwest Anatolia + Eastern Bal-kans, (2) Northeast Anatolia + Caucasus and (3) Crimea. Fourth, after evaluating morphological and song phenotypes we considered 21 species in P. bosphoricus group constituting three subgroups: (1) P. sureyanus and P. kocaki (+ P. athos),

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(2) P. turcicus + P. turciae and (3) P. bidens, P. bischoffi, P. bosphoricus, P. cervus, P. demirsoyi, P. geoktschajcus, P. hei-nrichi, P. istanbul, P. miramae, P. pliginskii, P. proximus, P. roseoviridis sp. n., P. scythicus, P. similis and P. tauricus (+ P. djakonovi). The following nomenclatural actions were made: (1) P. roseoviridis Chobanov & Kaya sp. n. described, (2)P. similis proximus Ünal, 2010 raised to species level as P. proximus stat. n., (3) P. naskrecki Ünal, 2001 syn.n. syn-onymised with P. demirsoyi Sevgili, 2001 (4) P. diversus Ünal, 2010 syn.n. and P. anatolicus Ramme 1933 syn.n. put in synonymy with P. sureyanus Uvarov, 1930, (5) P. oligacanthus Miram, 1938 syn.n. and P. tereckensis Stshelkanovtzev, 1910 stat.rev. resynonymised with P. similis Retowski, 1889, (6) P. beybienkoi Tarbinsky, 1932 syn.n. and P. kusnezoviMiram, 1929 syn.n. synonymised with P. tauricus Retwoski, 1888, and (7) P. boldyrevi Miram, 1938 syn.n. synonymised with P. pliginskii Miram, 1929.

Introduction

Poecilimon Fischer is one of the most speciose genera of Phaneropterinae (Tettigoniidae, Orthoptera) including around 150 species (Bei-Bienko 1954; Eades et al. 2011). It is distributed in Eurasia, but the core areas of its range are Anatolia and Southeast Europe. For example more than 55 of the Anatolian species are endemic (Çıplak et al. 1999). The high rate of endemism is possibly due to the radiation history of the genus. It is suggested that Poecili-mon radiated from some ancestral stocks in the Old Aegeid plate in Tertiary, but main radiation has occurred during glacial periods (La Greca 1999; Çıplak 2004). Such a radiation may cause a confusing taxonomy due to the follow-ing reasons. First, a recent radiation indicates young species that may be poorly differentiated from each other. Sec-ond, there may be several slightly diverged, but reproductively not yet isolated populations, some of which were erroneously described as distinct species. Third, range changes during climatic cycles might have caused isolation of some populations in one period, their secondary contact in the following period, and so finally their hybridiza-tion. All these possible events may cause a confusing classification and prevent advanced phylogenetic and phylo-geographic studies.

Ramme (1933) was the first who revised Poecilimon listing 68 species. After this date many new species have been described and this number has doubled (Bei-Bienko 1954; Heller et al. 1998; Çıplak et al. 1999; Ünal 2003, 2004, 2005, 2010; Eades et al. OSF—2011). However, in previous studies on Poecilimon, especially for the Anato-lian species, mostly the morphology has been considered in a traditional approach, and modern methods or approaches are rarely used. Only recently some species groups were studied (Chobanov & Heller 2010; Heller & Sevgili 2005; Heller et al. 2006, 2008; Zhantiev & Korsunovskaya 2005; Ullrich et al. 2010) using new techniques.

Ramme (1933) included 15 species in his Group VIII and later (Ramme 1951) transferred them to the genus Eupoecilimon Tarbinsky 1932, adding some species described thereafter. However, this classification has not been followed in subsequent publications and all these species were retransferred to Poecilimon (Bei-Bienko 1954; Harz 1969). These species share several similarities, especially those belonging to the index vertex/scapus, shape of pro-notum and male cerci (Fig. 1). The monophyly of this group is also supported by a molecular phylogenetic analysis (Ullrich et al. 2010). However, some of the characters used in species diagnoses are variable and the species cannot be easily distinguished from each other. For example, both elevation and width of metazona significantly vary within species and have been used in doubtful delimitation of species (e.g. compare Ramme 1951 for P. similis andP. richteri). Thus, it is necessary to reconsider the accuracy of characters used in species diagnoses.

The species united by Ramme (1951) in genus Eupoecilimon, which was later synonymised (see above), represent by our opinion a natural group of related taxa and following Ramme (1951) the group have to be named P. anatolicus group. However, according to the newly collected data (present paper), P. anatolicus has to be synonymised with P. sureyanus. Ünal (2010) has already used the name P. sureyanus group, though he listed here only few of the species formerly united in genus Eupoecilimon. He (l.c.) splits this natural grouping into few groups, including here only the species found in Turkey. He also refers only to morphology (including variable characters—see below) and thus, the latter grouping cannot reflect the natural relationships. Taking into account the published information (including molecular data—Ullrich 2010) and new information (see below) we outline the group and use the oldest published name for the group—P. bosphoricus.

The species P. flavescens (Herrich-Schäffer, 1838), type species of Eupoecilimon Tarbinsky, is certainly a member of the bosphoricus group. However, since the type (one female), which was figured and described by Fis-cher (1853), seems to be lost and the locality information is quite imprecise (“Südrußland” fide Ramme 1951= southern Russia), it has to be treated as species incerta, following Ramme (1933). Ünal (2010) proposes again to use the name Eupoecilimon as subgenus name for the bosphoricus group, but for such an action the relatonship to



other species groups should be known. According to Ullrich et al. (2010) the bosphoricus group is deeply nested among the species with mute females. The P. bosphoricus group is distributed in the circum-Black-Sea-Basin. However, the highest species number is concentrated to Anatolia and the Balkans (Ramme 1951; Harz 1969; Çıplak et al. 1999; Unal 1999, 2001, 2003, 2004, 2010). The remaining few species occur in Caucasus, Crimea and the North Black Sea Basin (Bei-Bienko 1954). In the present study the following taxa (the names valid until this study) are considered belonging to this group according to data by Ramme (1951), Bei-Bienko (1954) and our considerations.

1—P. anatolicus Ramme, 1933; NW Anatolia and Thrace2—P. athos Tilmans, F. Willemse & L. Willemse, 19893—P. beybienkoi Tarbinsky, 1932; Crimea4—P. bidens Retowski, 1889; Thrace5—P. bischoffi Ramme, 1933; NE Anatolia6—P. boldyrevi Miram, 1938; Crimea7—P. bosphoricus Brunner von Wattenwyl, 1878; NW Anatolia 8—P. cervus Karabağ, 1950; NW Anatolia9—P. diversus Ünal, 2010; NW Anatolia10—P. demirsoyi Sevgili, 2001; NW Anatolia11—P. djakonovi Miram, 1938; Caucasus12—P. geoktschajcus Stshelkanovtzev, 1910; Caucasus, Adzerbaijan13—P. heinrichi (Ramme, 1951); Thrace14—P. istanbul Ünal 2010; Thrace15—P. kocaki Ünal, 1999; NW Anatolia and Thrace16—P. kusnezovi Miram, 1929; Crimea17—P. miramae Ramme, 1933; NW Anatolia and Thrace18—P. naskrecki Ünal, 2001; NW Anatolia19—P. scythicus Stshelkanovtzev, 1911; Ukraine and southern Russia20—P. similis Retowski, 1889; NE Anatolia and Caucasus21—P. similis proximus Ünal, 2010; NE Anatolia22—P. sureyanus Uvarov, 1930; Anatolia23—P. tauricus Retowski, 1888; Crimea24—P. tereckensis Stshelkanovtzev, 1910; Caucasus25—P. turciae (Ramme, 1951); Anatolia and Thrace26—P. turcicus Karabağ, 1950; Anatolia and Thrace

Presently there is no key including all species of the group. The previous one (Bei-Bienko 1954) was not doubtlessly working, possibly due to variation in characters used, and is outdated due to many newly described species. Additionally some species (such as P. bidens and P. miramae) have never been clearly diagnosed or are diagnosed according to different species in different publications (compare Bei-Bienko 1954; Harz 1969; Ramme 1933). Accordingly, the distribution of several species is still not clear. In this study we aim to review the species of P. bosphoricus group using data from previous studies, studying museum collections and new material. The charac-ters frequently used in species diagnoses are herewith comparatively presented and their accuracy in species identi-fication is evaluated in the light of bioacoustic data. Available song data are given. A key to species of the group is prepared. Species distribution is redefined in the light of new data.

Material and Methods

MorphologySpecimens examined or type material are deposited in the following collections: (1) AUZM—Akdeniz Üniversite-si, Zooloji Müzesi, Antalya, Turkey (Collection of B. Çıplak), (2) CDC—Collection of D. Chobanov, (3) CH—Collection of K. G. Heller, (4) NHM—Natural History Museum, London, (5) ZMB—Museum für Naturkunde zu


http://orthoptera.speciesfile.org/Common/basic/Taxa.aspx?Taxo


Berlin, (6) NHMW—Naturhistorisches Museum, Vienna, (7) ZIN St. Petersburg, (8) CESA—Centre for Entomo-logical Studies Ankara and (9) AIBUEM—Abant Izzet Baysal University Entomological Museum, Bolu, (10) NMNHS—National Museum of Natural History, Sofia, (11) HMB—Historical Museum of Blagoevgrad. Addition-ally, photos of type specimens given in Orthoptera Species File—OSF2 (Eades et al. 2011) and SysTax-DORSA were examined.

Morphological structures were examined and measured using stereomicroscopes. In previous studies, descrip-tions and diagnoses of the species of P. bosphoricus group were based on qualitative examination of three struc-tures; the pronotum (mainly in male), the male subgenital plate and male cerci. To provide a more objective definition of these structures the terminology in previous studies is specified as follows: elevation and widening of the metazona according to prozona (Fig. 2), characteristics of the male cerci (Fig. 3) transformed to numerical data (measurements or coded character states), male anal tergite (Fig. 4) and apical structure of male subgenital plate (Fig. 5) from real material or photos obtained from the websites OSF2 and SysTax-DORSA. Although we have not coded any characters from female structures we provided illustrations for female subgenital plates and ovipositor.

SongFor sound recording in the laboratory a FOSTEX FR-2 (frequency response 22.05-192kHZ) digital recorder was used with a G.R.A.S. Type 40BF microphone (frequency response 10 Hz—40 kHz ± 1.0 dB, 4 Hz—100 kHz ± 1.0 dB). Other sound recordings were made with a Racal store 4 D tape recorder with microphones Brüel & Kjær 4133 and 4135 (frequency response flat up to 40 resp 70 kHz). Additionally, laboratory recordings are made with an electret condenser microphone (Knowles BT-1759-000) equipped with a custom-made preamplifier, connected to a PC through an external soundcard (TransitUSB, “M-Audio”) (96 kHz) and field recordings—using the same microphone connected to a ZOOM H2 recorder (ZOOM Corporation). Some songs were also recorded using an Uher tape recorder 4200 IC with microphone Uher M 645 (frequency response flat up to 20 kHz). Wing move-ments were registered by an opto-electronic device (Helversen & Elsner 1977, modified as in Heller 1988). Due to the heating effect of the registration lamp, only a lower limit for the body temperature can be given (see Heller 1988; measurements marked by “>”). Oscillograms and sound analysis were made using a PC and Turbolab (Stem-mer AG) and CoolEdit Pro. V. 2.0 (Syntrillium Software Corporation). In song descriptions the following terms were used (Fig. 6): Calling song—spontaneous song produced by an isolated male, syllable—the song produced by one opening-closing movement cycle of the tegmina, impulse—a simple, undivided transient train of sound waves, s—second, ms—millisecond.

Eight song characters were studied (Fig. 6; Table 3): (1) syllable duration (SD), (2) impulse number per sylla-ble (INS), (3) duration of early part of the syllable (DEP), (4) impulse number in the early part of the syllable (INE), (5) period of impulses in the early part of syllable (PIE), (6) duration of late part of the syllable (DSP), (7) impulse number in the late part of the syllable (INL), (8) period of impulses in the late part of syllable (PIL). To elucidate the differences between populations we produced two additional characters: (9) ratio of durations of the early/late parts (RDEL) and (10) ratio of the periods of the impulses in the early/late parts (RPEL). Since the spe-cies with syllable Type-1 (the populations representing P. sureyanus; and though song of P. kocaki and P. athoswere not available to include into analyses they are in syllable Type-1) do not clearly exhibits an early and a late part they were excluded from the analyses of characters 3–10. Additionally, P. turciae was excluded from statistical analyses due to insufficient number of the records, but later added to the graphs with the aim of comparison.

It is known that differences in recording temperatures are in potential of effecting temporal characteristics (durations and periods) of the song. Thus temporal and non temporal characters were analysed using different sta-tistical procedures. Each variable for each analysis group was tested for normality and homogeneity applying log or inverse transformation when required. Differences in values not affected by temperature (non-temporal INS, INE and INL; the rational RDEL and RPEL) were analysed using one-way ANOVA. The post-hoc analysis was done using Tukey test.

To determine the possible effect of the temperature and to exclude it series of statistical analyses were per-formed for five temporal parameters (SD, DEP, PIE, DSP and PIL). For the traits (SD, DEP, PIE, DSP and PIL), where the effect of temperature was found statistically significant (p<0.05), multiple regression analysis was per-formed including the temperature as a linear co-variate. k-1 (k, the number of populations) dummy variables were created in order to investigate the effect of each population. Empirical distribution of residuals was used to check the normality assumption of the error term, equal variance assumption, model over-fitting, model under-fitting, and



outlier detection. Estimation of the Outliers and influential observations were determined via Cook’s distance (Cook 1977) and Leverage statistics (HAT matrix) provided by the software. When an influential observation was detected it was dropped out from the analysis. Significant dummy variables (P< 0.05) for the populations were taken as evidence of differences between the populations (Suits, 1957). Determination coefficient was used to assess the model adequacy. To determine the most suitable regression model between possible competitive models we used Mallow’s Cp criterion (Mallows 1973), Akaike information cirterion (AIC) (Akaike 1973) and Bayesian (BIC) or also Swardz information criterion (Swardz 1978). The parameters which are obtained from the regression analysis were used to calibrate the temperature effect and eliminate the population differences on each song charac-ter. Standardized variables were analysed using one-way ANOVA. The pair-wise comparison of populations for these five characters was done using Tukey test. All these statistical analyses of song data were performed in SAS version 9.1.

Results

MorphologyThe P. bosphoricus group can be defined by the combination of following characters (Fig. 1); (i) tegmina in female rudimentary, not touching each other, (ii) small auditory spiracles, (iii) fastigium width roughly as 0.5 of scapus, (iv) robust male cerci with several apical denticles, (v) female lamella (basal fold of the lower ovipositor valve) dis-tinctly flattened and laterally protruded; (vi) song consisting of syllables which are clearly separated from each other; (vi) pronotum constricted in prozona and widened/elevated in metazona; (vii) typical coloration of the pro-notum with red lateral corners of metazona of the disc, a black spot/pattern along the hind margin or mid of the disc, (viii) cerci reddish brown with black apex at least along the denticles and (ix) tegmina yellow peripherally and black/brown in the middle. The species of this group resemble those of the P. ampliatus group (Heller & Lehmann 2004) of which they differ in characters (v), (iv), and partly (vi). Even bigger similarities are shared by the P. pro-pinquus group (Heller 1984, Lehmann 1988). The latter can be distinguished by less or not flattened female lamella (in P. bosphoricus group—lamella distinctly laterally protruded in different shape and reddish coloured), the more or less bulged metazone (constantly widened in P. bosphoricus group), coloration of cerci and tegmina and the distribution of male cercal teeth.

In previous publications the characters used in species descriptions/diagnoses belong mostly to three struc-tures; pronotum, male cerci and male subgenital plate. We defined three characters of male pronotum (Character 1–3 in Table 1; Figs 7–65); (i) structure of caudal margin (concave, convex or straight), (ii) width of metazona accord-ing to prozona and (iii) elevation of metazona according to prozona. All these characters are variable and one spe-cies alone may show two or three different states. None of the species can be distinguished from others by any character defined from pronotum except P. turcicus by the non elevated metazona (Table 1, Figs 8, 25, 39, 54). There is a correlation between elevation of metazona and covering the cubital vein of male tegmen by pronotum. Several species exhibit both character states, so the covered/non covered cubital vein cannot be doubtlessly used as a diagnostic character.

Male tenth tergum is uniform in the species group as in other Poecilimon except for P. tauricus, expressing variable development of protruding its medial part (formerly described for P. kusnezovi, syn.n.) (Fig. 4) The inci-sion of male subgenital plate at its caudal margin is another character which was commonly mentioned in previous studies. This character is not uniform per species since it is variable both in presence/absence and in the depth of incision. But the straight or somewhat convex (sometimes weakly incised) apical margin of male subgenital plate is characteristic to P. sureyanus and P. turcicus. Length of male subgenital plate according to cerci is also variable, but can be used to distinguish some species in combination with other characters (Table 1).

Male cerci are still the most productive structure in producing diagnostic characters, although caution is neces-sary when delimiting the species. We defined 12 characters belonging to male cerci most of which are not applica-ble to P. cervus, since cerci are with two apical branches in this species (Fig. 86). Of the other 11 cercal characters the robustness and the apical width are relatively subjective in description or variable in some species, so not very useful to diagnose the species. But the widened cercal apex is typical for P. turciae and P. heinrichi. The angle of curvature overlaps in several species. The number of denticles on each row is variable and may overlap in different species (see Table 1, Character 15, 16). Contrary to the numbers, the orientation of denticles and presence of one or



two enlarged apical denticles seem to be the most doubtlessly applicable key characters (Character 13, 14). In P. turcicus, P. bidens, P. pliginskii the cerci are denticulate only along the externo-apical margin. In P. demirsoyi (and P. naskrecki syn. n.) the cerci have two rows of denticles, one along the apico-external and the other along the apico-dorsal margins, both very close to each other. The cerci in P. cervus are somewhat similar to P. demirsoyi in respect to the orientation of each row. In P. roseoviridis sp. n. one row of denticles is located apico-externally; the second is slightly dorsally, but contrary to P. demirsoyi– rather close to the internal margin. In the other species the cerci have also two rows of denticles, but one along the apico-external and the other along the apico-internal mar-gin. The internal and external rows of denticles join at apex in several species exhibiting the latter character state excluding P. bosphoricus and P. kocaki. In P. scythicus and P. tauricus, (incl. P. kusnezovi and P. beibienkoi as syn-onyms) the internal row of denticles is rather apically disposed on a truncate projected margin.

Some metric data were gathered for the structures commonly measured in Orthoptera (Table 2) either from new material or obtained from literature. These structures are rather similar in the P. bosphoricus group and thus uninformative for its internal taxonomy (Table 2). Contrary to metric data, the number of stridulatory pegs provides some objective data. P. sureyanus, P. kocaki and P. athos are clearly different from others by the small number (48–87) of stridulatory pegs. The peg number in P. turcicus and P. turciae (89–117) is higher than the previous group, but smaller than that of the remaining species. The peg number in species other than given above ranges roughly from 120 (rarely less) to 215 and largely overlaps between species (Table 1). The highest counts belong to P. bos-phoricus, P. bidens and P. heinrichi. Although the peg number data are inadequate for some species diagnoses it is a potential character to distinguish certain taxa.

SongSong pattern is relatively similar at group level and consists of irregular syllable series. Three types of syllables are distinguishable. Syllable Type-1 is definable by the presence of low number of impulses, periods of which gradu-ally decrease from beginning to the end and is observed in populations representing P. sureyanus (Fig. 159). Sylla-ble Type-2 is definable by a long early and a short late part. The impulse number is higher and impulse period is longer in early part than those in the late. Additionally, the early part is clearly longer than the late part. Intensity of the impulses gradually increases from beginning and reach maximum at the end of the early part, then, gradually decrease towards end in the late part. This type of syllables is observed in P. turcicus and P. turciae (Figs 160, 161). Syllable Type-3 also consists of two subparts. Contrary to the Type-2, the period of the impulses in the early part is clearly shorter than that in the late part. Intensity of the impulses in the early part gradually increases from begin-ning toward the end of the part, but it is always lower than that of the impulses in the late part. Species other than mentioned above exhibit this type of syllable (Figs 162–174). The stridulatory movements by which these three syllable types are produced differ slightly. Syllable Type-1 and 3 are produced by a more or less continuous closing movement of the tegmina (Figs 6D, E, F; see also examples in Heller 1988), while in Syllable Type-2 the speed of the movement increases abruptly to produce the late part with its short impulse periods.

Independent of the song pattern, songs were examined in respect to the duration and the impulse number per syllable (data per populations were included into analyses independent of valid taxa). The multiple regression anal-yses suggested a significant correlation between the temperature and the values of temporal characters (Table 4). After calibration of the temperature affected values of temporal characters, data for all characters were analysed using ANOVA. The results suggested significant variation among populations for each of total 10 characters (Table 4). A Tukey test was performed using syllable duration, independent of syllable type, suggested four clusters of species (Table 3 and Fig. 175). P. proximus stat. n. constitutes one cluster by the shortest syllable duration. The remaining species are divided into three clusters (from the lowest to highest); the first includes P. cervus, P. demir-soyi, P. miramae, P. roseoviridis sp.n., P. bischoffi and P. similis; the third—P. heinrichi, P. bosphoricus and P. tau-ricus, and the second the remaining species. A Tukey test applied to the impulse number per syllable segregated seven clusters (Table 1 and Fig. 176). P. sureyanus represents one cluster by the least number of impulses. P. tur-ciae and P. proximus (possibly also P. turcicus, which has not been included into the statistical analyses due to insufficient number of measurements) constitute the second cluster with an impulse number higher than the previ-ous and lower than the next. Each of P. roseoviridis sp.n., P. heinrichi and P. pliginskii represents a single cluster. P. bosphoricus and P. bischoffi constitute another cluster. The last cluster shows a great variation in impulse number per syllable and includes the remaining nine species.



We performed further statistical tests for nine different parameters of the syllable Type-3. The two subparts of the syllable are indistinguishable in P. sureyanus. Additionally, the impulse pattern per syllable is different in P. turcicus and P. turciae. Including these species into statistical analyses together with the species exhibiting Type-3 may cause a bias, thus, they were excluded from the following analyses. First, a Tukey analysis applied to the duration of early part of the syllable suggested eight distinct clusters; from lowest to the highest values these are: (1) P. proximus, (2) P. miramae (Bulgaria), (3) P. cervus, P. miramae (Turkey), P. roseoviridis sp.n., P. demirsoyi (+ P. naskrecki) and P. similis, (4) P. bischoffi, (5) P. kusnezovi syn.n., and P. pliginskii, (6) P. scythicus, (7) P. heinrichiand (8) P. bosphoricus and P. tauricus (Fig. 177). Second, the analysis of the impulse number in the early part of the syllable resulted in three distinct clusters; (1) P. proximus, (2) P. roseoviridis sp.n. and (3) the remaining 14 populations with a great variation (Fig. 178). The third analysis was for the impulse period in the early part of the syllable and it separated P. scythicus, P. tauricus and P. kusnezovi, from the remaining populations which also exhibited great variation (Fig. 179). The fourth and the fifth analyses included the duration and the impulse number in the late part of the syllable and both segregated P. kusnezovi from all others (Figs 180, 181). The Tukey test applied to the period of impulses in the late part of the syllable segregated P. miramae, P. roseoviridis sp.n. and P. proximus from the remaining taxa (Fig. 182). The ratio of durations of the early/late parts suggested three different clusters; (1) P. proximus and P. miramae (Bulgaria) at the lower end, (2) P. tauricus at the highest end, and (3) the remaining all in the middle (Fig. 183). The ratio of periods of impulses in the early/late parts suggested two distinct clusters, the first including P. roseoviridis sp.n., P. kusnezovi syn.n. and P. tauricus and the second all others (Fig. 184).

TaxonomyIn the light of the morphology and song data, which were studied on the basis of populations, the species listed below were considered in the P. bosphoricus group. Remarks for taxonomic considerations are given along each species. A key to species is presented following the species list. Data produced in the concept of present study were insufficient for P. athos Tilmans et al., 1989, P. djakonovi Miram, 1938 and P. geoktschajcus Stshelkanovtzev, 1910(resembling P. similis) and thus we do not include them into the key.

1—P. athos Tilmans, F. Willemse & L. Willemse, 19892—P. bidens Retowski, 1889 3—P. bischoffi Ramme, 19334—P. bosphoricus Brunner von Wattenwyl, 1878 5—P. cervus Karabağ, 19506—P. demirsoyi Sevgili, 2001 (=P. naskrecki Ünal, 2001 syn. n.)7—P. djakonovi Miram, 1938 8—P. geoktschajcus Stshelkanovtzev, 19109—P. heinrichi (Ramme, 1951)10—P. istanbul Ünal, 201011—P. kocaki Ünal, 199912—P. miramae Ramme, 193313—P. pliginskii Miram, 1929 (=P. boldyrevi Miram, 1938 syn. n.)14—P. proximus Ünal, 2010 stat. n.15—Poecilimon roseoviridis sp. n. 16—P. scythicus Stshelkanovtzev, 191117—P. similis Retowski, 1889 (=P. tereckensis Stshelkanovtzev, 1910 stat. rev., P. oligacanthus Miram, 1938 syn. n.)18—P. sureyanus Uvarov, 1930 (=P. anatolicus Ramme, 1933 syn. n. and P. diversus Ünal, 2010 syn. n.)19—P. tauricus Retowski, 1888 (=P. kusnezovi Miram, 1929 syn. n., P. beybienkoi Tarbinsky, 1932 syn. n.)20—P. turciae (Ramme, 1951)21—P. turcicus Karabağ, 1950


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Key to species (males only)

1a Male cerci not branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21b Male cerci with two branches apically (Fig. 86) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .cervus2a Elevation of metazona constitute an angle <160°; cercal apex not as in Figs 67, 84 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32b Elevation of metazona constitute an angle >170° (Figs 39), 54; cercal apex as in Figs 67, 84 . . . . . . . . . . . . . . . . . . . . turcicus3a Apico-internal row of the denticles (1–3) strongly detached from apex leaving a gap in between (Figs 71, 83, 88) . . . . . . . . . 43b Both of apico-internal and apico-external rows joint to each other (Figs 66, 68–70, 72–82, 85, 87, 89–93). . . . . . . . . . . . . . . . 54a There is one detachet denticle on apico-internal row (Figs 71, 88) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . bosphoricus4b There are 2–4 detachet denticles on apico-internal row (Fig. 83). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .kocaki5a Cerci with one row of the denticles along the apico-external margin, one-three denticles on proximal part of the row are dis-

tinctly larger than the others (Figs 72, 79–81, 89) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65b Cerci with two rows of denticles, all denticles more or less similar in size (Figs 66, 68–70, 73–79, 82, 85, 87, 90–93) . . . . . . 86a The cercal apex not distinctly widened and the externo-apical row of denticles lies on slightly convex edge (Anatolia +Thrace)

(Figs 72, 89) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .bidens6b The cercal apex is distinctly widened and the externo-apical row of denticles lies on a concave or sinuately shaped edge

(Crimea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77a Cerci with 1–2 large apical and 7–17 small denticles (Fig. 81); the apical denticles does not seem to be placed on a protruded

part of the cercus; the subgenital plate usually does not protrude over cerci; Eastern Crimean Mountains. . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pliginskii (incl. boldyrevi syn.n.)

7b Cerci with 1–6 larger apical and 9–14 smaller denticles (Figs 79, 80); the apical denticles are placed on a protruded part of the cercus; the subgenital plate protrude over cerci; Western Crimean Mountains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tauricus (incl. kusnezovi and beybienkoi syn.n.)

8a Of the two rows of denticles, one located on the apico-external and the other on the apico-internal row (Figs 66, 68, 69, 73–78, 82, 85, 90–93) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

8b Both rows of denticles located on the apico-external margin parallel to each other (Figs 70, 87) . . . . . . . . . . . . . . . . . demirsoyi9a Male stridulatory file with more than 90 pegs (mostly more than 110; Table 1), if it is less than this number then apex of cerci

is distinctly widened (Fig. 68); song is of syllable Type-2 or Type-3 (Figs 160–174) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109b Male stridulatory file with less than 90 pegs (less than 87; Table 1); song is of syllable Type-1 (Fig. 159) . . . . . . . . . sureyanus10a The apical internal and external margins of cercus (and thus rows of denticles) joint to each other at a right or an obtuse angle

(Figs 68, 69) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1110b The apical internal and external margins of cercus (and thus rows of denticles) joint to each other at an acute angle (Figs 73–

80, 85, 90–93) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1211a Male stridulatory file with 190–210 pegs (Table 1), syllable is of Type-3 (Fig. 162) . . . . . . . . . . . . . . . . . . . . . . . . . . .heinrichi11b Male stridulatory file with ca. 98 pegs (Table 1), the syllable is of Type-2 (Fig. 161). . . . . . . . . . . . . . . . . . . . . . . . . . . . turciae12a Cerci not acutely tapered to a pointed apex; the inner row of the denticles is shorter than or as long as the outer one (Figs 73–

76, 78–80, 85, 90–92) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1312b Cerci acutely tapered to a pointed apex; the inner row of the denticles is longer than the outer one (Figs 77, 93) . . . . . bischoffi 13a Externo-apical row of denticles straight or even concave (Figs 73, 78–80, 85) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1413b Externo-apical row of denticles convex (Figs 74–76, 90–92) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1714a Syllable duration is more than 50 ms (usually over 100 ms) (Table 3); cerci other than in Fig. 85. . . . . . . . . . . . . . . . . . . . . . 1514b Syllable duration is 50 ms or less (Table 3); cerci as in Fig. 85 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . proximus15a Cerci enlarged at apex (Figs 73, 78–80) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1615b Cerci not enlarged or tapering at apex (Fig. 78) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .scythicus16a Cerci with 2–3 denticles on inner row (Fig. 73) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . istanbul16b Cerci with no denticles on the inner row; if sometimes some denticles tend to locate toward the inner margin, then they are on

a projecting apex (Fig. 79, 80). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tauricus (kusnezovi/beybienkoi syn.n.)

17a Pronotum weakly elevated in metazona, its hind margin straight or slightly concave (Figs 15, 16, 34, 35, 45, 46, 62–63); cerci more or less flattened at apex (Figs 74, 75, 90, 91), if cerci not distinctly flattened apically then the apico-internal row of den-ticles located dorsalward (Fig. 74, 75, 90, 91) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

17b Pronotum strongly elevated in metazona, its hind margin strongly concave (Figs 17, 36, 47, 64); cerci rounded at apex and gradually but distinctly pointed to apex (Figs 76, 92). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .similis

18a Stridulatory row with more than 140 pegs; the internal row of denticles is not dorsalward (Figs 74, 90) . . . . . . . . . . . .miramae18b Stridulatory row with less than 140 pegs; cerci with internal row of denticles located dorsalward (Fig. 75, 91) . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . roseoviridis sp.n.



Subgroup 1: Species with syllable Type-1.Syllable Type-1 (Fig. 6A) is definable by the presence of low number of impulses, period of which gradually decrease from beginning to the end and is observed in P. sureyanus (including its synonyms) (Fig. 159). Congruent with this syllable type, the number of stridulatory pegs in these species is lower that those exhibited by the species producing the other two syllable types (Table 3). Song of P. kocaki was not available to include in statistical analy-ses during preparation the manuscript. However, it also consists of Syllable Type-1 (data available after the manu-script was finished).

Poecilimon sureyanus Uvarov, 1930(Figs 7, 23, 38, 52, 66, 82, 94, 109, 122, 141, 159, 175, 176, 185)

Poecilimon sureyanus Uvarov, 1930: 350.Eupoecilimon sureyanus (Uvarov); Ramme, 1951: 339.Poecilimon sureyanus Uvarov; Bei-Bienko, 1954: 337–338.Poecilimon anatolicus Ramme, 1933: 555. syn.n.Poecilimon diversus Ünal, 2010: 159. syn. n.Poecilimon rufonitens Ingrisch & Pavicevic, 1985: 51–54. syn. with anatolicus by Heller, 1988: 62. Tilmans et al. 1989: 29.

Type information: Holoype—male; Turkey, Izmit (Kocaeli) (NHM). Photos of the type specimens from OSF examined.

Material examined: TURKEY: 1 male, 1 female syntypes of P. anatolicus; Bursa province—4 males, 3 females, Ulufer Cayiy south of. Ulu Dagh (40° 05’ N, 29° 04’ E), 400 m, 21–22.7.1983 (leg. K.G. Heller), 11 males, 2 females , Uludağ Mt., (40°7’ N, 29°6’ E), 1600–2000 m, 23–24.7.1983, (leg. K.G. Heller), 4 males, 4 females, Uludağ Mt., Uludağ National Park, 22.7.1998 (leg. B. Çıplak); 3 males, 2 females, 1800 m, 22.7.2003 (collector unknown); 3 females, 1903 m, 27.7.2006 (collector unknown); 14 males, 9 females, 1732–1903 m, (40° 07’ N, 29° 08’ E) 29.6.2008 (leg. S. Kaya et al.); 1 male, 11.8.2008 (leg. G. Deniz & D. Sirin), 9 males, 7 females, 1732 m, 18.7.2009 (leg. B. Çiplak et.al); 4 males, 2 females, 5 km nordwestl. Inegöl (40°6'N, 29°29'E), 360 m, 24.7.1983, (leg. K.G. Heller), 2 males, 2 females, Keles-Inegöl road, 1319 m (39° 56’ N, 29°18’ E) 19.7.2009 (leg. B. Çiplak et al.); 2 males, 1 females, Orhaneli, 555 m (40° 02’ N, 28° 53 E,) 29.6.2008 (leg. S. Kaya et al.); 1 male, Kestel, 4.6.1998 (leg. B. Çıplak); Balıkesir province—5 males, 8 females, Edremit Kazdagi Mt., 1950 m, 20.7.1998 (leg. B. Çıplak); 1 male, 4 females, 23.7.2001 (leg. A. Mol) (all in AUZM), 17 males, 11 females, Edre-

mit Kazdagi Mt., (39o 42’ N, 026o 53’ E) 1230 m. 05.07.2011, (leg. B. Çıplak, S. Kaya) (in alcohol in AUZM); Bil-ecik province—12 males, 5 females, Bozhöyük, Yirice Yaylası1700–1900m, 29.7.2000 (leg. A. Mol) (AUZM), 22 males, 11 females Bozüyük, Kömürsu yaylası, (39° 51’ N, 29° 44’ E,) 1630 m, 12.8.2010 (leg. B. Çıplak, S. Kaya & Z. Boztepe) (in alcohol in AUZM). GREECE: Kavalla province—5 males, 2 females Umg. Lekani (41° 09’ N, 24° 33’ E), 870 m, 27.6.1980 (leg. K.G. Heller & M. Volleth), 4 males, 1 female Pangaion Geb. 1100 m, 1.8.1982 (leg. K.G. Heller); 1 male, 1 female Pangaion Geb. 1100 m, 24.7.2001 (leg. K.G. Heller); Drama province—3 males, 2 females Kato Vrondou nordöstl. Serrai (41° 16’ N, 23° 44’ E), 1.6.1983 (leg. O. v. Helversen)

Description: For morphology see Table 1, 2, Figures Figs 7, 23, 38, 52, 66, 82, 94, 109, 122, 141; for song see Table 3 and Figures 159, 175, 176. Additional description can be found mainly in Ramme (1933, 1951) and Bei-Bienko (1954).

Distribution (Fig. 185): Known from Northwest Anatolia, Thrace and Greek Makedhonia (Ramme 1933, 1951; Bei-Bienko 1954; Karabağ 1958; Willemse 1984, 1985; Naskrecki 1991; Heller 1984 (as cf. similis), 1988: the record of Gelibolu is based on a mis-identification and refers in fact to P. turciae); Ingrisch & Pavicevic, 1985; Sevgili 2001; Ünal 1999, 2003, 2004, 2005, 2010; Willemse 1984, 1985; Willemse & Willemse 2008). Ramme (1933), based on the specimens by Adelung (1907), reported the presence of P. sureyanus in Artvin province of Turkey in Northeast Anatolia. In the light of the material examined the specimens from Artvin are referred to P. similis and thus P. sureyanus occurs only in northwest Anatolia.

Remarks: This species (and its synonyms) well differs from all others by the male calling song and the lower number of stridulatory pegs. The male calling song consisting of a few impulses and the stridulatory file with 48–87 pegs are characters not shared by other species except P. kocaki and P. athos (song not known). Although P. ana-tolicus is suggested to be different from P. sureyanus by the less number of cercal denticles there is no clear cut and



the number of denticles on both of internal and external margin overlaps. The apex of male cerci also does not pres-ent any species specific structure and different types can be found in the specimens from the same locality. All these statements are also valid for the recently described P. diversus (Ünal, 2010). Although P. diversus is slightly different in cercal apex, the shape of the latter remains in the variation limits of a single species. Especially, the song of a P. diversus like population (recently published by Sevgili et al. 2011 as P. anatolicus) is very similar both in the pattern and in temporal parameters. Additionally, a preliminary analysis of the COI sequences supports this statement (Çıplak et al.: unpublished data). Thus, P. anatolicus Ramme, 1933 and P. diversus Ünal, 2010 are put in synonymy with P. sureyanus Uvarov, 1930.

Poecilimon kocaki Ünal, 1999(Figs 24, 53, 83, 110, 123, 142, 185)

Poecilimon kocaki Ünal, 1999: 244, 245, 253.

Type information: Holotype—male; Turkey, Sakarya, Geyve, Şerefiye (AIBUEM).Description: For morphology see Table 1, 2 and Figures 26, 55, 84, 110, 122, 136. Additional description can

be found mainly in Ünal (1999).

Material examined: TURKEY: Adapazarı province—3 males, 2 females between Gevye Şerefiye (40o 34’ N, 030o 19’ E) 74 m, 07.07.2011, (leg. B. Çıplak, S. Kaya & D. Chobanov).

Distribution (Fig. 185): Endemic to northwest Anatolia (Ünal 1999, 2005).Remarks: Ünal (1999) reported this species to be most similar to P. bosphoricus but later he (Ünal 2010) men-

tioned it in his P. sureyanus group. The inner row of denticles not joining to the apico-external row (separated by a non denticulate gap) may indicate a synapomorphy of P. kocaki and P. bosphoricus but according to other mor-phocharacters and song, it is rather a homoplasy. In P. kocaki the inner row contains 3–4 denticles, while in P. bos-phoricus it has only a single huge one. However, song of P. kocaki (it was available after the manuscript finished) includes syllable Type-1 as in P. sureyanus.

Poecilimon athos Tilmans, F. Willemse & L. Willemse, 1989(Fig. 186)

Poecilimon athos Tilmans, F. Willemse & L. Willemse, 1989: 26–30.

Type information: Holotype—male; “Hellas, Athos, 1500 m/Mt. Athos, Panaya,/ leg. J.M. Tilmans, 30/31-VII-1983 (Collectio Tilmans)”; Allotype female, “Hellas (Athos)/Mt. Athos 1200–1500 m/L. Willemse 18-VII-1986 (Collection F. Willemse)”. Paratypes: 1 male (Collectio Tilmans) labelled as holotype, 5 males, 1 female (Collec-tion F. Willemse), 1 male, 1 female (Natural History Museum, London), 1 male (Instituut voor Taxonomische Zoölogie, Amsterdam) labelled as holotype, 3 females (Collection Tilmans), 1 female (Instituut voor Taxonomis-che Zoölogie, Amsterdam), “Hellas, Athos, 900–1200 m/Mt. Athos, below Panayia,/leg. J.M. Tilmans, 30/31-VII-1983”; 1 female (Collectio Tilmans) “Hellas, Athos, 1300 m/Mt. Athos, below Panayia,/leg. J.M. Tilmans 31-VII-1983”; 3 males (Collectio F. Willemse) “Hellas (Athos)/Mt. Athos 850–1150 m/L. Willemse 19-VII-1986”; 2 males, 2 females (Collection F. Willemse) “Hellas (Athos) 1500–/2000 m Mt. Athos ab. Panayia/L. Willemse 18-vii-1986”; 1 female (Collectio F. Willemse) “Hellas (Athos)/600 m Kerasia/L. Willemse 18-VII-1986.

Description: See Tilmans, F. Willemse & L. Willemse (1989).Distribution (Fig. 186): As far known the species occurs on the peninsula of Akti (Athos), part of the

Khalkidiki Peninsula in Northern Greece.Remarks: Though the authors (Tilmans, F. Willemse & L. Willemse 1989) have difficulties to define the clos-

est relatives of this species, according to the available morphological description and figures P. athos is probably closely related to P. sureyanus subgroup. The species is rather similar to P. sureyanus by the shape of male cerci and number of stridulatory pegs, yet differing by the short male subgenital plate and, to some extent, by the female lamella.

Using the synapomorphies, shared by this species and P. sureyanus, we tentatively place the species under Sub-group 1. New data is needed for evaluating its phylogenetic position within the group.



Subgroup 2: Species with syllable Type-2Syllable Type-2 (Fig. 6B) is definable by a long early and a short late part. The impulse number is higher and the impulse period is longer in the early part than those in the late one (the ratio of the periods of the impulses in early part:late part of the syllable is more than 4). Intensity of the impulses gradually increases from beginning and reach maximum at the end of the early part, then, gradually decrease towards the end of the late part. This type of sylla-bles is observed in P. turcicus (Fig. 160) and P. turciae (Fig. 161) and species specific song parameters can be found in Table 3 and Figs 175–184. The number of the stridulatory pegs in these two species is higher than those with syllable Type-1, but is lower than those with the Type-3.

Poecilimon turcicus Karabağ, 1950(Figs 8, 25, 39, 54, 67, 84, 95, 111, 124, 143, 160, 175–185)

Poecilimon turcicus Karabağ, 1950: 152.Poecilimon turcicus Karabağ; Bei-Bienko, 1954: 375.

Type information: Holoype—male; Turkey, Bursa; Karacabey (NHM) (photo in OSF examined).Material examined: TURKEY: Edirne province—5 males, 5 females, Havsa, (44° 33’ N, 26° 49’ E),

11.7.2009 (leg. B. Çıplak) (AUZM); Tekirdağ province—1 male, 15 km südl. Corlu (41° 04’ N, 27° 47’ E), 1–31.5.1996, (leg. Holger Braun), Kırklareli province—1 male, 10 km westl. Lüleburgaz (Abzweigung nach Saricaali) (41° 25’ N, 27° 15’ E); 1–31.5.1996, (leg. Holger Braun); GREECE: Lesbos/Mytilini province—8 males, 5 females Umg. Larissos (Kolpos Geras) (39° 07’ N, 26° 26’ E), 28.5.1993, (leg. K.-G. Heller)

Description: For morphology see Table 1, 2 and Figures 8, 25, 39, 54, 67, 84, 95, 111, 124, 143; for song see Table 3 and Figures 160, 175–184. Additional description can be found in Ramme (1951) and Bei-Bienko (1954).

Distribution (Fig. 185): Known from Northwest Anatolia (including the island of Lesbos) and Thrace (Karabağ 1950; Naskrecki 1991; Sevgili 2001; Ünal 2004; Willemse & Willemse 2008).

Remarks: The prominent autapomorphy of this species is the typical apex of cercus with a single row of denticles apico-externally and the tip bearing a large tooth. However, it is similar to P. turciae by sharing the presence of sylla-ble Type-2, as well as to P. sureyanus + P. kocaki + P. athos by the smaller number of stridulatory pegs (see Table 1).

Poecilimon turciae (Ramme, 1951)(Figs 9, 26, 40, 68, 125, 161, 175–185)

Eupoecilimon turciae Ramme, 1951: 339.Poecilimon turciae (Ramme); Bei-Bienko, 1954: 326–327.

Type information: Holotype male and allotype female; Turkey, Bursa (ZMB).Material examined: TURKEY: Holotype—male; Çanakkale province—1 male, 1 gynander, Umg. Gelibolu

(40° 24’ N, 26° 40’ E), 09.5.1985 (leg. E. Bluemm).Description: For morphology see Table 1, 2 and Figures 9, 26, 40, 68, 125; for song see Table 3 and Figures

161, 175–184. Additional description can be found in Ramme (1951) and Bei-Bienko (1954).Distribution (Fig. 185): Known from Northwest Anatolia (Ramme 1951; Sevgili 2001; Ünal 1999, 2003,

2004, 2005, 2010).Remarks: This species is very similar to P. heinrichi in the shape of its cerci. However, they are clearly differ-

ent from each other in the number of stridulatory pegs and the syllable pattern (see Tables 1, 3 and Figures 175–184). The low number of stridulatory pegs and the syllable Type-2 are shared by P. turciae and P. turcicus.



Subgroup 3: Species with syllable Type-3Syllable Type-3 (Fig. 6C) also consists of two subparts (see Figs 161–174). Contrary to the Type-2, the ratio of the periods of the impulses in early:late part is less than 2 (more than 4 in syllable Type 2). Similarly, the impulse num-ber in the early part of the syllable is roughly twice the number in the late part. The intensity of impulses in the early part gradually increases from beginning toward the end of the part, but it is always lower than that of the impulses in the late part. The following species exhibit this syllable type; P. proximus, P. bosphoricus, P. miramae, P. roseoviridis sp.n., P. cervus, P. demirsoyi, P. similis, P. bischoffi, P. heinrichi, P. pliginskii, P. scythicus and P. tauricus. As in the other two groups, the number of stridulatory pegs is similar in this group and is higher than in the other two groups. For further details and differences between species in the subgroup see Table 3 and Figures 175–184. There are two prominent indications in these data. First, differences between species occur in the early part of the syllable. Second, of the species given above, P. proximus, P. heinrichi and to some extend P. tauricus are different from others in several song parameters, while others are relatively similar.

Songs of P. bidens, P. istanbul, and P. geoktschajcus were not included into statistical analyses and are not presented in this study. Although this is so, recently available records indicate that P. bidens produces a song of this type. P. istanbul is closely related to P. bosphoricus by several morphological features, thus, possibly it belongs to Subgroup-3. Morphological data for P. geoktschajcus, P. tereckensis (song data for these taxa can be found in Zan-thiev & Kursunovskaya 2005) and P. oligacanthus are considered in this study according to previous literature (Ramme 1933; Bei-Bienko 1954) and photos of type material in OSF (Eades et al. 2011). In our oppinion these species constitute forms of P. similis (for taxonomical consideration see under P. similis) and are thus also consid-ered in this subgroup.

Poecilimon heinrichi (Ramme, 1951)(Figs 10, 27, 41, 55, 69, 96, 126, 144, 162, 175–184, 186)

Eupoecilimon heinrichi Ramme, 1951: 340.Poecilimon heinrichi (Ramme); Bei-Bienko, 1954: 328–329

Type information: Holotype—male, 2 females (allotype and paratype), Bulgaria, Strandja-Balkan, b. Konak (ZMB).

Material examined: SE BULGARIA: Strandzha Mts—1 male (holotype), 2 females (allotype, paratype), Byala Voda (Konak) Village, 350 m [Ramme 1951] (41° 10’ N, 27° 27’ E), 7.1935 (G. Heinrich; W. Ramme det. P. miramae; ZMB); 1 male, Fakiya Village (42° 11’ N, 27° 05’ E), 11.7.1974 (leg. G. Peshev; NMNHS); 6 males, 3 females, 1 male nymph last instar, Gradishteto Peak, 400–600 m (41° 57’ N, 27° 29’ E), xeromesophyte under-growth (Apiaceae gen.sp., Rubus sp.) in xerothermic oak and xeromesic Fagus orientalis—wood, 26.6.2007, rare (leg. D. Chobanov; NMNHS); 2 males, 1 female, Kovach Place between Malko tarnovo and Zvezdets Villages [labelled “Strandzha”] (42° 05’ N, 27° 25’ E), 5.7.1955 (leg. M. Josifov; NMNHS); observed, Brushlyan Village, 300 m (42° 02’ N, 27° 26’ E), 25.6.2007 (obs. D. Chobanov); 1 male, Malko Tarnovo, 16.6.1963; 4 males, same place, 18.6.1973 (both G. Peshev; NMNHS); 1 female, N of Malko Tarnovo, 350 m (42° 00’ N, 27° 31’ E), mes-opyte herbal and bush associations in mesophyte forest, 26.5.2005, nymphs 4th–5th instar on Urtica dioica (leg. D. Chobanov; CDC); 14 males, 1 female, Chernogorovo Place S of Malko Tarnovo, 330 m (41° 57’ N, 27° 33’ E), 19.6.1973; 3 males, Kostina Luka Place near Malko Tarnovo, 18.6.1973 (both leg. G. Peshev; NMNHS).Other material: Poecilimon sp. aff heinrichi; SW BULGARIA: Strouma Valley— 1 male, Melnik, 25.–26.6.1969 (leg. J. Deckert; A. Lehmann det. P. heinrichi; ZMB).

Description: For morphology see Table 1, 2 and Figures 10, 27, 41, 55, 69, 96, 126, 144; for song see Table 3 and Figures 162, 175–184. Additional description can be found in Ramme (1951), Bei-Bienko (1954) and Harz (1969).

Distribution (Fig. 186): Known from Strandzha Mts at the border of NW Turkey and SE Bulgaria (Ramme 1951, Buresch & Peshev 1958; Karabağ 1958; Peshev & Djingova 1974). The locality in Southwest Bulgaria remains doubtful for the slight morphological differences of the specimen concerned.

Remarks: This species was described from Istranca (Strandzha) Mts. at the border of Turkey and Bulgaria. It is very similar to P. turciae, known from Northwest Anatolia, in the shape of male cerci. Ramme (1951) diagnosed these species by two characters; the width of cercal apex (also mentioned in the key) and the length of male sub-



genital plate. Yet, there are other differences leading to the presumption that the above-mentioned similarities of these two species are not homologous. First, one of the highest numbers of stridulatory pegs belong to P. heinrichi, while that of P. turciae is much lower. Second, the syllable pattern and the other temporal parameters of the song of P. heinrichi are distinctly different from those of P. turciae. The latter species is similar to P. turcicus in respect to song parameters and stridulation structures. Since the traditional characters are rather variable in several species we think that the sister species status suggested in previous publication (Ramme 1951; Bei-Bienko 1954) is not valid.The only specimen recorded outside the Strandzha Mountains was collected in Southwest Bulgaria (see above). This specimen differs slightly in the shape of male cerci and colouration of the typical P. heinrichi yet we refrain from judging about its taxonomic identity until new material will be available.

Poecilimon proximus Ünal, 2010 stat. n.(Figs 28, 56, 85, 112, 163, 175–185)

Poecilimon similis proximus Ünal, 2010: 158.

Type information: TURKEY: Bolu, Aladag yolu, 1225 m, 40°39’601” N, 31°38’398” E, 5.7.2001 (AIBUEM).Material examined: TURKEY: Bolu province— 2 males, road from Bolu to Düzce, Bolu Mt pass, (40° 44’

N, 31° 24’ E), 962 m, 14.8.2010 (leg. B. Çıplak, S. Kaya & Z. Boztepe) (in alcohol in AUZM).Description: For morphology see Table 1, 2 and Figures 28, 56, 85, 112; for song see Table 3 and Figures 163,

175–184. Additional morphological description can be found in Ünal (2010).Distribution (Fig. 185): Ünal (2010) reported his new species from several localities in Northwest Anatolia. Remarks: This species was described as a subspecies of P. similis by Ünal (2010). However, apart of the gen-

eral characteristics of P. bosphoricus group, there are no synapomorphic characters shared by P. proximus and P. similis except the syllable Type-3. However, of the 16 species producing syllable Type-3 the most aberrant syllable pattern and temporal parameters are demonstrated by P. proximus (see Figs 163, 175–184). Thus, it is raised to a species level.

Poecilimon cervus Karabağ, 1950(Figs 29, 57, 86, 113, 127, 145, 164, 175–185)

Poecilimon cervus Karabağ, 1950:153

Type information: Holotype—male; Turkey, Ankara, Nallihan (NHM).Material examined: TURKEY: Bolu province—1 male (nymph), 3 females (2 nymphs), Bolu to Yedigöller

National Park (40° 52’ N, 31° 40’ E), 1713 m, 20.7.2009 (leg. B. Çıplak, S. Kaya & E. M. Korkmaz); 21 males, 16

females, Road from Bolu to Yedigöller National Park, (40° 53’ N, 31° 40’ E), 1670 m, 15.8.2010 (leg. B. Çıplak, S. Kaya & Z. Boztepe) (in alcohol in AUZM); 1 male Yığılca (40o 55’ N, 031o 23’ E), 663 m, 08.07.2011, (leg. B. Çıplak, S. Kaya & D. Chobanov), 8 males, 5 females (40o 58’ N 031o 33’ E), 499 m 08.07.2011, (leg. B. Çıplak, S. Kaya & D. Chobanov), 22 males, 17 females, Yedigöller National Park Mengenden road (40o 59’ N 031o 37’ E), 900 m, 08.07.2011, (leg. B. Çıplak, S. Kaya & D. Chobanov) (in alcohol in AUZM); 1 male, “Iskilip”, 28.6.1951; 1 female, Ankara, Nalhkay, 28.6.1949 (both leg. T. Karabağ) (NHMW).

Description: For morphology see Table 1, 2, Figures 29, 57, 86, 113, 127, 145; for song see Table 3 and Fig-ures 164, 175–184. Additional description can be found in Karabağ (1950), Ramme (1951) and Bei-Bienko (1954).

Distribution (Fig. 185): Endemic to northwest Anatolia.Remarks: P. cervus is the most aberrant species of the group by the male cercus with two apical branches. The

transversal elongation of cerci at apex and the apico-dorsally located two rows of cercal denticles are shared also by P. demirsoyi (including its new synonym P. naskrecki). More prominent statistical analyses suggested no differences between P. cervus and P. demirsoyi (plus its synonym P. naskrecki) in all song parameters. However, they are considered as distinct species by their different cercal morphology. Although P. cervus and P. demirsoyiwere found to occur allopatrically in some localities, in Boludag Mts these species were found along an altitudinal



gradient. At lower elevations we found typical P. naskrecki, at the top typical P. cervus, but at mid elevations some forms similar to P. demirsoyi occurred. More importantly, a preliminary phylogenetic analysis of COI sequences (Çıplak et al.: unpublished) indicates a very weak divergence between P. cervus and P. demirsoyi (plus its synonym P. naskrecki).

Poecilimon demirsoyi Sevgili, 2001(Figs 11, 30, 42, 58, 70, 87, 97, 114, 128, 146, 165, 175–185)

Poecilimon demirsoyi Sevgili, 2001: 18. Poecilimon naskrecki Ünal, 2001: 1–4. syn. n.

Type information: Holotype—Male; Turkey, Kastamonu, Ilgaz Mts. 1600-1850 m (HUBM).Material examined: TURKEY: Bolu province— 19 males, 3 females, Road to Yedigöller National Park, 855

m (40° 51’ N, 31° 34’ E), 30.6.2008 (leg. S. Kaya et al.); 2 male, 2 females, 1713 m, (40° 52’ N, 31° 40’ E), 20.7.2009 (leg. B. Çıplak et al.); 2 males, 2 females, Road from Bolu to Yedigöller National Park, (40o 52’ N, 31o

40’ E), 1291–1519 m, 15.8.2010 (leg. B. Çıplak, S. Kaya & Z. Boztepe) (in alcohol in AUZM). Description: For morphology see Table 1, 2 and Figures 11, 30, 42, 58, 70, 87, 97, 114, 128, 146; for song see

Table 3 and Figures 165, 175–184. Additional description can be found in Ünal (2001) and Sevgili (2001).Distribution (Fig. 185): Endemic to northwest Turkey (Ünal 2001, 2003; Sevgili, 2001). Remarks: The typical character of this species is the location of the rows of cercal denticles similar to that of

P. cervus, both parallel, placed apico-externally one over the other (in other species of the group the second row is internally disposed). The apico-dorsal row of denticles is typical for both P. naskrecki and P. demirsoyi which were described in the same year (Ünal 2001; Sevgili 2001). From the description by Sevgili (2001) P. demirsoyi is simi-lar to P. cervus by the outer apical projection of cerci, which is somewhat similar to the prominent projection of the latter. However, this character seems to be unuseful in diagnosing these species. Thus, the characters used to diag-nose these species seem to represent local variations (See Remarks section of P. cervus) and objective data suggest P. demirsoyi and P. naskrecki to be synonyms.

Deciding about the name to be retained requires determining the day of publications since both descriptions were published in the same year. The issue of Centre For Entomological Studies Miscellaneous Papers including the paper by Ünal (2001) was published on 2nd of July (according to the information in that issue) and therefore later than the issue of the Journal of Orthoptera Research that includes the contribution by Sevgili (according to information in that issue: June 2001). Thus, the priority has to be given to P. demirsoyi including the name P. naskrecki as a junior synonym.

Poecilimon bosphoricus Brunner von Wattenwyl, 1878(Figs 12, 31, 43, 59, 71, 88, 98, 115, 129, 147, 166, 175–185)

Poecilimon bosphoricus Brunner von Wattenwyl, 1878: 43.Eupoecilimon bosphoricus (Brunner von Wattenwyl); Ramme, 1951: 417.Poecilimon bosphoricus Brunner von Wattenwyl; Bei-Bienko, 1954: 342.

Type information: Male holotype, 5 males and 3 females paratypes—Turkey, Istanbul, both sides of Bosphorus (NHMW).

Material examined: 1 male (holotype), 4 males, 3 females (paratypes), “Giöksu (Bosph.), Br.v.W.”; 1 male (paratype), “Therapia” (all NHMW); [1 female (labeled as paratype) without locality label is not P. bosphoricus but was identified as P. cf. marmaraensis, det. D. Chobanov,]; 1 male, “Konstant., Türk 1870”; 1 male, “Konstant., Br.W.”; 1 male, 1 female, “Constantinopel, Belgrader Wald vii.00, Dr. Werner”; 1 male P. bosphoricus/bidens, det. D. Chobanov, “Belgrader Wald, Eahringer, 26.vii.1910”, Ramme det. as P. bosphoricus; (all NHMW); TURKEY: Bolu province—1 male, 1 female, Aladağ Mt., 1600 m, 22.7.2004 (leg. A. Mol); 8 males, 15 females, Kartalkaya, Mt., 1674–2026 m, (40° 39’ N, 31° 46’ E), 21.7.2009 (leg. B. Çıplak et al.); Kastamonu province—9 males, 4 females , Ilgaz-Geb. (41°3'N, 33°40'E), 1800–2000 m, 15–16.8.1983 (leg. K.-G.Heller), 21 males, 12 female, Ilgaz



Tosya road, Ilgaz pass (41o 07’ N, 034o 04’ E) 1626m 10.07.2011, (leg.: B. Çıplak, S. Kaya & D. Chobanov ) (in alcohol in AUZM); Çankırı province—11 males, 4 females, Ilgaz Mt., 1900 m, 10.8.1996 (leg. B. Çıplak); 3 females, 1980 m, 20.7.2003 (leg. A. Mol); 5 males, Işıkdağı Mt., 1350 m (40° 77’ N, 32° 87’ E), 22.6.2008 (leg. B. Çıplak); Istanbul province—1 male, Kilyos 19.7.1977 (leg. S. Salman) (AUZM); 3 males, 3 females, 40 km NE Istanbul, 25 km of Sile, 115 m, undergrowth of Quercus wood, 2.7.1997 (leg. S. Petrov; CDC).

Description: For morphology see Table 1, 2 and Figures 12, 31, 43, 59, 71, 88, 98, 115, 129, 147; for song see Table 3 and Figures 166, 175–184. Additional description can be found in Ramme (1933), Bei-Bienko (1954) and Harz (1969).

Distribution (Fig. 185): Known from northwest Anatolia and Thrace (Brunner von Wattenwyl 1878; Ebner 1919; Bei-Bienko 1954; Karabağ 1958; Karabağ et al. 1971; Heller 1988; Naskrecki 1991; Sevgili 2001; Ünal 1999, 2003, 2004, 2005, 2010).

Remarks: P. bosphoricus and P. kocaki are similar by sharing the apico-internal row of denticles not joint to the apico-external one, but they well differ in syllable pattern. In P. bosphoricus the cerci with a huge single denti-cle on internal row, but sometimes there are two or three (in one male one cercus with one and the other with two) denticles. In such cases P. bosphoricus and P. istanbul can not be distinguished from each other.

Poecilimon bidens Retowski, 1889(Figs 13, 32, 44, 60, 72, 89, 99, 116, 130, 148, 185)

Poecilimon bidens Retowski, 1889: 221 (Ramme, 1933)Eupoecilimon bidens (Retowski) Ramme, 1951: 417.Poecilimon bidens (Retowski); Bei-Bienko, 1954: 351.

Type information: Holotype—male; Turkey, Istanbul, Bosphorus, Büyükdere (probably lost, not in ZIN St Petersburg). When Retowski (1889) described bidens as a variety of P. bosphoricus he mentioned “most of them around Bosphorus near to Bujuk-Dere” without any indication to the number of the specimens. When Ramme (1933) raised it to a species level he based his description on 5 males and 2 females (Ramme leg. 1914) from the same locality (photos of two males used by Ramme in ZIN St Petersburg provided by A. Gorochov and O. Korsunovskaya).

Material examined: TURKEY: 1 male, 1 female (locality unclear, possibly near Sile, 7.1997) (leg. S. Petrov; CDC).

Description: For morphology see Table 1, 2 and Figures 13, 32, 44, 60, 72, 89, 99, 116, 130, 148. Additional description can be found mainly in Ramme (1933), Bei-Bienko (1954) and Harz (1969).

Distribution (Fig.185): Known from Thrace and Northwestern Anatolia (Retowski 1889; Ramme 1933; Bei-Bienko 1954; Ünal 2004, 2010).

Remarks: This species was originally defined as a variety of P. bosphoricus by Retowski and later raised to species level by Ramme (1933). The subsequent publications refer to the description by Ramme (1933). P. bidensand P. bosphoricus differ from each other mainly by the shape of male cerci. All denticles (including the huge api-cal one) of male cerci are located on one margin in P. bidens, while the large denticle is located on the inner margin with a gap between those on apico-external margin in P. bosphoricus. Although the position of the large apical tooth was thought to be a species specific character, we observed some specimens intermediate for this feature between P. bosphoricus and P. bidens. The variation of the latter character was observed also in the Crimean taxa (P. pliginskii (=P. boldyrevi) and P. tauricus (=P. kusnezovi + P. beybienkoi) (see below).

Poecilimon istanbul Ünal, 2010(Figs 14, 33, 61, 73, 100, 117, 149, 185)

Poecilimon istanbul Ünal, 2010: 160.

Type information: Holotype—male; Turkey-in-Europe, Istanbul, European side, Fenertepe (AIBUEM).Description: For morphology see Table 1, 2 and Figures 14, 33, 61, 73, 100, 117, 161, 149. Additional

description can be found in Ünal (2010).



Material examined: TURKEY: European Turkey—2 males, 1 female, ”Belgrader Wald bei Istambul June 1954 Candon, 1955” (NHMW).

Distribution (Fig.185): Known from Thracian Turkey (Ünal 2010). Remarks: Ünal (2010) described this species for some specimens earlier reported by him as P. bidens (Ünal

2004). Description of the species largely overlaps with that of P. bosphoricus and P. bidens also from the European side of the Bosphorus. However, it differs from these two by; (1) presence of 1–5 denticles on interno-apical row of cerci, which are in similar size with the rest (against 1 large denticle in P. bidens and P. bosphoricus) and (2) absence of a gap between the two rows of cercal denticles. Thus, the species diagnose will remain unclear until song of this taxon is available.

Poecilimon miramae Ramme, 1933(Figs 15, 34, 45, 62, 74, 90, 101, 118, 132, 150, 167, 175–184, 186)

Poecilimon miramae Ramme, 1933: 563, 564.Eupoecilimon miramae (Ramme); Ramme, 1951: 417 (comb.n. in table).Poecilimon miramae Ramme; Bei-Bienko, 1954: 339–340 (partim).Poecilimon miramae Ramme; Harz, 1969: 159, 162 (partim).

Type information: Holotype—male; European Turkey, İstanbul, Kilyos on the Black Sea near Bosphorus (NHMW).

Material examined: TURKEY: European Turkey— Holotype male, 1 male (paratype), “Europ. Türkei 1911, Kilia, Fahringer”, 1 male, 1 female (paratypes), “Europ. Turkey, Jarim Burgas, 8.viii.1909, Fahringer”; 1 male (paratype), “Kleinasien, Jalova, 31.vii.1910”; 1 female paratype, 1 male paratype (OSF) “Europ. Türkei, Kütschük Tschekmedje, Fahringer 1911”; 1 female P. cf. miramae, “Kleinasien, Göck Daghi, Fahringer”, Ramme det. as Poecilimon sp. (NHMW); Sakarya province—1 male, 1 female, Sapanca Gölü südl. Adapazari (40°41'N, 30°24'E), 15.6.1986 (leg. Otto von Helversen); BULGARIA: E Rhodope Mts—3 males, 5 females, Mandritsa Village, 80 m, mesophyte ruderalized associations of Sambucus ebulus (41° 23’ N, 26° 07’ E), 05.6.2005 (collected as nymphs last instar and imago); 13 males, 5 females (leg. D. Chobanov; CDC), 3 males, 1 female (leg. D. Chob-anov; NMNHS), Mandritsa Village (41° 23’ N, 26° 07’ E) and 4 km N of it (41° 25’ N, 26° 07’ E), 60–80 m, mes-ophyte associations of Sambucus ebulus, 22.6.2008 (common).

Description: For morphology see Tables 1, 2 and Figures 15, 34, 45, 62, 74, 90, 101, 118, 132, 150; for song see Table 3 and Figures 167, 175–184. For species description see Ramme (1933). The additional descriptions and previous records by Bei-Bienko (1954) and Harz (1969) are herewith referred to two species—P. miramae and P. roseoviridis sp.n. (see below). The situation appears from the description of P. miramae by Bei-Bienko (1954), who uses specimens belonging to the new species but identified by Ramme as P. miramae. Thus, Bei-Bienko (1954) and Harz (1969) mixed the features of these two taxa using both the original description by Ramme (1933) and the material used by Bei-Bienko (l.c.). After comparing the type material of P. miramae preserved in the NHMW, we could diagnose P. miramae and P. roseoviridis sp.n. (see below).

Diagnosis: P. miramae resembles morphologically P. bidens in the shape of male pronotum, the variably shaped male subgenital plate and colouration. From the latter it differs in the few (1–5) small apico-internal denti-cles on male cerci (Fig. 74, 90), while in P. bidens there is a single large apical denticle (Fig. 72, 89). However, P. miramae was recently mixed with an undescribed species of the group, P. roseoviridis sp.n., P. miramae differs from P. roseoviridis sp.n. in the shape of pronotum, tegmina, number of stridulatory pegs, male subgenital plate, female lamellae (see Diagnosis of P. roseoviridis sp.n. below).

Distribution (Fig. 186). This species occurs on both sides of the Bosphorus, along Marmara Sea and the Dar-danelles up to the easternmost part of the Rhodope Mountains in Bulgaria and Greece (along Byala and Luda Reka/Erithropos Rivers) (Willemse 1982; own new data).



Poecilimon roseoviridis Chobanov & Kaya sp.n.(Figs 16, 35, 46, 63, 75, 91, 102, 119, 133, 151, 168, 174–184, 186)

Type material examined: Holotype, male; Bulgaria, Strandzha Mts, Malko Tarnovo, 18.6.1973, leg. G. Peshev (Strandja | M. Tarnovo | 18.6.1973). Paratype: BULGARIA: N Black Sea coast— 4 males, 1 female, Pobitite Kamani place near Varna (43° 13’ N, 27° 42’ E), 8.7.1969 (leg. G. Peshev; NMNHS); S Black Sea coast—8 males, Arkoutino Lake [Peshev & Djingova 1974] (42° 20’ N, 27° 43’ E), 19.6.1963; 5 males, Ropotamo Reserve [Peshev & Djingova 1974] (42° 19’ N, 27° 44’ E), 19.6.1963; 1 male, same place, 23.6.1963; 1 female, same place, 24.6.1966; 5 males, 3.7.1964; 11 males, 5 females, 7.7.1965; 52 males, 15 females, 8.7.1964 (all leg. G. Peshev; NMNHS); 1 male, 1 female, Beglik Tash place near Primorsko (42° 17’ N, 27° 45’ E), mesophyte herb associations (Urtica dioica, Rubus sp., Melissa officinalis) in mesophyte deciduous forest (Quercus sp., Ulmus minor и др.), 9.6.2005, nymphs 5th–6th instar, imago (leg. D. Chobanov; CDC); 19 males, 16 females, Veleka River Estuary [Peshev & Djingova 1974] (42° 04’ N, 27° 58’ E), 15.6.1968; 1 male, Nessebar—the forestry, 13.7.1968 (both leg. G. Peshev; NMNHS); Strandzha Mts—1 male [possibly Fakiya Village, 11.7.1974 or Veleka Estuary, 17.7.1974], wrongly labelled as Pirin, Banderitsa Lodge; 5 males, Fakiya Village (42° 11’ N, 27° 05’ E), 11.7.1974 (leg. G. Peshev; HMB); 4 males, 1 female, Gradishteto Peak, 600–700 m (41° 57–58’ N, 27° 29’ E), xeromesophyte under-growth (Apiaceae gen.sp., Rubus sp.) in xerothermic oak and xeromesic Fagus orientalis—wood, 26.6.2007; 3 males, 1 female, Brushlyan Village, 300 m (42° 02’ N, 27° 26’ E), 25.6.2007 (both leg. D. Chobanov; CDC); 3 males, 4 females, Byala Voda (Konak) Village, 350 m [Bei-Bienko 1954 as P. miramae] (41° 10’ N, 27° 27’ E), 7.1935 (G. Heinrich; W. Ramme det. P. miramae; ZMB); 6 males, Malko Tarnovo, 18.6.1973, 2 males, same place, 8.7.1967; 8 males, 2 females, same place, 14.7.1975 (leg. G. Peshev; NMNHS); 5 males, 4 females, N of Malko Tarnovo, 350 m (42° 00’ N, 27° 31’ E), mesopyte herbal and bush associations in mesophyte forest, 26.5.2005, nymphs 4th–5th instar; 3 males, same place, 11.6.2005, common—imago and nymphs 5th and 6th (last) instar on Urtica dioica (both leg. D. Chobanov; DCD); 2 males, 1 female, Chernogorovo Place S of Malko Tarnovo, 330 m (41° 57’ N, 27° 33’ E), 19.6.1973 (leg. G. Peshev; NMNHS); observed (rare), same place, xeromesophyte scrub of Quercus sp., Prunus spinosa, Carpinus orientalis, Rosa sp., 25.6.2007 (obs. D. Chobanov); 28 males, 2 females, Kostina Luka Place near Malko Tarnovo, 18.6.1973; 1 male, 1 female, Kachoul Place SW of Gramatikovo Village (42° 01’ N, 27° 37’ E), 20.6.1973 (both leg. G. Peshev; NMNHS).

Other localities: TURKEY: European Turkey, Istrandja Mts— Demirkoy (specimen in OSF online; as P. miramae).

Description: Male (Figs 16, 35, 46, 63, 75, 91, 102, 119). Medium sized for the genus, over medium sized for the group. Integument moderately shiny to dull.

Head: fastigium of vertex well produced anteriorly with a groove, its lateral margins parallel or slightly con-verging, its width half the width of scapus.

Thorax: pronotum long (both pro-and metazona elongated), moderately constricted in prozone, metazona gradually but strongly widened and considerably raised with its surface slightly domed; metazona long, covering the tegmina at least to the stridulatory vein. Posterior margin of pronotum gently concave. Lower margin of prono-tal lateral lobe slightly wavy with small nose-like projection in the frontal part. Auditory spiracle very small, its diameter smaller than the width of apical limbs of palpi. Tegmina conspicuously bulged for the group, reaching or slightly extending over the hind edge of first abdomunal tergite. The stridulatory file has comparatively sparse pegs, smoothly but fastly getting bigger from the apex to 1/5th to the base and become largest in the midlle of the file. The stridulatory row is about 2.6–2.9 mm long and bears 120–136 stridulatory pegs. Fore femur slightly longer than pronotum. Hind femora lack ventral spines.

Abdomen: margins of tergites straight or slightly convex. Cerci (Fig. 75, 91) wide in the base but generally comparatively slender and long for the group; they are almost straight in the basal 1/3–1/4 and slightly incurved apically; the apex is slightly bulged just before the end and then gently tapered to the end from both external and internal margins; the apex has two rows of fine teeth 8–11 externally and 5–6 dorso-internally. Subgenital plate (see Fig. 119) is long, extending over the tip of cerci, smoothly tapering to a narrow, slightly triangularly concave hind margin.

Colouration. General colouration pale-green to greenish with pink hue. Vertex, pronotum (except metazona), tergites and legs with big dark brown spots. Antennae yellowish-green, annulated with dark rings. Head and prono-tal dorsum with two lateral and a medial thin pale (almost white) stripes, the lateral ones are broadening backwards



in metazona; such sripes but intermittent and greenish may be observed on abdominal tergites. Dorsolateral corners of pronotum in metazona with big reddish-pink spots almost touching each other at the hind margin of metazona. Tegmina yellow with dark brownish-black stridulatory area. Femora and tibiae green getting yellowinsh or pink apically. Abdomen green with the first and sometimes all tergites with a black spot mediobasally. Cerci and partly the 10th abdominal tergite reddish-pink. Cerci with black teeth and rarely the apical fourth of cerci darkened.

Female (Figs 133, 151). Pronotum almost cylindrical but slightly flattened and widened in metazona, which has straight or insignificantly concave hind margin. Pronotal lateral plates as in male and straight dorsally with the transverse sulcus cutting the median line after the middle of pronotum. Tegmina fully covered by pronotum, reduced to scale-like appendages. Cerci surpassing the epiproct, bulged in the basal 3/2–3/4 and strongly attenuated apically being slightly incurved. Subgenital plate short, transverse, widely oval, apically blunt (fresh material observed). Ovipositor more than twice longer than pronotum, apical part with short, stout teeth dorsally, ventrally and laterally, as in allied species. Lamella (basal fold of dorsal margin of lower ovipositor valve) (Fig. 151) short, flattened, laterally widened, bearing a short, nose-like projection (observe from below). Colouration: as in male, usually reddish-pink colour less conspicuous.

Measurements—mm (mean±stdev) for 10 individuals each structure: pronotum (along the median line) ♂ 4.4–5.2 (4.8±0.2), ♀ 4.5–5.4 (4.8±0.3); male tegmen (visible part) 1.8–2.9 (2.4±0.4); hind femur ♂ 14.4–17.0 (15.6±0.8), ♀ 15.8–17.4 (16.5±0.6); male subgenital plate 3.9–4.9 (4.3±0.3); ovipositor (from base of lamella to tip) 5.0–6.1 (5.6±0.3).

Bioacoustics. Male calling song (Fig. 168; Table 3) consists of a sequence of syllables of type 3 (see above). At 26 °C the syllables last 94–152 ms and have 31–68 impulses. The early syllable part lasts 50–113 ms and has 10–53 impulses with a period between 2.0 and 7.7 ms; the late part is 13–51 ms long and has 10–23 impulses with a period of 0.4–2.3 ms. The song frequencies have a peak at 21.5–25.2 kHz (for additional information see Table 3).

Diagnosis: Poecilimon roseoviridis, sp.n., is a typical representative of the P. bosphoricus group having wid-ened metazona, typical reddish-pink pattern on pronotum and yellowish-brown tegmina as well as cerci ending with many teeth. The species cannot be easily distinguished similarly to many others within this group. By the shape and colouration of pronotum and body in general it is most similar to P. istanbul; by the shape of cerci it resembles most P. similis and some specimens of P. miramae differing from both in denser and much higher num-ber of teeth (see Table 1) and from P. miramae in the dorso-apical position of the inner row of teeth against externo-apical in miramae. Male subgenital plate is similar to that in some specimens of P. miramae though in the latter its apical excision is variable and might be quite deep, while in P. roseoviridis it has very shallow excision. Female subgenital plate is widely rounded and thus similar to many species of the group, mostly resembling P. miramae. However, from the latter the new species differs in the shape of lamellae (lateral processes of the base of the lower ovipositor valve), which are wider and shortly pimple-like protruded, while in P. miramae they are more widely oval and not so laterally expanded (see Fig 151). The song of the new species resembles that of P. miramae differ-ing mostly in the lower number of impulses having longer periods in the early part of the syllable (but similar char-acteristics of the late part) and thus the lower mean impulse number of the whole syllable and the significantly higher ratio of the periods of the syllables in the early/late part of syllables (about 1.5 against 0.5–1.0 in P. mira-mae) (compare Table 3, Figs 167, 168, 175–184).

Generally, P. roseoviridis seems to be most closely related to P. miramae differing from it in the following fea-tures; body generally larger; pronotum longer with wider and slightly dome-shaped metazona lacking black pattern at the hind edge (present in many specimens of miramae); pronotum covers the stridulatory vein of male tegmina, while in P. miramae the stridulatory vein is visible; male tegmina are strongly bulged with higher (wider) Costo-medial area; rare appearance of yellow area on the first abdominal tergite in female (common for P. miramae); lower number of stridulatory pegs (compare Table 1); the structure of the base of female ovipositor (see above).

Distribution and ecology. The new species occurs in SE Bulgaria and NW Turkey ranging from the region of Varna in North along the Black Sea coast to Strandzha (Istrandja) Mountains, where it is best represented (Fig. 186). It inhabits the undergrowth of the typical Pontic mesophyte deciduous forests in the region, keeping on high herbs (Urtica dioica, Sambucus ebulus etc.) or low bush-or tree branches. In Strandzha Mountains it frequently coexists with P. heinrichi. The nymphs emerge in March-April and the imago occurs from late May to July.

Etymology. Named after its typical colouration.



Poecilimon similis Retowski, 1889(Figs 17, 36, 47, 64, 76, 92, 103, 120, 134, 152, 169, 175–184, 186)

Poecilimon similis Retowski, 1889: 220. (Ramme 1933).Poecilimon caucasicus Adelung, 1907: 136. syn. by Bei-Bienko (1954).Poecilimon tereckensis Stshelkanovtzev, 1910: 52, syn. by Bei-Bienko (1954), confirmed.Poecilimon richteri Ramme, 1933: 557. syn. by Unal (2010).Poecilimon oligacanthus Miram, 1938: 351, syn. n.Poecilimon paramonovi Ramme, 1951: 343, syn.n. (after synonymisation of P. oligacanthus).Eupoecilimon similis (Retowski); Ramme, 1951: 342. Poecilimon similis (Retowski); Bei-Bienko, 1954: 334–337.

Type information: Syntypes—1 male and 1 female Sinop-Turkey and 1 male from Batum—Georgia (unknown). Although three syntypes were mentioned by Retowski (1889), Bei-Bienko (1954) suggested Batum-Georgia as the type locality. Since some other species occur in Sinop province of Turkey (northwest Anatolia) and the Caucasus seems to be the core area of the range of P. similis we consider the decision by Bei-Bienko (1954) appropriate.

Material examined: ARMENIA: Kotayk province—1 female, 15 km ne Geghard, Mt. Azhdahak (40°13'N, 44°53'E), 2800 m, 18.8.2005, (leg. Mark Kalashian), 1 male, 5 km n Arzakan, Aghveran gorge (40°30'N, 44°37'E), 1820 m, 11.7.2005, (leg. Mark Kalashian); Syunik province—1 male, 10 km se Lernadzor (39°7'N, 46°14'E), 15.8.2005, (leg. Mark Kalashian), 1 male, 5 km w Goris (39°30'N, 46°20'E), 4.7.2005, (leg. Mark Kalashian); Tavush province—1 male, 1–2 km s Dilijan (40°43'N, 44°52'E), 11.7.2005, (leg. Mark Kalashian), 1 male, 1 female, 5–8 km sse Dilijan (40°43'N, 44°50'E), 6.8.2005, (leg. Mark Kalashian); GEORGIA: West Georgia— 2 males, “Kaukasus, Ober-Jvanefien, Beco” = Becho, Svaneti, 10.8.1913 (leg. J. Komarek; NHMW); Transcauca-sia, Borjomi district—1 males, Bakuriani Resort (41°45’ N, 43°32’ E), 1600–2000 m, 10.–13.9.1927 (leg. W. Ramme; ZMB), 1 male, 1 female, same place and date (leg. W. Ramme; NHMW), 1 female, Tbilisi, Tsodoreti lake (41°46'N, 44°39'E), 2005, (leg. David Tarknishvili); TURKEY: Ardahan province—3 males, 2 females, Pass östl. Ardahan (41°7'N, 42°59'E), 2000 m, 10.8.1983, (leg. K.-G. Heller), 2 males, 1 female, Çam-Pass, Ostseite (41°4'N, 42°19'E), 2200–2400 m, 10–11.8.1983, (leg. K.-G. Heller); 1 male Çıldır road (41o 07’ N, 042o 54’ E), 1940 m, 15.07.2011, (leg. B. Çıplak, S. Kaya, D. Chobanov, E. Mahir Korkmaz); Artvin province—1 females, Ardanuç, 350 m, 25.6.1974 (leg. S. Salman); 2 males, 1 female, Borçka, 150–520m, 24–30.7.1974 (leg. S. Sal-man), 1 male, 1 female, Şavşat, 1800 m, 5–10.8.1974 (leg. Z. Bahçeci); 3 females, Yalnizcam-Pass (41°4'N, 42°18'E), 9.8.1987, (leg. K. Reinhold), 3 male, 1 female, Yalnizcam-Pass, Nordwestseite (41°4'N, 42°16'E), 1200–1800 m, 12.8.1983, (leg. Heller), 21 males, 23 females, Şavşat (41o 14’ N, 042o 22’ E), 1226 m, 15.07.2011, (leg. B.

Çıplak, S. Kaya, D. Chobanov, E. Mahir Korkmaz), 2 males, 5 females, Ardahan road, 41o 13’ N, 042o 24’ E), 1640 m, 15.07.2011, (leg. B. Çıplak, S. Kaya, D. Chobanov, E. Mahir Korkmaz); Giresun province—1 male, Bergtal bei Tandere (Teknecik) (40°31'N, 38°23'E), 1500 m, 1.8.1983, leg. K.-G. Heller; 5 males, 2 females, Gebirge zwi-schen Kümbet u. Tandere (40°32'N, 38°23'E), 1200–1400 m, 1–2.8.1983, (leg. K.-G. Heller); Kars province—1 female, Yalnızçam, 2100 m, 20.8.1974 (leg. S. Salman); Ordu province—1 male, Kumru, 1130m, 3.7.2004 (leg. A. Mol); 1 female, Unye-Niksar, 1350 m, 31.7.2005 (A. Mol); Rize province—4 males, 1 female, İkizdere—Cimil 1851 m (40°44’ N, 40°44’ E), 24.7.2005 (leg. B. Çıplak et al.), 4 males, 1 female, Flusstal unterhalb Ikizdere (40°47'N, 40°32'E), 420 m, 3.8.1983, (leg. K.-G. Heller) (all in CH or AUZM).

Description: For morphology see Table 1, 2 and Figures 17, 36, 47, 64, 76, 92, 103, 120, 134, 152; for song see Table 3 and Figures 169, 175–184. Additional description can be found mainly in Ramme (1933, 1951) and Bei-Bienko (1954).

Distribution (Fig. 186): This is possibly the most widespread species of the P. bosphoricus group, range of which covers nearly the whole eastern half of the Black Sea basin in Anatolia and the Caucasus (Adelung 1907; Avakyan 1981; Heller 1988; Karabağ 1958; Salman 1978; Chernyakhovskiy 1994, Chernyakhovskiy et al. 1994), though not all of these records are shown on the distribution map. However, its presence in Bolu, Düzce, Sinop and Zonguldak provinces (Sevgili 2001; Ünal 2003, 2004, 2010) located in northwest Anatolia is doubtful and requires confirmation by examining song characteristics since these records may concern the related P. miramae or P. rose-oviridis sp.n..

Remarks: Several species, similar to P. similis, were described from the Caucasus and the northeastern part of Turkey. Some of them were synonymized with P. similis (such as P. richteri and P. caucasicus) or later re-estab-



lished (compare Ramme 1933, 1951; Bei-Bienko 1954; Zhantiev & Korsunovskaya 2005). P. tereckensis is such a species considered as a synonym of P. similis by Ramme (1933) and Bei-Bienko (1954) and later re-established by Zhantiev & Korsunovskaya (2005) by the presence of a more curved stridulatory file. From the material examined we think that curvature of the stridulatory file is not uniform at species level and the peg number in P. tereckensisfits within the variation range of that character of P. similis. Again, song parameters of P. tereckensis presented in Zhantiev & Korsunovskaya (2005) retain within the variation limits of P. similis. The traditionally used morpho-logical characters also do not suggest it to be a distinct species. Therefore we consider P. tereckensis as a synonym of P. similis in agreement with Ramme (1933) and Bei-Bienko (1954).

There are further some species, which were given as related to P. similis such as Poecilimon geoktschajcusStshelkanovtzev, 1910 and Poecilimon oligacanthus Miram, 1938 (Bei-Bienko 1954). Presently there are no data about the song and morphology of stridulatory file of P. oligacanthus. During this study no specimens of this spe-cies were available, but photos of type material from OSF2 (Eades et al. 2011) were examined. Since the tradition-ally used morphological characteristics are highly variable and the type material cannot be distinguished from P. similis, we also considered Poecilimon oligacanthus under the first species. Unfortunately we have no data of the song and stridulatory structures for P. geoktschajcus, but, from the description and the account by Bei-Bienko (1954) the above statements are possibly valid also for this species. However, an exact taxonomical decision should wait further data since Bei-Bienko (1954) stated that it may be intermediate between P. similis and P. scythicus.Below this species is mentioned according to Bei-Bienko (1954).

Poecilimon geoktschajcus Stshelkanovtzev, 1910(Fig. 186)

Poecilimon geoktschajcus Stshelkanovtzev, 1910: Trav. Lab. Cab. Zool. Univ. Varsovie 1909: 53.Poecilimon zawadskji Stshelkanovtzev, 1910: 56 syn. by. Bei-Bienko (1954).Poecilimon bidens geoktshajcus Stshelkanovtzev. 1914. Rev. Rus. Ent. 14:267. Poecilimon bosphoricus geoktshajcus Stshelkanovtzev. 1914. Rev. Rus. Ent. 14:269. Poecilimon geoktschajcus Stshelkanovtzev, 1910; Ramme, 1933:561.Eupoecilimon geoktschajcus (Stshelkanovtzev, 1910); Ramme. 1951: 341, 417.Poecilimon geoktschajcus Stshelkanovtzev, 1910; Bei-Bienko, 1954: 351.

Type specimen information: Unspecified primary type male, female; Asia-Temperate, Caucasus, Transcaucasus, Azerbaijan: Geokchai Steppe (MZPW Warsaw) (Bei-Bienko 1954).

Description and distribution: See Bei-Bienko (1954), Harz (1969) and Avakyan (1981).

Poecilimon djakonovi Miram, 1938(Fig. 186)

P. djakonovi Miram, 1938: 352. Zool. Zhurnal 17: 351 (Russian)/365.P. djakonovi Miram, 1938:331 (Bei-Bienko 1954).

Type information: Holotype—male; Northwest Caucasus, Sofiya River (tributary of Bolshoi Zelenchuk) and foot-hills of Oshten Mt. (Bei-Bienko 1954).

Bei-Bienko (1954) considers P. djakonovi most closely related to P. oligacanthus (herewith considered syn-onym of P. similis) and P. similis. We support this opinion though further consideration about its taxonomic status requires studying its song.

Remark: In OSF (2011/03/29) together with photos of a paratype of P. djakonovi (NHM) some photos of a specimen identified by Demirsoy (ZMB) are presented. However, from the large spiracle and the relatively small tegmina with black markings it is evident that the latter is not a member of the P. bosphoricus group. This specimen may represent a taxon related to P. varicornis.

Description: See Bei-Bienko (1954).Distribution (Fig. 186): NW Caucasus, Abkhazia and NE Turkey (Bei-Bienko 1954; Demirsoy 1975).



Poecilimon bischoffi Ramme, 1933(Figs 18, 37, 48, 65, 77, 93, 104, 121, 135, 153, 170, 175–184, 186)

Poecilimon bischoffi Ramme, 1933: 572.Eupoecilimon bischoffi (Ramme); Ramme, 1951: 417.Poecilimon bischoffi (Ramme); Bei-Bienko, 1954: 338.

Type information: Holotype—male; Turkey, Trabzon, Polathane (ZMB) (Holotype (stridulatory file) and photos of type material presented in DORSA were examined).

Material examined: Holotype—male; TURKEY: Rize province—4 males, İkizdere, Ovit Mt., (40o41’ N,

40o 24’E), 1627 m, 29.7.2010 (leg. S. Kaya) (in alcohol in AUZM), 8 males, 7 females, (40o 43’ N, 040o 36’ E), 1009 m, 13.07.2011, (leg. B. Çıplak, S. Kaya, D. Chobanov, E. Mahir Korkmaz) (in alcohol in AUZM); UKRAINE: Crimea—1 male, Central Crimean Mountains, Angarskiy Pereval Pass, (44o 45’ N, 34o 20’ E) 765 m, 25.7.2010 (leg. D. Chobanov) (in alcohol in CDC).

Description: For morphology see Table 1, 2 and Figures 18, 37, 48, 65, 77, 93, 104, 121, 135, 153, 170; for song see Table 3 and Figures 175–184, 186. Additional description can be found mainly in Ramme (1933, 1951) and Bei-Bienko (1954).

Distribution (Fig. 186): This species is known from its type locality Turkey, Trabzon (Ramme 1933) and Rize (present paper). The specimen collected from Crimea, at the opposite of Anatolia Black Sea coast, was tentatively placed under this taxon and further molecular studies may add to its systematic position.

Remarks: So far this species was known only from the type material and it is identifiable from P. similis by the typical apex of male cerci. New specimens from Rize Province (Turkey) were identified as P. bischoffi by the cer-cal apex with the inner row of denticles being longer than the outer one. Some specimens from other localities show intermediate features between P. similis and P. bischoffi. Thus, it is difficult to doubtlessly distinguish both species. On the other hand, although the syllable pattern is similar in both species they differ from each other in several song parameters (Figs 169, 170, 175–184), thus P. bischoffi is retained as a distinct species. Furthermore, a specimen collected at the opposite side of the Black Sea (Crimea) is similar to P. bischoffi both in morphology (especially male cerci) and song (Table 3 and Figures 170). Though there are some differences in the shape of pro-notum and song (syllable duration—see Fig. 175), it was considered under P. bischoffi until new data is gathered.

Poecilimon scythicus Stshelkanovtzev, 1911(Figs 19, 49, 78, 105, 136, 154, 171, 175–184, 186)

Poecilimon scythicus Stshelkanovtzev, 1911: 18.

Type information: Unknown number of syntypes —males, females; Ukraine, SE, Proval'skii stud farm, Donetk Hills (ZIN St. Petersburg) (not examined).

Material examined: UKRAINE: Lugansk District—2 males, Umg. Lugansk, Lugansk Reserve, (47° 21’ N, 37° 03’ E) 12.6.1996 (leg. A. Benediktow); Donetska Oblast District—2 males, 7 females, Goreliy Forest, 26.6.2006 (leg. ?) (in alcohol in CDC).

Description: For morphology see Table 1, 2 and Figures 19, 49, 78, 105, 136, 154; for song see Table 3 and Figures 171, 175–184. Additional description can be found in Ramme (1933), Bei-Bienko (1954) and Harz (1969).

Distribution (Fig. 186): This is the most widespread species in the northern Basin of the Black Sea present in Ukraine, Central and Southeast European Russia to Volga River and Western and Central Caucasus (Bei-Bienko 1954; Zhantiev & Korsunovskaya 2005).

Remarks: Bei-Bienko (1954) presented a comprehensive description and a detailed account about P. scythicusand mentioned that it is similar to P. tauricus. Both species share the presence of syllable Type-3, but well differ in temporal parameters, which confirm the species status of both taxa.



Poecilimon tauricus Retowski, 1888(Figs 20, 21, 50, 79, 80, 106, 107, 137, 138, 156, 172, 173, 175–184, 186)

Barbitistes sanguinolenta Fischer von Waldheim, 1846. Syn. by Bei-Bienko (1954: (nomen nudum)Poecilimon tauricus Retowski, 1888: 408. Poecilimon bosphoricus Retowski, 1888: Jacobson & Bianchi 1905; Kirby, 1906: 379 (both partim).Poecilimon kusnezovi Miram, 1929: 463, syn.n.Poecilimon beybienkoi Tarbinsky, 1932: 184, syn.n.Poecilimon scythicus Retowski, 1888; Bei-Bienko. 1941: 147.Poecilimon ajpetri Stshelkanovtzev, 1911. Syn. by Bei-Bienko (1954).

Type information: Unknown number of syntypes—males, females; Krym, [Crimea: S, Dvukhyakornyi (near Kok-tebel), Krasnokamenka (Kiziltash) north of Otuz, Novyi Svet (west of Sudak) and Okhontich'e (Buragan)] (Bei-bienko 1954).

Material examined: UKRAINE: Crimea—for P. tauricus —11 males, 8 females, Crimea, Western Crimean

Mountains, Aj-Petri Yayla, (44o 28’ N, 34o 02’ E), 1250 m, 30.7.2010; for P. kusnezovi/P. beybienkoi—13 males, 6 females, Western Crimean Mountains, Babugan Yayla, (44o 36’ /44o 37’ N, 34o 14’ / 34o 17’ E), 1440/1360 m, 27.7.2010 (leg. D. Chobanov) (in alcohol in CDC).

Synonymy and description: This taxon has been described under several names (see the synonymic list above), the recent ones used being P. tauricus, P. kusnezovi and P. beybienkoi. During an extensive field trip over the mountains of South Crimea held in July 2010 material matching all these forms has been collected. From ear-lier literature data (e.g. Bei-Bienko 1954; Harz 1969; Zhantiev & Korsunovskaya 2005) it is very difficult to out-line the range of these taxa; they were reported from same localities even in one paper, e.g. the couples P. tauricus-kusnezovi from Babugan Yayla and P. kusnezovi-beybienkoi from Yalta (see last citations). Considering their very similar morphology misidentification may frequently be involved in these records. Subsequently, we collected typ-ical P. kusnezovi and P. beybienkoi, as well as intermediate forms at the same place and habitat (Babugan Yayla); Bei-Bienko (1954) also investigated and reported both P. kusnezovi and P. tauricus from this place. The latter author (l.c.) has possibly concerned this as a proof that these are good species. However, we think this is not the case. We observed smooth transition in the shape of the tenth tergum ranging from typical kusnezovi to typical bey-bienkoi (see Fig. 4). Male cerci were also quite variable in all populations, ranging from typical for kusnezovi/bey-bienkoi (Fig. 80) or tauricus (Fig. 79) to intermediate (compare Figs 79 and 80). The songs though having some differences between animals from different populations are generally very similar and yet quite variable (also unpublished data from laboratory reared animals). Finally, new records on possible hybrids between “typical” tau-ricus and kusnezovi, as well as unpublished data on the molecular phylogeny of the Crimean taxa (Chobanov et al., data in preparation) support the synonymic state of P. tauricus, P. kusnezovi and P. beybienkoi.

For morphological data of the species see the above given references, Table 1, 2 and Figures 20, 21, 50, 79, 80, 106, 107, 137, 138, 156; for song see Table 3 and Figures 172, 173, 175–184.

Diagnosis: The species is most similar to P. scythicus and P. pliginskii. From both it differs in the shape of male cerci (cf. Figs. 79, 80 for tauricus with 78 for scythicus and 81 for pliginskii). From scythicus it further differs in the well-expressed medial constriction of male pronotum with strongly widened pro-and metazona (cf. Figs. 20, 21 with 22). From pliginskii ts differs also in the longer subgenital plate, which however in some cases may be obscure together with the cercus shape due to their variability. Unpublished data (Chobanov et al., data in prepara-tion) support the strong genetic similarity of P. tauricus and P. pliginskii. Females of the three mentioned taxa can-not be distinguished.

Distribution (Fig. 186): Endemic to the mountains and south coast of Western Crimea—from Alushta and Babugan Yayla westwards.



http://orthoptera.speciesfile.or


Poecilimon pliginskii Miram, 1929(Figs 22, 51, 81, 108, 139, 140, 157, 158, 174, 175–184, 186)

Poecilimon tauricus Miram, 1929: 452 (partim) (Ramme 1933; Bei-Bienko 1954; Eades et al. 2011).Poecilimon pliginskii Miram, 1929: Bei-Bienko, 1954: 349.Poecilimon boldyrevi Miram, 1938: 363, 366, syn.n.

Type information: Holotype —male; Krym, [Crimea: South, Taushan-Bazar] (ZIN St. Petersburg).Material examined: UKRAINE: Crimea—for pliginskii, 4 males and 8 females, Eastern Crimean Moun-

tains, Echki Dag Mountain, (44o 54’ N, 35o 07’ E), 600 m, 22.7.2010 (leg. D. Chobanov) (in alcohol in CDC); 8

males and 10 females, Crimea, Central Crimean Mountains, Chetyr Dag Mountain, (44o 44’ N, 34o 19’ E), 1250 m, 25.7.2010 (leg. D. Chobanov) (in alcohol in CDC); for boldyrevi, 4 males, 8 females, Crimea, Northern slope of the Crimean Mountains, Krasnolesje village S of Simgeropol, (44o 46’ N, 34o 11’ E), 700 m, 30.7.2010 (leg. D. Cho-banov) (in alcohol in CDC).

Synonymy and description: Bei-Bienko (1954) considered P. pliginskii and P. boldyrevi distinct taxa regard-ing mostly the shape of male cerci. We found considerable variation of this character (see Fig. 81) even in one place. For example, typical “boldyrevi-type” of cerci (Fig. 81A—specimen from Krasnolesje, Central Crimea) is almost identical with a specimen of P. pliginskii from Echki-Dag (near Karadag, Eastern Crimea) (Fig. 81E). The songs of both taxa (compare Zhantiev & Korsunovskaya 2005: Figs 10–11 with 18–20) have identical syllable structure; the different syllable grouping varies in all Crimean taxa (see Figs 174–184; unpublished data). Addi-tionally, unpublished data on genetic phylogeny (Chobanov et al., data in preparation) support the identity of both taxa. The great variation and genetic similarity (Ramme 1933; own unpublished data) suggest close relationships also of P. tauricus and P. pliginskii and their possible infraspecific relationships but we remain this question open until new data are gathered.

For morphology see Table 1, 2 and Figures 22, 51, 81, 108, 139, 140, 157, 158; for song see Table 3 and Fig-ures 174, 175–184.

Diagnosis: P. pliginskii is most similar to P. tauricus; for differences see the latter.Distribution (Fig. 186): Endemic to the South Crimea, occurring along the South coast and the mountains

mostly in the eastern part of the peninsula; further records are known westwards to Taushan-Bazar.

Discussion

MorphologyThe Poecilimon bosphoricus group was first defined by Ramme (1933) within his generic revision as “Group VIII”. Most of the subsequent publications on these species (e.g. Ramme 1951; Bei-Bienko 1954; Harz 1969) refer mainly to the characters used in this revision. Other publications on the species group (e.g. Ramme 1951; Karabağ 1958; Karabağ et al. 1971, 1981; Ünal 2003, 2004, 2005, 2010) are faunistic lists or descriptions of new species which also use the character set defined by Ramme (1933). It should be mentioned that the only molecular study including these species is that by Ullrich et al. (2010) in which phylogenetic analyses (for nuclear genes ITS1 + ITS2 and mitochondrial 16S rDNA+tRNA Val+12S rDNA separately) were conducted to test the monophyly of Poecilimon. Monophyly of the species presented in this study is well supported in the analysis of ITS1 + ITS2 sequences. However, Ullrich et al. (2010) do not present any conclusion on the internal taxonomy of the group, thus its taxonomic clarification requires reconsidering these characters and testing them with other character sources.

In general, the taxonomy developed for the group is based on three morphological structures in males; prono-tum, cerci and subgenital plate. The three characters defined in pronotum do not diagnose any species from others, except the non-elevated metazona in P. turcicus (see Character 1–3 in Table 1). The incision of male subgenital plate at caudal margin is another character which was commonly used in species diagnoses. However, on the basis of material examined and the descriptions given in previous publications this character is also variable both in pres-ence/absence and in the depth of the incision (see Table 1). Thus, the characters defined from pronotum and male subgenital plate are insufficient to be used alone for the internal taxonomy of the group.



The structure of male cerci is still the most productive diagnostic character, although caution is necessary when delimiting the species. Of the eleven characters defined from male cerci, the robustness, the apical width and the curvature are either subjective (yet applicable in some cases) or variable within species, thus, may be defined dif-ferently in different publications. The number of denticles on each row is variable and the variations in different species may overlap (see character 15, 16). However, we think that these characters can also be used in combina-tion with other characters or they may diagnose few species from each others. On the other hand, the orientation of the cercal denticles as well as the number of denticular rows and their location and the presence/absence of large denticles in some species are of the potential of more objective limitation of species (characters 9–14 in Table 1). Thus, these are among the main characters used in the key to species.

During this study some metric characters were also studied. The length of pronotum, tegmina and hind femur, structures commonly measured in Orthoptera, are overlapping between species. This is also the case for the ratio fastigium-scapus (see Table 2). Thus, such morphological characters are also uninformative in revealing the inter-nal taxonomy of P. bosphoricus group. However, the number and the structure of the stridulatory pegs provide some reliable information (see Heller 1988) about relationships of the species and can be used to diagnose at least some species from others. P. sureyanus is clearly different from all others by the least number of stridulatory pegs (Table 2). The peg number in P. turcicus and P. turciae is intermediate, higher than in the previous two species and lower than in all others though there is not a clear cut. The peg number in the species other than given above is vari-able and overlapping between species (Table 1), but it still diagnoses some species from others.

Why morphology (qualitative or quantitative) is not useful in description of species? First, description of some characters (such as elevated metazona, or convex or concave hind margin of pronotum, incised male subgenital plate ect.) is subjective in the absence of numerical values. Measuring these structures (such as the angle of prono-tum dorsally) allowed a more objective definition of the character states and showed that they overlap between related species. Second, several species were known from a few specimens (it is still valid for some) and deciding about the limits of variation of the qualitative characters in such cases is misleading. Thus, variations become prominent for some species if additional material is examined. Third, if the specimens examined are from a single locality then it is difficult to define variation limits at species level properly. So, inclusion of the material from new localities may be another reason to increase the known variation in some species. Fourth, a great emphasize has been given to morphology in species descriptions/diagnoses and so the descriptions of new species provided a more confusing taxonomy of the group. Finally, the possible hybridisation between species in the past and present (e.g. between P. cervus x P. demirsoyi, P. sureyanus x P. turcicus) might have lead to appearance of populations with mixed or intermediate character states. This is a hypothesis to be tested by genetic data. Another hypothesis emerging from the present study and worth to be tested by an appropriate method is coming from the habitat pref-erences and the similar morphology of the species in the group. The latter typically occur (and possibly evolved) in habitats dominated by Euxinic vegetation types characterised by similar microclimate conditions and this may lead to a speciation that retained less variable morphology due to similar selection pressures. Yet, a few species expanded their ranges to Mediterranean like habitats.

SongPrevious song data were available for a few species in the group (Heller 1988; Zanthiev & Korsunovskaya 2005). The new song data for nearly all species allowed a thorough evaluation of this character and testing the classifica-tion based on traditional morphological characters. At group level the song is more or less uniform in its general pattern (consisting of irregularly repeated syllables). Yet, the syllable structure shows definable patterns, which seem related to the wing movements. Changing the duration and speed of the closing movements, three types of syllables may be produced as presented in the Results section. Our results yielded four major outcomes. First, there is a correlation between the number of stridulatory pegs and the syllable type. The Type-1 is characteristic for the species with the least peg numbers (P. kocaki and P. sureyanus). Type-2 is found in P. turcicus and P. turciae where the peg number is higher than in the previous two species and lower than in those producing syllable Type-3. For this syllable type, however, a modified neuromuscular program is necessary with two speeds of movements. The third syllable type is characteristic for the species with higher peg numbers. Second, the syllable duration is vari-able and provides data to distinguish some species from others. Statistical analyses suggested four major clusters of the species with syllable Type-3 (Fig. 175). However, the number of impulses per syllable is much more variable in the group than the syllable period, thus much more informative for the taxonomy of the group. Congruent with this



statement, a Tukey analysis applied to the impulse number suggested seven distinct clusters (Fig. 176). Third, dif-ferentiation between species exhibiting syllable Type-3 in temporal song parameters is expressed mainly in dura-tion of the early part of the syllable since statistical analyses applied to this character suggested several distinct clusters (Fig. 177). Analyses of the other parameters belonging to the early or late parts of the syllables separated only a few species from others. Therefore, the late part is possibly more conservative than the early part (see Figs 180–182). Fourth and the last, song characteristics provided a more objective delimitation of the species and hence more objective taxonomy. Thus, definition of the subgroups is mainly based on these characters and decisions were made according to the song when song contra morphology.

Taxonomy and species distributionsBy combination of bioacoustics and morphological characters (mainly orientation of cercal denticles) we listed 21 species within P. bosphoricus group. One of these one, Poecilimon roseoviridis sp. n., is new to science. P. proxi-mus stat. n. has been raised to a species level from subspecies. We also proposed eigth synonyms; P. anatolicussyn. n. and P. diversus syn. n. to P. sureyanus, P. naskrecki syn. n. to P. demirsoyi; P. tereckensis stat.rev. and P. oligacanthus syn.n. to P. similis, P. beybienkoi syn.n. and P. kusnezovi syn.n. to P. tauricus, P. boldyrevi syn.n. to P. pliginskii.

We defined three subgroups within P. bosphoricus group according to characteristics of the song and stridula-tion file (see Results section). The species group and subgroups established in this study on the basis of song type and stridulation structures do not agree with that suggested by Ünal (2010); however, he has not given any charac-ters or objective synapomorphies for the groups proposed and these groups are sometimes in conflict with the data presented by Ullrich et al. (2010). It should be noted that the relationships defined here agree with the most defined by Ramme (1933) or Bei-Bienko (1954). There are two or three species within each of the first and second sub-groups; (1) P. sureyanus + P. kocaki + P. athos and (2) P. turcicus +P. turciae. The third group includes the remain-ing 15 species. It is obvious that the latter can be split further into interclades, however, this remains preliminary until a thorough phylogenetic analysis will be accomplished. The possible intraclade relationships among species in the third subgroup were mentioned in the species accounts. These are as follows: (1) P. cervus + P. demirsoyi (by sharing the second row of the denticles located dorso-apically), (2) P. similis + P. bischoffi + P. miramae + P. roseo-viridis (similar both in cercal morphology and song parameters), (3) P. bosphoricus + P. istanbul + P. bidens (pres-ence of one or more huge teeth apico-internally), (4) P. scythicus + P. tauricus + P. pliginskii (the external margin of male cercus is nearly straight and the song parameters are similar). Of the remaining species P. proximus and P. hei-nrichi are aberrant from that mentioned above. Available data about P. djakonovi do not allow a statement. Some of the results indicate a conflict between bioacoustic and qualitative morphological characters. P. heinrichi and P. turciae were considered sister species by Ramme (1951) and others for the widened apex of male cerci, however, they distinctly differ in their syllable patterns and the number of stridulatory pegs. This was also confirmed by molecular phylogenetic analyses (Ullrich et al. 2010). An opposite case is true for P. cervus and P. demirsoyi. The cercal morphology is distinctive, but the song parameters are very similar. The presence of individuals with inter-mediate morphology between these two species leads to questioning the importance of the cercal morphology in the reproductive isolation. Another similar case is observed for P. kocaki and P. bosphoricus. The apical rows of the denticles not joining to each other is shared by these two species, but, the first produces a song in syllable Type-1 and the second Type-3. Thus, we think defining species in P. bosphoricus species group still requires an iterative testing of species cohesion by further data (Yeates et al. 2011).

There are either allopatric or sympatric species in the group (Figs 185, 186). However, although some species were recorded from the same geographical unit, rarely two or more species were found syntopically. The latter is common case for the co-occurrence of P. roseoviridis and P. heinrichi in Strandzha Mountains, as well as for P. proximus which has frequently been found together with other species of its group (see Unal 2010: together with P. turciae, P. miramae, P. demirsoyi (as P. naskrecki), P. cervus and P. bosphoricus).

On the other hand, some records of co-occurrence reported in previous studies, especially those of closely related species, require re-examination of the specimens. It is worth mentioning that established or possible mis-identifications in previous publications are not rare. Thus, a phylogeographic study will allow more prominent statements for the speciation pattern in this taxonomically difficult group.

Another interesting aspect of the distribution pattern is related to the total range of the group. The group is typically represented through the whole Black Sea Basin in a ring-area. There seem to be three core areas of the



range of the P. bosphoricus group. P. similis, P. geoktschajcus + P. djaconovi (and other forms now given under P. similis) are known from the Caucasus and the eastern half of the Anatolian Black Sea Basin. According to our opinion there are four species in the northern Basin of the Black Sea (mainly Crimea); P. tauricus, P. pliginskii(endemic for Southern Crimea), P. scythicus (widely distributed north of the Caucasus) and P. bischoffi. Since Anatolian and Crimean populations of P. bischoffi are very similar in phenotype (morphology and song), a recent dispersal is possible, however, a more proper explanation can be done after a molecular study. The remaining species are confined to Northwest Anatolia and the Southeast Balkans (the region of Thrace), both divided by a sea barrier including the Dardanelles + Marmara Sea + the Bosphorus. Of these P. heinrichi, P. roseoviridis, P. athosand P. istanbul were recorded only from the western side of this barrier. P. sureyanus, P. bidens, P. miramae, P. bosphoricus and P. turcicus are represented on both sides with populations not prominently diverged. Also, Anatolian and Lesbos’s populations of P. turcicus do not exhibit any prominent phenotypical differences. The remaining species, P. cervus, P. kocaki, P. demirsoyi and P. proximus are endemic to Northwest Anatolia. From this distribution pattern the following conclusions can be made as working hypotheses for future studies. First, there are three radiation centres of the group; (1) Caucasus + Northeast Anatolia, (2) Crimea + North Caucasus, and (3) Northwest Anatolia + Thrace. Second, the barrier constituting by Dardanelles + Marmara Sea + Bosphorus is an important entity influencing the speciation of the group. However, there are species found on both sides, so the barrier might have not been permanent in the radiation course of the group. A similar role can be attributed to the Caucasian Mountains and the Black Sea itself since nearly all species of the Northern and Southern Black Sea coast are endemic to the respective site (only the distribution of P. bischoffi conflicts with this). Third, judging from the preference of a certain type of habitat, the distribution pattern of the group and the weak morphoacoustic differentiation between related species, it might be concluded that the climatic fluctuations during the Pliocene-Pleistocene are main factor leading to the speciations within this group. All these considerations offer hypotheses to be tested by further data.

Acknowledgements

Collecting data for this study has taken more than a decade period and during this period we received help/support from different persons/organisations. Some specimens used in this study are from Dr. Selahattin Salman (Kırşehir) and Dr. Abbas Mol (Samsun). A great help has been provided by the Russian scientists Dr Andrej Gorochov and Dr Olga Korsunovskaya by sending photos of P. bidens from ZIN, St. Petersburg. The young orthopterists Zehra Boz-tepe has made some technical contributions. The statistical test would have been difficult without the help by Dr M. Ziya Fırat (Antalya). Dr. Nurşah Çokbankir and Teyfik Turgut helped us in translation of previous German publica-tions. Special thanks are due to Prof. Dr. Sergey Ivanov (Taurida National University “V.I. Vernadsky”, Simferopol, Crimea, Ukraine) and Dr. Yury Budashkin (Karadagh Nature Reserve, Ukrainian Natinal Academy of Sciences, Feodosya, Crimea, Ukraine) for their kind hospitality and invaluable help during the expedition to the Crimea. We are grateful to all.

Field studies during which we collected material were financed by both Akdeniz University Research Fund and Turkish Scientific and Technical Research Council (TUBITAK) for other projects. D. Chobanov was supported by grant N N303 611738 by the Ministry of Science and Higher Education of Poland; Synthesys grants DE-TAF-3950 and AT-TAF-546; grant DO 02-259/08 of the Ministry of Education, Youth and Science of Bulgaria; BG051PO001-3.3.04/41 (European Social Fund through the Ministry of Education, Youth and Science of Bul-garia).

Author contributions: B.Ç. conceived the ideas; S.K., D.C., B.Ç. and K-G.H. collected the specimens and recorded the songs; S.K. produced the data; S.K. and B.Ç. analysed the data; and B.Ç. led the writing.

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FIGURE 1. General appearance of members of the P. bosphoricus group. A—P. sureyanus (male,) B—P. kocaki (male), C—P. miramae (male), D—P. miramae (female), E—P. cervus (male), F—P. cervus (female)



FIGURES 2–4. Terminology followed in this study. 2—male pronotum (A—P. heinrichi, B—P. bosphoricus), 3—Male cerci (A—P. heinrichi, B—P. naskrecki, C-P. bosphoricus), 4—Male anal tergum (A—P. pliginskii, B, C—P. tauricus ), 5—Male subgenital plate (P. sureyanus).



FIGURES 6A–C. Terminology for male calling song. A—P. sureyanus, B—P. turcicus, C—P. cervus



FIGURES 6D–F. Oscillograms of stridulatory movements and song [synchronous registration of left tegmen movement (upward deflection represents opening, down-ward closing) and sound]. D—P. sureyanus Ulu Dagh, E—P. turcicus Lesbos, F—P. similis Giresun: Kümbet.



FIGURES 7–14. Hand drawings of male pronotum, from above (scale = 2 mm): 7—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A—Bilecik, B-Bursa, C, D-Balıkesir), 8—P. turcicus (Turkey: Edirne), 9—P. turciae (Turkey: Bursa), 10— P. heinrichi (Bulgaria: A, B-Strandzha), 11—P. demirsoyi (Turkey: Bolu), 12—P. bosphoricus (Turkey: A, B, C-Istanbul, D, E-Çankırı), 13—P. bidens (Turkey: Istanbul), 14—P. istanbul (Turkey: Istanbul).



FIGURES 15–22. Hand drawings of male pronotum, from above (scale = 2 mm): 15—P. miramae (Bulgaria: Rhodopian Mts.), 16—P. roseoviridis sp. n. (Bulgaria: Strandzha), 17—P. similis (Turkey: Rize), 18—P. bischoffi (Turkey: A-Trab-zon, Ukraine: B-Crimea), 19—P. scythicus (Ukraine: Crimea), 20—P. tauricus (Ukraine: A, B-Crimea), 21—P. tauricus (= P. kus-nezovi syn.n.) (Ukraine: A, B-Crimea), 22—P. pliginskii (Ukraine: A, B-Crimea) (=P. boldyrevi syn.n. Krasnolesje) (Ukraine: C, D-Crimea).



FIGURES 23–25. Photos of male pronotum, from above (scale = 2 mm): 23—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A, B, C-Bilecik, D, E-Bursa, F, G-Balıkesir, H-Greece: Lekomi), 24—P. kocaki (Turkey: Sakarya), 25—P. turcicus (Tukey: A-Edirne, B-Balıkesir, Greece: C-Mytillini).



FIGURES 26–31C. Hand drawings of male pronotum, from above (scale = 2 mm): 26—P. turciae (Turkey: Bursa), 27—P. heinrichi (Bulgaria: Strandzha), 28—P. proximus (Turkey: A, B-Bolu), 29—P. cervus (Turkey: Bolu), 30—P. demirsoyi (Tur-key: A, B, C, D-Bolu), 31—P. bosphoricus (Turkey: A, B—Istanbul, C-Bolu).



FIGURES 31D–36E. Hand drawings of male pronotum, from above (scale = 2 mm): 31—P. bosphoricus (Turkey: A, B—Istanbul, C-Bolu, D-Kastamonu), 32—P. bidens (Turkey: Istanbul), 33—P. istanbul (Turkey: Istanbul), 34—P. miramae (Tur-key: A-holotypus, B, C-Sakarya), 35—P. roseoviridis (Bulgaria: Strandzha), 36—P. similis (Turkey: A-Rize, B-Artvin, C-Ordu, Georgia: D, E- Bakuriani, Russia).



FIGURES 36D–37. Hand drawings of male pronotum, from above (scale = 2 mm): 36—P. similis (Georgia: F-Bakuriani, Rus-sia: G-Krasnaya Polyana), 37—P. bischoffi (Turkey: A-Trabzon, B, C-Rize, Ukraine: D-Crimea).



FIGURES 38–47. Hand drawings of male pronotum, from lateral view (scale = 2 mm): 38—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A-Bilecik, B-Bursa, C-Balıkesir), 39—P. turcicus (Turkey: Edirne), 40—P. turciae (Turkey: Bursa; from OSF online), 41—P. heinrichi (Bulgaria: A, B, C-Strandzha), 42—P. demirsoyi (Turkey: Bolu), 43—P. bosphoricus (Turkey: A, B-Istanbul, C-Çankırı), 44—P. bidens (Turkey: Istanbul), 45—P. miramae (Bulgaria: Rhodopian Mts.), 46—P. roseoviridis (Bulgaria: Strandzha), 47—P. similis (Turkey: Rize).



FIGURES 48–51. Hand drawings of male pronotum, from lateral view (scale = 2 mm): 48—P. bischoffi (Turkey: A-Trabzon, Ukraine: B-Crimea), 49—P. scythicus (Ukraine: Crimea), 50—P. tauricus (Ukraine: A, B-Crimea), (= P. kusnezovi syn.n.) (Ukraine: C, D-Crimea), 51—P. pliginskii (Ukraine: A—Crimea) (= P. boldyrevi syn.n. —Krasnolesje), (B, C, D, E—Crimea).



FIGURES 52–56A. Photos of male pronotum, from lateral view (scale = 2 mm): 52—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A, B, C-Bilecik, D-Bursa, E—Balıkesir; Greece: F-Lekomi), 53—P. kocaki (Turkey: Sakarya), 54—P. turcicus (Turkey: A-Edirne, B-Balıkesir, C-Mytillini), 55—P. heinrichi(Bulgaria: Strandzha), 56—P. proximus (Tur-key: A-Bolu).



FIGURES 56B–62A. Photos of male pronotum, from lateral view (scale = 2 mm): 56—P. proximus (Tur-key: A, B-Bolu), 57—P. cervus (Turkey: Bolu), 58—P. demirsoyi (Turkey: A, B-Bolu), 59—P. bosphoricus (Turkey: A-Istanbul, B-Bolu, C-Çankırı, D-Kastamonu), 60—P. bidens (Turkey: A-Istanbul, B-Kocaeli), 61—P. istanbul (Turkey: Istanbul), 62—P. miramae(Turkey: A-Holotype).



FIGURES 62B–65. Photos of male pronotum, from lateral view (scale = 2 mm): 62—P. miramae (Turkey: A-Holotype; B—Sakarya), 63—P. roseoviridis (Bulgaria: Strandzha), 64—P. similis (Turkey: A-Rize, B, C-Artvin, D-Ordu, Russia: E-Kras-naya Polyana, Georgia: F-Bakuriani), 65—P. bischoffi (Turkey: A-Trabzon, Ukraine: B-Crimea)



FIGURES 66-77A. Hand drawings of male cerci (scale = 1 mm): 66—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A—Bilecik; B, C, D-Bursa, E-Balıkesir), 67—P. turcicus (Turkey: Edirne), 68—P. turciae (Turkey: A, B-Bursa), 69—P. heinrichi (Bulgaria: A, B, C-Strandzha), 70—P. demirsoyi (Turkey: A, B, C-Bolu), 71—P. bosphoricus, (Tur-key: A, B, C, D, E-Istanbul, F, G-Çankırı, H, J—Bolu), 72—P. bidens (Turkey: A, B-Istanbul), 73—P. istanbul (Turkey: Istan-bul), 74—P. miramae (Bulgaria: A, B, C-Rhodope Mts), 75—P. roseoviridis sp.n. (Bulgaria: A, B, C-Strandzha) 76—P. similis (Turkey: A, B-Rize, C-Ordu, D-Artvin), 77—P. bischoffi (Turkey: A-Trabzon).



FIGURES 77B-81. Hand drawings of male cerci (scale = 1 mm): 77—P. bischoffi (Turkey: A, B-Trabzon, Ukraine: C-Cri-mea) 78—P. scythicus (Ukraine: A-Donetsk, B, C, D-Ramme 1951), 79—P. tauricus (Ukraine: A, B, C-Crimea, D, E, F, G-Ramme 1951), 80—P. tauricus (=P. kusnezovi syn.n. and P. beybienkoi) (A, B, C, D, E, F, G-Ukraine: Crimea, H-Ramme 1933), 81—P. pliginskii (Ukraine: A—Crimea) (=P. boldyrevi syn.n. —Krasnolesje) (Ukraine: B, C, D, E-Crimea).



FIGURES 82–88A. Photos of male cerci (scale = 1 mm): 82—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A, B-Bilecik; C, D, E, F: Bursa, G-Balıkesir; Greece: H-Lekomi), 83—P. kocaki (Turkey: Sakarya), 84—P. turcicus (Tur-key: A-Edirne, B-Balıkesir, Greece: C-Mytillini), 85—P. proximus (Turkey: A, B-Bolu), 86—P. cervus (Turkey: A, B-Bolu), 87—P. demirsoyi (Turkey: A, B, C-Bolu), 88—P. bosphoricus (Turkey: A, B-Istanbul).



FIGURES 88B–93. Photos of male cerci (scale = 1 mm): 88—P. bosphoricus (Turkey: A, B-Istanbul, C, D-Bolu, E, F-Çankırı), 89—P. bidens (Turkey: A-Istanbul, B-Kocaeli), 90—P. miramae (Turkey: A-holotype; B-Sinop, C, D-Sakarya), 91—P. roseoviridis sp.n. (Bulgaria:Strandzha), 92—P. similis (Turkey: A-Rize, B-Artvin, C-Ordu, Russia: D-Krasnaya Poly-ana), 93—P. bischoffi (Tur-key: A, B, C-Rize).



FIGURES 94-108. Hand drawings of male subgenital plate (scale = 2 mm): 94—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A, B, C, D-Bilecik; E, F-Bursa), 95—P. turcicus (Turkey: A-Edirne) 96— P. heinrichi (Bulgaria—Strandzha Mts), 97—P. demirsoyi (Turkey: A, B, C, D-Bolu), 98—P. bosphoricus (Turkey: A, B-Istanbul, C-Bolu, D-Çankırı), 99—P. bidens (Turkey: Istanbul), 100—P. istanbul (Turkey: Istanbul), 101—P. miramae (Bulgaria: Rhodope Mts), 102—P. roseoviridis sp.n. (Bulgaria: Strandzha Mts), 103—P. similis (Turkey: A, B-Rize), 104—P. bischoffi (Ukraine: Crimea), 105—P. scythicus (Ukraine: Crimea), 106—P. tauricus (Ukraine: A, B-Crimea), 107—P. tauricus (=P. kusnezovi syn.n.) (Ukraine: A, B-Crimea), 108—P. pliginskii (Ukraine: A (=P. boldyrevi syn.n.—Krasnolesje), B, C-Crimea).



FIGURES 109–114. Photos of male subgenital plate (scale = 2 mm): 109—P. sureyanus (= P. anatolicus syn.n. and P. diver-sus syn. n.) (Turkey: A, B-Bilecik, C, D, E-Bursa, F-Orhaneli, G-Balıkesir, Greece: H-Lekomi), 110—P. kocaki (Turkey: Sakarya), 111—P. turcicus (Turkey: A-Edirne, B-Balıkesir, Greece: C-Mytilini), 112—P. proximus (Turkey: A, B-Bolu), 113—P. cervus (Turkey: Bolu), 114—P. demirsoyi (A, B, C, D, E-Bolu).



FIGURES 115–121. Photos of male subgenital plate (scale = 2 mm): 115—P. bosphoricus (Turkey: A, B, C-Istanbul, D, E-Bolu, F, G-Çankırı), 116—P. bidens (Turkey: A-Istanbul, B-Kocaeli), 117—P. istanbul (Turkey: Istanbul), 118—P. miramae(Turkey: A-paratype B-Sinop, C-Sakarya), 119—P. roseoviridis sp.n. (Bulgaria: Strandzha Mts), 120—P. similis (Turkey: A-Rize, B-Art-vin, Russia: C-Krasnaya Polyana, Ukraine: D-Crimea), 121—P. bischoffi (Turkey: A, B-Rize).



FIGURES 122–129A. Photos of female ovipositor (scale = 2 mm): 122—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A, B-Bilecik; C, D, E-Bursa, F, G-Balıkesir), 123—P. kocaki (Turkey: Sakarya), 124—P. turcicus (Turkey: Balıkesir), 125—P. turciae (Turkey: Bursa), 126— P. heinrichi (Bulgaria: Strandzha) 127—P. cervus (Turkey: A, B-Bolu), 128—P. demirsoyi (Turkey: Bolu), 129—P. bosphoricus (Turkey: A-Istanbul).



FIGURES 129B–140. Photos of female ovipositor (scale = 2 mm): 29—P. bosphoricus (Turkey: A-Istanbul, B-Bolu, C-Çankırı, D-Kastamonu), 130—P. bidens (Turkey: Kocaeli), 131—P. istanbul (Turkey: Istanbul), 132—P. miramae (Turkey: Sakarya), 133—P. roseoviridis sp.n. (Bulgaria—Strandzha Mts), 134—P. similis (Turkey: Rize), 135—P. bischoffi (Turkey: Rize), 136—P. scythicus (Ukraine: Crimea), 137—P. tauricus (=P. ajpetri syn.n. —Ukraine: Crimea), 138—P. tauricus (=P. kusnezovi syn.n.) (Ukraine: Crimea), 139—P. pliginskii (Ukraine: Crimea), 140—P. pliginskii (=P. boldyrevi syn.n.) (Ukraine: Crimea).



FIGURES 141–150. Female subgenital plate and basal valves of the ovipositor (scale = 1 mm): 141—P. sureyanus (=P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A-Bilecik; B-Bursa, C-Balıkesir), 142—P. kocaki (Turkey: Sakarya), 143—P. turcicus (Turkey: Edirne), 144— P. heinrichi (Bulgaria: Strandzha) 145—P. cervus (Turkey: Bolu), 146— P. demirsoyi (Turkey: A, B-Bolu), 147— P. bosphoricus, (Turkey: A, B-Bolu, C-Kastamonu), 148—P. bidens (Turkey: Kocaeli), 149—P. istanbul (Turkey: Istanbul), 150—P. miramae (Turkey: A, B-Sakarya).



FIGURES 151–158. Female subgenital plate and basal valves of the ovipositor (scale = 1 mm): 151—P. roseoviridis sp.n. (Bulgaria—Strandzha Mts), 152—P. similis (Turkey: A-Ardahan B-Artvin, C-Rize), 153—P. bischoffi (Turkey: Rize), 154—P. scythicus(Ukraine: Crimea), 155—P. tauricus (=P. ajpetri syn.n.) (Ukraine: Crimea), 156—P. tauricus (=P. kusnezovisyn.n.) (Ukraine: Crimea), 157— P. pliginskii (Ukraine: Crimea), 158—P. pliginskii (=P. boldyrevi syn.n) (Ukraine: Crimea).



FIGURES 159–160A. Male calling song. 159—P. sureyanus (= P. anatolicus syn.n. and P. diversus syn. n.) (Turkey: A Bil-ecik, B-Bursa, Greece: C-Kavalla), 160—P. turcicus (Greece: A-Mytillini).



FIGURES 160B–163. Male calling song. 160—P. turcicus (Greece: A-Mytillini, Turkey: B-Kırklareli), 161—P. turciae (Tur-key: Çanakk-ale), 162— P. heinrichi (Bulgaria: Strandzha), 163—P. proximus (Turkey: Bolu).



FIGURES 164–166. Male calling song. 164—P. cervus (Turkey:Bolu), 165—P. demirsoyi (Turkey: A, B-Bolu), 166—P. bos-phoricus (Turkey: Bolu).



FIGURES 167–169. Male calling song. 167—P. miramae (Turkey: A-Sakarya; Bulgaria: B-Rhodopian Mts), 168-P. roseo-viridis sp.n. (Bulgaria: Strandzha), 169—P. similis (Turkey: Rize),



FIGURES 170–172. Male calling song. 170—P. bischoffi, (Turkey: A-Rize, Ukraine: B-Crimea), 171—P. scythicus (Ukraine: Lugansk), 172—P. tauricus (Ukraine: Crimea).



FIGURES 173–174. Male calling song. 173—P. tauricus (=P. kusnezovi syn.n.) (Ukraine: Crimea), 174—P. pliginskii(Ukraine: Crimea).

FIGURES 175–176. Box plots for song parameters of P. bosphoricus group species (Statistical analyses were made on the basis of populations prior to taxonomical decisions made in this study. Thus, the synonymised species are given in old names in the plots). 175—Syllable duration (SD), 176—Impulse number per syllable (INS).



FIGURES 177–184. Box plots for song parameters of P. bosphoricus group species (Statistical analyses were made on the basis of populations prior to taxonomical decisions made in this study. Thus, the synonymised species are given in old names in the plots). 177—Duration of early part of the syllable (DEP), 178—Impulse number in the early part of the syllable (INE), 179—Period of impulses in the early part of syllable (PIE), 180—Duration of late part of the syllable (DSP), 181—Impulse number in the late part of the syllable (INL), 182—Period of impulses in the late part of syllable (PIL), 183—Ratio of dura-tions of the early/lat parts (RDEL), 184—ratio of the periods of the impulses in the early/late parts (RPEL).



FIGURES 185–186. 185—Distribution of Poecilimon bosphoricus group species in Turkey and Greece, 186—Distribution of Poecilimon bosphoricus group species in Black see basin.



TABLE 1. Qualitative/meristic morphological characters used in internal taxonomy of P. bosphoricus species group (Characters were given on the basis of populations and the synonymised species are given in old names in the table)

1- Caudal margin of pronotum in male: (0) concave, (1) straight, (2) convex.2- Elevation of male pronotum in metazoan (as the angle of dorsal margin): (0) >1700, (1) <1600.3- Index ratio width metazoan/prozona in male: (0) <1.5, (1) >1.5. 4- Cubital vein of tegmen in male: (0) covered by pronotum, (1) visible (not covered).5- First abdominal tergite in female: (0) monochromatic, (1) banded.6- Male subgenital plate at hind margin: (0) straight, (1) incised (roundly, triangularly or quadrangularly) (2) convex.7- Male tenth tergum at hind margin; (0) straight, (1) with two conical processes 8- Cerci at apex; (0) not branched, (1) branched.9- Structure of male cerci: (0) slender, (1) moderately robust, (2) robust.10- Curvature of male cerci at apex: (0) 90-119o, (1) 120-150o.11- Width of male cerci at apex: (0) roughly in the same width of proximal part or narrower, (1) widened12- Tapering of male cerci: (0) narrowing from external margin, (1) narrowing from both of external and internal margins.13- Male cerci at apex: (0) denticulate only along externo-apical margin, (1) denticulate along both externo-apical and interno-

apical margins, (2) denticulate both along externo-apical and dorso-apical margins; (3) denticulate along externo-apical and apical margins

14- The apical denticle(s) of male cerci: (0) same or roughly as large as the others, (1) obviously larger than all others. 15- Number of denticles on externo-apical margin (number per species).16- Number of denticles on dorso-apical or interno-apical margin (number per species). 17- Coloration of male cerci at apex: (0) black only along the basis of denticles, (1) black roughly at apical 1/3.18- Length of subgenital plate according to cerci: (0) shorter, (1) equal-nearly equally, (2) longer.19- Number of stridulatory pegs (range and median per species)



TABLE 4. Results of Regression and one-way ANOVA analyses

** P<0.0001, DEPS: Duration of early part of syllable, INEPS: Impulse number in early part of syllable, PIEPS: Period of impulses in the early part of syllable, DLPS: Duration of late part of syllable, INLPS: Impulse number in the late part of syllable, PILPS: Period of impulses in the late part of syllable, DRELPS: Duration ratio of early and late part of syllable, IPRELSP: Impulse period ratio of early and late part of syllable, Trns: Transversion, dfM: degrees of freedom of model, dfP: degrees of freedom of population,

dfE: degrees of freedom of error, F: F test result,CP: Mallow’s CP, Adj-r2: Adjusted r square, AIC: Akaike information criterion, BIC:Bayesian or also Schwardz information criterion.

Regression ANOVA

Character Trns dfM dfE F CP Adj-r2 AIC BIC dfP dfE F

Frq - - - - - - - 20 1550 116.37**

SD LnSD 21 1542 800.58** 18.0717 0.9148 -5424.4360 -5421.8676 20 1543 534.50**

INS - - - - - - - - 20 1546 1744.56**

DFPS InvDFPS 14 1026 445.05** 12.8120 0.8567 -11018.938 -11016.435 15 1025 1061.09**

INEPS - - - - - - - - 15 1028 518.83**

PIEPS InvPIEPS 12 1031 199.62** 11.4725 0.6956 -4490.3552 -4487.9054 15 1028 135.79**

DLPS InvDLPS 14 1026 222.68** 14.2596 0.7193 -10864.682 -10862.221 15 1023 256.62**

INLPS - - - - - - - - 15 1013 184.80**

PILPS LnPILPS 15 1008 72.52** 15.4466 0.5119 -3725.7673 -3723.2421 15 1008 146.38**

DRELPS - - - - - - 15 1010 216.77**

IPRELSP - - - - - - 15 1004 113.40**


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