H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

Turk J Zool 2011; 35(5): 631-652 © TÜBİTAK Research Article doi:10.3906/zoo-0912-5

Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Hasan SEVGİLİ1,*, Ali İsmet DEMİRSOY2, Yusuf DURMUŞ2 1Ordu University, Faculty of Arts and Sciences, Department of Biology, Cumhuriyet Campus, Ordu - TURKEY 2Hacettepe University, Faculty of Science, Department of Biology, 06532 Beytepe, Ankara - TURKEY

Received: 02.12.2009

Abstract: Th e and Mantodea fauna of Kazdağı National Park (Balıkesir) were investigated. Field studies were conducted between 2004 and 2005. During this study, 34 species of Orthoptera (16 , 1 Oecanthidae, 2 Pamphagidae, 1 Tetrigidae, and 14 Acrididae) and 3 species of Mantodea were investigated. Isophya cania, Poecilimon luschani egrigozi, Anadolua schwarzi, Rhacocleis acutangula (Tettigoniidae), Paranocaracris rubripes burri (Pamphagidae), and one recently described species, Chorthippus (Glyptobothrus) kazdaghensis (Acrididae), are endemic taxa. Metrioptera (Broughtonia) arnoldi,, known from the Balkan fauna until now, is a new record for Turkish fauna. Th e calling songs of Metrioptera arnoldi, Rhacocleis acutangula, Anadolua schwarzi, Isophya cania, and Poecilimon luschani are described here for the fi rst time. Th e recordings of the some species are also given along with habitat observations and altitudinal distribution. From a zoogeographical perspective, Kazdağı holds both Balkan fauna elements as well as those typical of Mediterranean-Anatolian fauna.

Key words: Fauna, bioacoustics, Kazdağı, Orthoptera, Mantodea, biogeography, Turkey

Kazdağı (İda) Milli Parkı Orthoptera ve Mantodea faunası ile bazı çalıçekirgelerinin çağrı sesleri

Özet: Bu çalışmada Kazdağı (İda) Milli Parkı (Balıkesir) Orthoptera ve Mantodea faunası incelenmiştir. Arazi çalışmaları 2004-2005 yıllarını kapsamaktadır. Çalışma sonucunda Orthoptera takımına ait 5 familyadan 34 tür, Mantodea takımına ait 2 familyadan 3 tür saptanmıştır. Orthoptera familyalarından Tettigoniidae familyasına ait 16 tür, Oecanthidae familyasına ait 1 tür, Pamphagidae familyasına ait 2 tür, Tetrigidae familyasına ait 1 tür ve Acrididae familyasına ait 14 tür belirlenmiştir. Çalışma alanından Tettigoniidae familyasına ait Isophya cania, Poecilimon luschani egrigozi, Anadolua schwarzi ve Rhacocleis acutangula, Pamphagidae familyasından Paranocaracris rupripes burri, Acrididae familyasından yakın bir zamanda tanımlanan Chorthippus (Glyptobthrus) kazdaghensis taksonları endemiktir. Kazdağı’ndan saptanan Tettigoniidae familyasına ait daha önceleri Bulgaristan’dan bilinen Metrioptera (Broughtonia) arnoldi Türkiye faunası için yeni kayıttır. Metrioptera arnoldi, Rhacocleis acutangula, Anadolua schwarzi, Isophya cania ve Poecilimon luschani egrigozi tür/alttürlerinin erkek çağrı sesleri ilk kez bu çalışmada tanımlanmıştır. Çalışmada türlere ait varsa eski kayıtların yanı sıra, habitat gözlemleri, yayılış yükseltileri ve taksonomik problemleri de verilmiştir. Çalışma alanı zoocoğrafi k açıdan değerlendirildiğinde, Kazdağı’nın tipik Akdeniz-Anadolu faunasının yanı sıra Balkan faunası elemanlarını da barındırdığı ortaya konmuştur.

Anahtar sözcükler: Fauna, biyoakustik, Kazdağı, Orthoptera, Mantodea, biyocoğrafya, Türkiye

* E-mail: [email protected]

631 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Introduction endemic and semiendemic species richness. More Damage to nature caused by human activities is than 60% of the bush cricket species (Tettigoniidae) a worldwide problem that has steadily increased in and several genera (e.g. Karabağ, 1958; Ünal, 2002) Turkey in recent years. Unsustainable agriculture, recorded from Turkey are endemic to Anatolia unregulated extractive industries, creeping (Çıplak et al., 1993; Çıplak and Demirsoy, 1995; urbanization, and the fragmentation of natural Çıplak et al., 2002a). environments are resulting in the loss of many However, when compared with other parts of species per year. Habitat loss and fragmentation pose Turkey, the Orthoptera fauna of the western part a serious problem for the proliferation of species of the Marmara region is rather poorly studied, since it limits the ability of many species to migrate especially on the Biga Peninsula and nearby islands to favorable conditions. In addition, the impact of of the Aegean Sea. Until now, no faunistic study global warming, resulting in numerous shift s in the concerning orthopterans has been conducted in this distribution and abundance of species, is now a well area, excluding a few taxonomic recordings (Heller documented fact (Th omas et al., 2004). Th erefore, et al., 2004; Sevgili, 2004a; Ünal, 2004; Çıplak et al., many countries have had to take preventive measures 2005). Th erefore, this study will make an important for protecting the diversity of natural habitats and contribution to our understanding of the Orthoptera against the loss of biodiversity in their regions. and Mantodea fauna of the region and to the Kazdağı National Park, the focus of this study, is one zoogeographical patterns of western Anatolia. of such important biodiversity hotspots in Turkey. In addition to taxonomic recordings, we also Study objectives recorded and described the acoustic signals of 11 Currently our knowledge of the biodiversity bush-cricket species within this area. Male songs of Kazdağı National Park is poor, excluding some are one of the most conspicuous elements of cricket plants (Özhatay and Özhatay, 2005; Satıl et al., and katydid behavior, conveying several kinds of 2006) and groups such as mammals (Yiğit information to the females for species identifi cation et al., 2006), amphibians, and reptiles (Hür et al., and mate choice (e.g. Heller, 1988, 1990, 2006; Ragge 2008), malacofauna (Gümüş, 2006), and butterfl ies and Reynolds, 1998; Gerhardt and Huber, 2002). and moths (Seven, 2006, 2007). Th e main objective Th e amplitude modulation of male calling songs of this study was to determine the Orthoptera and are thus rather diverse and strictly species specifi c Mantodea fauna of the national park. Th e Orthoptera in many genera such as (e.g. Ragge, 1990; and Mantodea fauna of Turkey has been studied Ragge and Reynolds, 1998; Çıplak et al., 2002b), extensively (e.g. Mantodea by Demirsoy, 1977a and Isophya (e.g. Heller, 1988; Sevgili, 2004a; Sevgili et Orthoptera by Ramme, 1933a, 1933b, 1951; Uvarov, al., 2006), Poecilimon (e.g. Heller, 1988; Heller and 1934; Bei-Bienko, 1954; Karabağ, 1958, 1964; Sevgili, 2005; Heller et al., 2006), Leptophyes (e.g. Karabağ et al., 1971, 1974, 1980; Demirsoy, 1973, 1975, 1977b; Salman, 1978; Naskrecki and Ünal, Sevgili, 2004b), and Chorthippus (Perdeck, 1958; Mol 1995; Çıplak et al., 1996a, 1996b, 2002a; Sevgili and et al., 2003). Hence, in addition to faunistic data and Çıplak, 2000; Ünal, 2000, 2002, 2004; Sevgili, 2001) remarks, oscillograms of several important bush- and orthopterans have served as a model in the cricket species are provided. biogeographical evaluation of the region (Demirsoy, 2002; Çıplak, 2003a, 2003b, 2008; Kenyeres et al., Materials and methods 2009). Many factors contribute to the diversity of faunal and fl oral elements in Anatolia, where the Study area number of endemic species is high and most species Kazdağı (1796 m), formerly known as Ida are restricted to small areas (e.g. for fl ora, Avcı, 2005; Mountain, is the highest peak of the Biga Peninsula for fauna, Demirsoy, 2002). For example, among separating the Aegean and Marmara regions from the groups studied to date, Orthoptera has a high each other (Figure 1). Th e national park (39°40ʹN,

632 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

0 10 20 km N WE S Figure 1. Map of Biga Peninsula and Kazdağı.

26°45ʹE) is mainly made up of deep valleys, and the Song recordings canyons situated within the highlands of Kazdağı Live specimens were collected for song recordings continue in a north-south direction towards the and kept in the laboratory in string cages. Recordings vicinity of Edremit. Th is region supports diverse were made on multiple days at irregular intervals at 29- and distinct fl ora and fauna, consisting mainly of fi r forests at elevations higher than 1000 m and pine 30 °C. Specimens were kept isolated from each other forests at lower elevations. Th ere are about 800 natural in diff erent cages. For sound recording, a Sony WM- plant taxa in Kazdağı National Park and 68 of them GX688 Walkman and a stereo microphone were used. are endemic to Turkey (Özhatay and Özhatay, 2005). Th is system does not record frequencies above 20 kHz. About 30 of the endemic taxa grow only in this park Th e microphone was placed approximately 15 cm from (Satıl et al., 2006). Th erefore, the area was classifi ed the cage. Aft er digitizing the songs on a PC computer, as a European “Important Plant Area” (http://www. oscillograms (aft er high-pass fi ltering, Bessel, the cut- plantlife-ipa.org). off frequency was set to 1000 Hz) and song analyses Specimens and localities were conducted using the programs Turbolab and CoolEdit on a PC computer. For more details on the Specimens collected during fi eld studies were methods used, see Ragge and Reynolds (1998). prepared as museum material by the standard methods and deposited in the Hacettepe University Song terminology followed that of Ragge and Zoological Museum (HUZOM), Ankara. Some Reynolds (1998). Calling song: the song was produced specimens were preserved in alcohol. Th is study by an isolated male. Syllable: the sound produced covers the results of fi eld studies conducted between by one complete opening and closing stroke of the 2004 and 2005, except for 2 species. Nearly all tegmina. Echeme: a fi rst order assemblage of syllables. specimens used in the study were collected by the Echeme-sequence: a fi rst-order assemblage of authors. follows the OSF database (Eades echemes. Impulse: a simple, undivided transient train and Otte, 2009). of sound waves. Click: an isolated distinct impulse.

633 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Taxonomy Remarks: Typically, the species was found from ORTHOPTERA July to August in suitable areas at low altitudes. However, according to our fi eld studies, population densities were low. Tettigonioidea Isophya cania Karabağ, 1975 Tettigoniidae Isophya cania: Karabağ, 1975. J. Nat. Hist. 9(3): 341. Acrometopa syriaca Brunner von Wattenwyl, Previous records: 2♂♂, Balıkesir: Kazdağı 1878 (Sarıkız yolu), 25.06.2002, 770 m; 2♂♂, 1♀, Sarıkız Acrometopa syriaca: Brunner von Wattenwyl, tepesi, 25.06.2002, 1650 m; 1♀, Çanakkale: Bayramiç 1878. Verh. der Zoologisch-botanischen Gesellsch (Kazdağı yolu), 27.06.2002 (M. Ünal) (Ünal, 2004). Wien 28: 85, 87. Material examined: 6♂♂, 1♀ (in alcohol), Material examined: 1♂ (in alcohol), Balıkesir: Balıkesir: Edremit (Kazdağı, Sarıkız), 03.07.2004; Edremit, Kazdağı, Zeytinli Stream, 17.08.2004 (H. 3♂♂ (in alcohol), same locality, 23.07.2004 (H. Sevgili). Sevgili & Y. Durmuş); 5♂♂, 4♀♀, same locality, 19- Calling song: We analyzed the songs of one male. 20.07.2005 (Y. Durmuş & H. Sevgili); 4♂♂, 7♀♀ Th e calling song was usually heard in the evening and (in alcohol), Tozlu altı, 23.07.2004; 2♂♂, 2♀♀, at night. Th e song pattern was described by Heller same locality, 19.07.2005 (Y. Durmuş & H. Sevgili); (1988) from Greece. Our song data from Kazdağı 3♂♂ (in alcohol), Tozlu, 24.07.2004; 5♂♂, 5♀♀ (in were similar to his data. Th e male calling song was a alcohol); 3 ♂♂ (in alcohol), Kazdağı, 03.07.2004, dense echeme followed immediately by strong clicks 1217 m (H. Sevgili & Y. Durmuş). (Figures 2A and 2B). Th e echeme began as crescendo Calling song: Th e songs from 6 males were syllables and then continued at a nearly constant recorded in the evening and at night in the laboratory syllable amplitude. Th e echeme usually lasted at 28-30 °C. Th e general pattern of the male calling between 555 and 892 ms (mean: 724 ms at 30 °C, n song was very similar to that of I. thracica (Heller, = 25). Th e echemes were repeated irregularly and the 1988; HS, unpublished data), but the duration periods of successive echemes were quite variable, between the main syllable and the “click” was much measuring between 4.5 and 20 s. Aft er an interval longer, about 598-880 ms (n = 24) and syllables of approximately 45-53 ms (mean: 48 ms, n = 25), were repeated at very variable intervals (4.2-29 s) a series of 11-17 clicks followed (duration of series: 15-23 ms, n = 25). Th e mean duration of the main (Figures 3A and 3B). Th e main syllables were of the echeme was 651 ms at 30 °C and it was composed of decrescendo type (Figures 3C and 3D), lasting about 52-81 syllables (n = 22). 30 ms (min-max: 24-37 ms, n = 36). Th e syllables were composed of about 5-19 impulses (mean: 12.85 ± 0.8, n = 35). Th e impulse period varied between 1 B and 8 ms. Th e “click” was not present in some calling songs.

A Remarks: I. cania (Figure 4) was very similar to 1 s I. thracica morphologically and bioacoustically, but the male tegmina and cercus were diff erent (Sevgili, 2004a). Both species were considered in the I. thracica subgroup within the I. rectipennis species group, B endemic to the western and Th racian parts of Turkey 100 ms (Sevgili, 2004a). Th e distribution of I. thracica is Figure 2. Acrometopa syriaca: A) 2 echemes; B) a faster known from the western part of Çanakkale Province oscillogram of the syllables indicated in A. and Th race.

634 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

C nymphs of the species can be found in various habitat types from April to May at lowlands and from May to July at highlands, and mature adults can be found A from May to July onwards. Populations reach peak 5s densities in July.

D Poecilimon anatolicus Ramme, 1933 Poecilimon anatolicus: Ramme, 1933. Mitt. Zool. Mus. Berlin 19: 555. B 2 s Material examined: 3♂♂, 1♀ (in alcohol), Balıkesir: Edremit, Kazdağı, Tozlu altı, 23.07.2004; 1♂ (in alcohol), same locality, 26.07.2004; 1♂ (in alcohol), Açöldüren, 23.07.2001, 1364 m (H. Sevgili). C 50 ms Calling song: Th e songs of 3 males were analyzed. Th e calling song consisted of an irregular sequence of syllables. Th e mean syllable duration was about 277 ms (30 °C, n = 8). Th e syllables were composed of 15- D 18 impulses (Figure 5A-C). Th e impulse period varied 100 ms between 3 and 30 ms (mean: 17.16, n = 125). Heller Figure 3. Isophya cania, oscillograms of the male calling song: A) 3 calling songs, B) 1 complete song from A, C) 3 (1988) recorded the calling song of P. anatolicus from calling songs with “click,” D) faster song and “click” of Bursa (Uludağ) and Çanakkale (Gelibolu) and found the syllables shown in Figure C. a similar song pattern. Remarks: Th is species resembles P. turc i c u s , which is known from the western Marmara and Th racian regions and was recently discovered from Lesbos Island, Greece (Willemse and Willemse, 2008), by the long denticles on the tip of male cerci (Sevgili, 2001). However, the apex morphology of the cerci was diff erent in both species and the male cerci were without denticles on the posterior margin of the apical part (Karabağ, 1950; Bei-Bienko, 1954).

B

Figure 4. Isophya cania (male). A 0.5 s

C Distribution and habitat: Until now, this species was known only from the Balıkesir and Çanakkale B provinces; its type locality is Gönen (Balıkesir) 0.5 s (Karabağ, 1975). According to fi eld observations, the species is patchily distributed and inhabits open parts of forests associated with bush vegetation (e.g. Rubus sp.), green herbaceous vegetation found on edges of small streams, and Astragalus sp. and Juniperus sp. C 100 ms from montane steppes of the national park, excluding Figure 5. Poecilimon anatolicus: A) 2 male calling songs, B) and the subalpine zone between 600 and 1700 m. Th e C) faster oscillograms of the syllables shown in A.

635 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Distribution and habitat: P. anatolicus is mountains of western Anatolia. Th e taxonomic status widespread in the western Marmara region and of these populations needs to be clarifi ed based on the Th racian parts of Turkey and Greece. We fi rst more reliable and detailed studies covering the whole collected this species from the bushes (Rubus sp.) in distributional area of the taxa. a glade at Kazdağı (near Açöldüren). Adults of this Distribution and habitat: Th e nominate species can be found mainly in June from Kazdağı subspecies is distributed in Gölbaşı, between Kale and also from Uludağ (about 1300 m; Heller, 1988; and Kaş in Antalya Province (Ünal, 2005). P. lu s chani HS, unpublished data). egrigozi is known from only Kütahya (type locality) Poecilimon luschani egrigozi Ünal, 2005 and Balıkesir provinces. Th is interesting subspecies Poecilimon luschani egrigozi: Ünal, 2005. Trans. lives only above 1000 m on the mountain of Kazdağı, Am. Entomol. Soc. 131: 440. and it is associated with small herbaceous plants in lowlands and montane steppes in and around Previous records: 1♂, Balıkesir: Edremit, Kazdağı, Sarıkız. Adults were found mainly on Astragalus Sarıkız yolu, 25.06.2002, 770 m; 1♂, Kazdağı, Sarıkız sp. in the open areas of Sarıkız in July and August. tepesi, 1650 m (M. Ünal) (Ünal, 2004, as P. lu s chani ). Nymphs were seen from mid-May. Males were Material examined: 1♂, 1♀ (in alcohol), bioacoustically active both in daytime and at night. Balıkesir: Kazdağı, Sarıkız, 23.07.2001 (H. Sevgili In the low localities, P. luschani egrigozi was collected & Y. Durmuş); 10♂♂ (in alcohol), same locality, from open-side plants in the forest. While the other 03.07.2004; 16♂♂, 7♀♀, same locality, 19-20.07.2005, 2 of the 3 Poecilimon species (P. pergamicus and P. 1730 m (Y. Durmuş & H. Sevgili); 2♂♂ (in alcohol), anatolicus) that are known from Kazdağı prefer low Kazdağı, Tozlu, 03.07.2004 (H. Sevgili). localities and the vegetation in open-side forests and Calling song: Th e songs of 5 males were analyzed. near water courses, P. l. egrigozi generally prefers high Th e calling song consisted of single crescendo altitudes. syllables (duration: 87-140 ms, mean: 97 ms, n = Poecilimon pergamicus Brunner von Wattenwyl, 32, impulse number: 25-53 at 28 °C), which were 1891 produced at quite variable intervals (Figures 6A and Poecilimon pergamicus: Brunner von Wattenwyl, 6B). Th e syllable period ranged from 444 to 964 ms 1891. Verh. der Zoologisch-botanischen Gesellsch (mean: 656, n = 32). Wien 41: 129. Remarks: Th is subspecies, recently described Previous records: 1♂, Balıkesir: Kazdağı, from Kütahya Province by Ünal (2005), is endemic Sarıkız yolu, 27.06.2002, 770 m; 1♂, Sarıkız tepesi, to northwestern Anatolia. Our materials showed 25.06.2002, 1650 m; 2♂♂, 1♀, Çanakkale: Bayramiç, morphological similarities with P. luschani egrigozi. Kazdağı, Ayazma, 27.06.2002, 1400 m (M. Ünal) P. lu s chani lives in distinct, isolated, local populations (Ünal, 2004). in various habitats at high altitudes throughout the Material examined: 2♂♂, 1♀ (in alcohol), Balıkesir: Kazdağı, Açöldüren, 23.07.2001, 1364 B m; 7♂♂ (in alcohol), Kazdağı, 03.07.2004, 1067 m; 3♂♂ (in alcohol), Kazdağı, Tozlu altı, 23.07.2004 (H. A Sevgili & Y. Durmuş); 1♂, same locality, 19.07.2005, 3 s 1330 m (Y. Durmuş & H. Sevgili); 1♂ (in alcohol), camping area, 03.07.2004 (H. Sevgili); 1♂ (in alcohol), Kazdağı, Hasan Boğuldu, 06.06.2005 (Y. Durmuş). B Calling song: Th e songs of 4 males were analyzed. 250 ms Th e calling songs were faint and consisted of a series Figure 6. Poecilimon luschani egrigozi: A) a sequence of 10 of syllables of variable lengths (Figures 7A and 7B). complete male syllables, B) 2 complete songs from A. Each syllable contained 14-17 impulses and lasted

636 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

B low herbaceous plants in glades and shrubs at the sides of streams between 500 and 1300 m. It has not A been recorded at altitudes over 1300-1400 within 2 s the national park. In some places, this species was found to occur syntopically with I. canina. It was B also collected at sea level from around Bergama 100 ms (İzmir) and Alibey Island (Ayvalık) in April (HS, Figure 7. Poecilimon pergamicus: A) oscillogram of the male unpublished data). calling song recorded at 30 °C, B) 2 syllables from A. lilifolia (Fabricius, 1793) Locusta lilifolia: Fabricius, 1793. Supplementum about 17 ms. Th e syllable period lasted between 0.4 Entomologiae Systematicae 2: 36. and 0.95 s (mean: 0.56, n = 32). Syllables were repeated Material examined: 1♂, 3♀♀ (in alcohol), fairly regularly at the rate of about 1-3/s. Th e calling Balıkesir: Edremit, Kazdağı, camping area, 800 song of P. pergamicus was recorded form Bergama m, 25-29.08.2004; 1♂ (in alcohol), same locality, (İzmir Province) by Heller (1988), who found shorter 23.07.2004; 1♂, same locality, 19.07.2005 (Y. Durmuş syllable duration and impulse numbers. & H. Sevgili); 1♀ (in alcohol), Kazdağı, camping Remarks: Th is species (Figure 8) was very area-Ayı Deresi Yolu, 25.08.2004 (H. Sevgili & Y. similar to the northwestern Anatolian endemic Durmuş). species P. kutahiensis, as they have a similar external morphology, especially the shape of the male cercus. Anadolua schwarzi Ramme, 1939 However, the shape of the male subgenital plate was clearly diff erent from P. kutahiensis. Male calling Anadolua schwarzi: Ramme, 1939. Mitt. Zool. songs, which consist of a fast sequence of single Mus. Berlin 24: 68. syllables, are also diff erent (Heller, 1990). Th e nymphs Material examined: 3♂♂, 3♀♀, Balıkesir: of P. pergamicus could be found from the second Edremit, Kazdağı, 25.08.2004 (H. Sevgili & Y. week of April in Kazdağı. Adults were observed Durmuş) (HUZOM); 1♂ (holotype), Muğla: from mid-May to the end of August. Adults reached southwestern Anatolia, Sandras Dağı, Köyceğiz, 2600 high densities in June and July and mating occurred m, 07.1938 (Dr. O. Schwarz); 1♀ (allotype), Konya: during this period. Males were bioacoustically active Anatolien, Akchehir Göl, 01.10.1934 (coll. Fuss) (in mostly in the evenings and during the night. Berlin Museum of Natural History); 1♂, Kazdağları, Distribution and habitat: Known from the İzmir, Koşara yöresi, 30.07.1977, 1250 m (S. Salman). Balıkesir, and Çanakkale provinces of Turkey and Calling song: Th e calling songs of 3 males were Lesbos Island of Greece. Th e species was found on recorded at 29 °C. Th e calling song consisted of a series of trisyllabic (sometimes disyllabic) echemes repeated regularly at the rate of about 6-7/s (Figures 9A-9C). Th e song was most oft en broken up into short echeme sequences separated by brief intervals of about 0.16-0.5 s, but was also emitted without interruption for long periods of up to 25 s or more. Echeme sequences usually began with disyllabic echemes; the remaining echemes consisted mainly of 3, but sometimes 4, syllables and the fi rst syllable was usually short and oft en quiet. Each trisyllabic echeme lasted about 55-65 ms and successive echemes were separated by intervals of about 70-113 ms (Figure Figure 8. Poecilimon pergamicus (male). 9D). Th e syllables in each echeme usually lasted

637 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

B

2 s A C

B 0.5 s D Figure 10. Anadolua schwarzi (female). C 100 ms

Calling song: Th e calling song of 1 male was D recorded at 22 °C by Heller (1988) from Macedonia. 25 ms Our data were diff erent from Heller’s recording in Figure 9. Anadolua schwarzi: A) a slow oscillogram of the male terms of repetition rate and syllable duration due calling song, B) a faster oscillogram of the echemes to diff erent temperatures; the male calling song, indicated in A, C) a faster oscillogram of the 7 echemes indicated in B, D) a faster oscillogram of the 1 complete produced both during the day and at night, consisted echeme of A. schwarzi. of a long series of isolated syllables repeated at intervals of about 2-10 s (mean: 4.88 s at 30 °C) (Figure 11A). Th e syllable consisted of 2 hemisyllables lasting about 15.7 ms (range: 10 to 20, n = 44), but the fi rst together about 69-110 ms (mean: 90 ms, n = 35) and syllables were usually shorter (10-13 ms). Within was repeated at the rate of about 13-28/min (Figure each echeme, the syllable repetition rate was between 11B). 18/s and 21/s. Th ere were no microsyllables. Remarks: Th is widely distributed species was Remarks: Th ese individuals diff ered from A. seen at altitudes of 500-1000 m, within forest gaps, schwarzi in that the structure of the pronotum and from July to August within Kazdağı. female ovipositor was shorter, with a diff erent shape. Decticus albifrons (Fabricius, 1775) Th eir identifi cation needs to be checked. Locusta albifrons: Fabricius, 1775. Supplementum Th e Anadolua is endemic to western Entomologiae Systematicae 2: 86. Anatolia (Karabağ, 1952). In Kazdağı, it was seen Material examined: 1♂, 1♀, Balıkesir: Edremit, only around Tozlu and densely in the vicinity of Kazdağı, camping area, 800 m, 27.07.2004 (H. Sevgili Sarıkız and Radar. Th is genus prefers the Astragalus & Y. Durmuş). sp. vegetations at high altitudes. Nymphs were seen from the end of July. Th e adults of this species (Figure Remarks: Th is species preferred the open-side 10) could be seen from mid-August to the end of gaps in forests below 1000 m. Adults were present at September. Th is species was usually bioacoustically active at night. B Bucephaloptera bucephala (Brunner von Wattenwyl, 1882) Th amnotrizon bucephala: Brunner von Wattenwyl, A 2 s 1882. Prodromus des Europaischen Orthopteren: 338. Material examined: 1♂, 1♀, Balıkesir: Edremit, Kazdağı, camping area, 800 m, 25.08.2004 (H. Sevgili B 100 ms & Y. Durmuş); 2♂♂ (in alcohol), same locality, Figure 11. Bucephaloptera bucephala: A) oscillogram of the male 23.07.2004 (H. Sevgili & Y. Durmuş). calling song, B) 1 complete song from A.

638 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

the end of July and August and they called during the within an echeme at the rate of about 18/s (range: 15 day in hot, sunny weather. Recordings were collected to 20, n = 28). Th e opening syllables were shorter, from Balıkesir-Edremit (Karabağ et al. 1971). not more than 6 ms. Th e duration of a single echeme Metrioptera arnoldi Ramme, 1933 period was about 213 ms (range: 196 to 225, n = 74) and the interval between 2 echemes was about 106 Metrioptera arnoldi: Ramme, 1933. Mitt. Zool. ms (range: 80 to 133, n = 74). Th e duration of echeme Mus. Berlin 18(3): 422. was about 106 ms (range: 80 to 129, n = 74). Th e Material examined: 2♂♂, 5♀♀, Balıkesir: duration of the echeme depended on the number of Edremit, Kazdağı, Sarıkız civarı, 27.07.2004; 1♂ (in syllables in each echeme. alcohol), same locality, 25.07.2004; 1♂ (in alcohol), Th e calling song of M. arnoldi was very similar to same locality, 23.07.2004 (H. Sevgili & Y. Durmuş) that of M. brachyptera, M. saussuriana, M. caprai, and (HUZOM); 1♂ (Type), Bulgarien: Pirin, Eltepe, M. bicolor, in that it had a long series of polysyllabic 06-07.08.1931 (A. Müller); 1♂, 1♀, Rita (or Rila) echemes (Ragge and Reynolds, 1998; Buzzetti et (Brunner von Wattenwyl); 14♂♂, 9♀♀, Macedonian: al., 2006). When the songs of these species were Vodno b. Skopje, 5-800 m, 10.08.1939 (W. Ramme); compared, they diff ered in syllable repetition rate, 1♀, Serbien: Suva Planina oberh Duschnik, 1200 echeme repetition rate, syllable number per echeme, m, 24.07.1939 (W. Ramme) (in Berlin Museum of and whether they had a microsyllable or not. Th ese Natural History). bioacoustic parameters are listed in Table 1. Diagnosis and affi nities: M. arnoldi belongs to Remarks: Th is species is known as a Balkan same subgenus, Broughtonia, as M. domogledi, known endemic species. According to previous authors, its from the Balkans. Its general appearance resembles distribution mainly covered Bulgaria at a considerable that of M. domogledi and the Bicolorana subgenus, range of altitudes, from sea level up to about 2200 m usually greenish in color with black lateral stripes on (Ramme, 1933b; Warchalowska-Sliva et al., 1987; the posterior part of the head and pronotum (Figures Heller et al., 1998). We collected our specimens at 12A-12C). Pronotum dorsally fl at, slightly widened high altitudes in Kazdağı. in metazona. Tegmina short, reaching almost to the Parapholidoptera castaneoviridis (Brunner von middle of the abdomen in males (Figures 12B-12D). Wattenwyl, 1882) Male cerci are curved inwards, with a robust apex and a small inner tooth that is slightly upcurved Th amnotrizon castaneoviridis: Brunner von (Figure 12E). Th is species is clearly related to M. Wattenwyl, 1882. Prodromus des Europaischen domogledi and can be easily distinguished from it Orthopteren: 337. by more robust teeth of the cercus and a subgenital Material examined: 8♂♂, 3♀♀ (5♂♂, 2♀♀ plate with a triangular excision in males (Figure 12F). in alcohol), Balıkesir: Edremit, Kazdağı, Sarıkız, Females are characterized by a subgenital plate that 23.07.2004 (H. Sevgili & Y. Durmuş); 1♂, same has a deep excision, but not reaching to the middle, locality, 19.07.2005 (Y. Durmuş & H. Sevgili); 1♂ (in as illustrated by Harz (1969). Diagnostic morphology alcohol), same locality, 25.07.2004; 1♂ (in alcohol), of our female specimens fi ts the descriptions given by Açöldüren mevkii, 39°42ʹN, 26°53ʹE, 1364 m, Ramme (1933b) and Harz (1969). 23.07.2001 (H. Sevgili & Y. Durmuş). Calling song: Th e songs of 3 males were recorded Calling song: Th e songs of 2 males were recorded. at 29 °C. Th e calling song consisted of long sequences Th e genus Parapholidoptera was revised by Çıplak of echemes repeated at an average rate of 4.6/s (range: (2000) and the song pattern of some species was 4 to 5, n = 28) (Figures 13A-13C). Each echeme adapted on the cladogram given by Çıplak and Heller consisted of 3 or 4 syllables (Figures 13C and 13D). (2006). Th e song pattern was oft en relatively similar Opening hemisyllables were usually present, though within the species groups (Heller, 2006). Heller (1988) oft en quiet. Th e closing hemisyllables always lasted recorded the male calling song and stridulatory longer than the opening hemisyllables, usually about movements of P. castaneoviridis from Bursa and 16.8 ms (range: 14 to 19, n = 30), and were repeated Edirne. Our recording was similar to his data. Th e

639 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Figure 12. Metrioptera arnoldi (male): A) head and pronotum from above, B) tegmina from above, C) head and pronotum in lateral view, D) tegmina in lateral view, E) cerci and apex of titillators, F) subgenital plate from below.

640 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

Table 1. Some bioacoustic parameters of Metrioptera arnoldi and some related taxa.

Syllable rep. Echeme rep. Microsyllables added Species Syllable/echeme Literature rate/s rate/s to an echeme

M. arnoldi 3-4 18 (15-20) 4.6 (4-5) no microsyllables Our data

M. brachyptera 3-4 35-55 4-7 no microsyllables Ragge and Reynolds (1998)

M. saussuriana 5-10 65 2-6 2-6 Ragge and Reynolds (1998)

M. caprai spp. 8-15 32.5-60 1.7-4.2 3-6 Buzzetti et al. (2006)

Heller (1988); Ragge and M. bicolor 3 80-120 15-25 no microsyllables Reynolds (1998)

B B

A A 0.5 s 2.5 s C

B B 50 ms D Figure 14. Parapholidoptera castaneoviridis: A) 2 echemes of the song recorded at 29 °C, B) a faster oscillogram of the echeme indicated in A. C 0.5 s

D 50 ms Figure 13. Song structure of male Metrioptera arnoldi: A) an echeme sequence of the male calling song, B) and C) faster oscillograms of the echemes, D) 2 complete echemes from C. male calling song consisted of echemes separated by variable intervals (several seconds; Figure 14A). Within each syllable group (5-8 syllables), the amplitude increased at the beginning, remained constant, and then slightly decreased toward the end (Figure 14B). Th e echeme duration was between 268 and 420 ms (mean: 335 ms at 29 °C, n = 6). Th e song period was longer and measured about 2.78-4.20 s. Figure 15. Parapholidoptera castaneoviridis (male). Remarks: Th is species (Figure 15) generally lives at high altitudes and is spread within the northern within forest gaps and bush at higher altitudes. It was Aegean region of Turkey, the island of Samothrace bioacoustically active at night. Nymphs appeared in (Greece), and the southern Balkans (Çıplak, 2000). June and adults were present from mid-July to the In Kazdağı, it was spread from approximately 800 m end of August. It has been recorded from the vicinity to higher al titudes, and it lived in scrub vegetation of Balıkesir, Savaştepe-Soma Road (Çıplak, 2000).

641 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Platycleis (Platycleis) intermedia intermedia B C (Serville, 1839) A Decticus intermedia: Serville, 1839 [1838]. 2 s Histoire naturelle des insectes. Orthoptères: 488. Previous records: 2♀♀, Balıkesir: Edremit yolu, Dereören-Eğmir kavşağı, 150 m, 13.07.1996; 1♂, Edremit, Kazdağı, Zeytinli Yayla mevki, 750-800 B m, 20.07.1998; 2♀♀, Edremit, Kazdağı, Zeytinli, Tuzlugöl mevki, 1400 m, 20.07.1998 (B. Çıplak) (Çıplak et al., 2002b). Material examined: 1♂, 4♀♀ (3♀♀ in alcohol), C Balıkesir: Edremit, Kazdağı, camping area, 800 0.5 s m, 25.08.2004; 2♂♂,3 ♀♀ (in alcohol), Kazdağı, camping area, 03.07.2004; 1♂ (in alcohol), same Figure 16. Platycleis affi nis: A) 2 complete echemes, B) and C) 2 faster, diff erent oscillograms indicated in A. locality, 24.07.2004; 1♂ (in alcohol), Kazdağı, Sarıkız, 23.07.2004; 1♂ (in alcohol), Kazdağı, camping area- Ayı Deresi yolu, 26.05.2005; 1♂, 1♀ (in alcohol), same locality, 25.08.2004 (H. Sevgili & Y. Durmuş). Rhacocleis acutangula Karabağ, 1957 Platycleis (Platycleis) affi nis affi nis Fieber, 1853 Rhacocleis acutangula: Karabağ, 1957. Comm. Fac. Sci. Univ. Ankara, Ser. C Sc. Nat. 8(1): 15. Platycleis affi nis: Fieber, 1853. Lotos 3: 150. Material examined: 3♂♂, 2♀♀, Balıkesir: Material examined: 4♀♀ (3♀♀ in alcohol), Edremit, Kazdağı, camping area-Ayı Deresi yolu, Balıkesir: Edremit, Kazdağı, camping area, ♂ ♀♀ 03.07.2004; 1♂ (in alcohol), same locality, 23.07.2004 25.08.2004 (H. Sevgili & Y. Durmuş); 1 , 4 , ♂ (H. Sevgili & Y. Durmuş); 4♀♀ (in alcohol), same Kazdağları, Ayı Deresi, 900 m, 30.07.1977; 1 , locality, 04.06.2005 (Y. Durmuş); 2♂♂, 5♀♀, same Kazdağları, Koşara yöresi, 1250 m, 30.07.1977 (S. locality, 19.07.2005 (Y. Durmuş & H. Sevgili). Salman). Calling song: We analyzed the songs of 2 males. Calling song: Th e songs of 2 males were recorded. Our data were similar to previous oscillograms Th e basic structure of the song was quite similar to published for this species (e.g. Heller, 1988; Ragge that of R. werneri from Greece and R. neglecta from and Reynolds, 1988; Oliveira et al., 2001) but slightly Italy (Heller, 1988; Ragge and Reynolds, 1998). Th e diff ered in the duration of the echemes and macro- quiet calling song, produced especially at night, microsyllables; the calling song consisted of mixed B echemes usually lasting about 0.6-4.35 s (mean: 2.68 s at 29 °C, n = 15) (Figure 16A). Th e echeme period was quite variable, between 3.3 and 13.5 s. Each A 5 s echeme consisted of macrosyllables (mean: 33) and C usually ended with a series of about 18 microsyllables (Figures 16B and 16C). Th e macrosyllables usually B lasted about 48-88 ms and were repeated at the rate of 1 s about 10-11/s. Th e duration of the microsyllables was about 1-5 ms and they repeated at the rate of about 29-33/s. C 250 ms Remarks: Although this species lives in the Figure 17. Song structure of male Rhacocleis acutangula: A) same habitat as the previous species, it prefers lower an echeme sequence of the male calling song, B) 10 altitudes. Adults could be seen in June-September complete echemes of the echeme sequence indicated in A, C) a faster oscillogram of the 4 echemes and were bioacoustically active at night and daytime. indicated in B.

642 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

Figure 19. Saga natoliae (male).

Figure 18. Rhacocleis acutangula (female). Material examined: 2♂♂, Balıkesir: Edremit, Kazdağı, camping area, 03.07.2004 (H. Sevgili & Y. consisted of a sequence of dense echemes, each Durmuş). lasting about 80-160 ms (mean: 136 ms, n = 27) Remarks: Th e range of this species covers the and consisting of about 3-6 diplosyllables (usually southern Balkans, northeastern Greece, some 5) (Figures 17A-17C). Th e period of echemes was eastern Aegean islands (Willemse, 1984), Th race, between 424 and 837 ms (mean: 542 ms at 29 °C, n Anatolia, and the Mediterranean’s eastern coast. = 27). Th e echemes were repeated regularly at a rate We collected this species (Figure 19) only from one of 2-3/s. locality associated with dense, tall grass or low shrubs Remarks: Th e genus Rhacocleis is represented by in the Yayla-Kamp area of the national park. Males 5 species in Turkey (Çıplak et al., 2002a; Otte et al., of S. natoliae began to call in the late aft ernoon and 2004). R. acutangula (Figure 18) and is an endemic evening. As with other Saginae species, this species is species spread throughout the vicinity of Balıkesir. In a carnivorous and showed nocturnal activity. Kazdağı National Park, this species lives in forest gaps It is quite rare in a few habitats and it is a very and along-roadside vegetation at altitudes between important predator for biocontrol in nature. It should 500-1000 m. It was bioacoustically active mainly be protected. at night. Adults were mostly present in August and Grylloidea September. Oecanthidae Tettigonia viridissima Linnaeus, 1758 Oecanthinae Gryllus (Tettigonia) viridissimus: Linnaeus, 1758. Oecanthus pellucens (Scopoli, 1763) Systema Naturae per Regna tria naturae (10th ed.) 1: 429. Gryllus pellucens: Scopoli, 1763. Entomologia carniolica, exibens Insecta Carniolae indigera: 32. Material examined: 1♂, Balıkesir: Edremit, Kazdağı, Tozlu, 03.07.2004 (H. Sevgili & Y. Durmuş). Material examined: 1♂ (in alcohol), Balıkesir: Edremit, Kazdağı, camping area, 23.07.2004 (H. Saginae Sevgili & Y. Durmuş). Saga natoliae Serville, 1839 Remarks: Th e tree cricket Oecanthus pellucens Saga natoliae: Serville, 1839 [1838]. Histoire has a wide distribution that is associated with the naturelle des insectes. Orthoptères: i-xviii, 1-776, pl. xerothermous ecological conditions of steppes and 1-14. woody steppe biotopes in Turkey. It lives typically in Previous records: 2♀♀, Balıkesir-Edremit yolu, bush trees and high herbs and stridulates during the 7. km, 14.07.1969 (Karabağ et al., 1971). night.

643 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Acridoidea Pamphagidae Orchaminae Orchamus davisi Uvarov, 1949 Orchamus davisi: B.P. Uvarov, 1949. Bull. Soc. Fauad l er. d’Ent. 33: 8.

Material examined: 3♂♂ (in alcohol), 1♀, Figure 21. Paranocaracris rubripes burri (female). Balıkesir: Edremit, Kazdağı, Milli Park girişi- camping area yolu, 500 m, 03.07.2004 (H. Sevgili & Remarks: P. rubripes burri (Figure 21) is an Y. Durmuş). endemic subspecies of Anatolia and widespread Remarks: Th is species (Figure 20) inhabits most in the northwestern part of Anatolia. Within the maquis and pine forest vegetation from sea level to Anatolian range of the genus, there are a large about 800 m. number of endemic subspecies having a restricted distribution (Demirsoy, 1973, 1977b; Ünal, 2002). Nocarodesinae Th is species can oft en be found around Sarıkız from Paranocaracris rubripes burri (Uvarov, 1949) May to September. Nocarodes burri: B.P. Uvarov, 1949. Bull. Soc. Acridoidea Fauadler. d’Ent. 33: 1-9. Acrididae Material examined: 4♂♂, 7♀♀ (in alcohol), Calliptaminae Balıkesir: Edremit, Kazdağı, Sarıkız, 1700 m, 23.07.2004; 7♂♂, 3♀♀ (in alcohol), same locality, Calliptamus italicus italicus (Linnaeus, 1758) 03.07.2004; 3♂♂, 4♀♀, same locality, 1730 m, 19- Gryllus italicus: Linnaeus, 1758. Systema Naturae 20.07.2005 (Y. Durmuş & H. Sevgili); 6♂♂, 2♀♀ (in per Regna tria naturae (10th ed.) 1: 432. alcohol), Kazdağı, Tozlu altı, 23.07.2004 (H. Sevgili Material examined: 6♂♂, 7♀♀ (in alcohol), & Y. Durmuş); 2♂♂, 6♀♀ (in alcohol), same locality, Balıkesir: Kazdağı, camping area, 800 m, 03.07.2004; 04.06.2005 (Y. Durmuş); 1♂, 2♀♀ (in alcohol), 1♂ (in alcohol), same locality, 23.07.2004 (H. Sevgili Kazdağı, Tozlu altı, 1360 m, 03.07.2004 (H. Sevgili & Y. Durmuş). & Y. Durmuş); 5♀♀ (in alcohol), same locality, 04.06.2005 (Y. Durmuş); 1♂, same locality, 1330 m, Remark: It occurs frequently in montane habitats 19.07.2005 (Y. Durmuş & H. Sevgili); 3♂♂, 1♀ (in and lives in dry, stony, open meadows. alcohol), Kazdağı, Tozlu mevki, 1400 m, 04.06.2005 Catantopinae (Y. Durmuş); 1♀, same locality, 20.07.2005 (Y. Durmuş & H. Sevgili). Pezotettix anatolica Uvarov, 1934 Pezotettix anatolica: Uvarov, 1934. Eos. 10: 112 Material examined: 1♂, 2♀♀ (in alcohol), Balıkesir: Edremit, Kazdağı, Zeytinli Çayı, 17.08.2005 (H. Sevgili). Remark: P. anatolica is a typical Mediterranean species. It prefers the ground of Pinus forests in lowlands to submontane areas. Cyrtacanthacridinae Anacridium aegyptium (Linnaeus, 1764) Gryllus aegyptium: Linnaeus, 1764. Museum Figure 20. Orchamus davisi (female). S.R.M. Ludovicae Ulricae Reginae, etc.: 138.

644 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

Material examined: 1♀, Balıkesir: Edremit, Akçay, 19.08.2005 (I. Acar). Remark: Afro-eremial species, typically found in dry habitats. Gomphocerinae Chorthippus (Glyptobothrus) biguttulus euhedickei Helversen, 1989 Chorthippus biguttulus euhedickei: Helversen, 1989. Articulata 4: 28. Material examined: 1♀, Balıkesir: Kazdağı, camping area, 19.07.2005, 800 m; 1♂♂, 4♀♀, Tozlu altı, 19.07.2005, 1330 m; 2♂♂, Tozlu mevki, 20.07.2005, 1400 m (leg. Y. Durmuş & H. Sevgili). Remarks: Th is subspecies, separated from the nominal one by bioacoustic characters, is known Figure 22. Chorthippus kazdaghensis (female). from Macedonia, northern Greece, and western Anatolia (Helversen, 1989; Willemse et al., 2009). Chorthippus (Glyptobothrus) bornhalmi Harz, Dociostaurus (Dociostaurus) maroccanus 1971 (Th unberg, 1815) Chorthippus bornhalmi: Harz, 1971. Atalanta Gryllus maroccanus: Th unberg, 1815. Mem. Acad. Munnerstadt 3: 336. Imp. Sc i. St. Peterburg 5: 211-301. Material examined: 1♂, 1♀ (in alcohol), Material examined: 5♂♂, 6♀♀ (1♂, 1♀ in Balıkesir: Kazdağı, camping area, 800 m, 04.06.2005 alcohol), Balıkesir: Kazdağı, camping area, 800 m, (Y. Durmuş); 8 ♂♂, 2 ♀♀, Kazdağı, Tozlu mevki, 03.07.2004; 3♂♂, 3♀♀, same locality, 19.07.2005 (H. 20.07.2005, 1400 m; 3♂♂, 1♀, camping area, Sevgili & Y. Durmuş). 19.07.2005, 800 m; 5♂♂, 6♀♀, Tozlu altı, 1330 m, Remark: It is a common, important plague locust 19.07.2005 (H. Sevgili & Y. Durmuş). in Turkey. Chorthippus (Glyptobothrus) kazdaghensis Mol Myrmeleotettix maculatus maculatus and Çıplak, 2005 (Th unberg, 1815) Chorthippus (Glyptobothrus) kazdaghensis: Mol Gomphocerus maculatus: Th unberg, 1815. Mem. and Çıplak, 2005. In: Çiplak, Mol, Sirin, Zeybekoglu Acad. Imp. Sci. St. Peterburg 5: 221. and Taylan. Trans. Amer. Entomol. Soc. 131(3-4): Material examined: 14♂♂, 20♀♀, Balıkesir: 470. Kazdağı, Sarıkız, 1730 m, 19.07.2005; 3♂♂, same Previous records: 6♂♂, 4♀♀, Balıkesir: Edremit, locality, 19.07.2005 (H. Sevgili & Y. Durmuş). Kazdağı, Sarıkız, 1700 m, 21.08.2003; 6♂♂, 3♀♀, Remark: Th is cold-tolerant species inhabits same locality, 1720 m, 23.07.2001; 2♂♂, 1♀, Kazdağı, highlands, and males called during the day in Açöldüren, 1400 m, 23.08.2001, Çıplak et al. 2005 (A. open areas within some alpine steppe vegetation in Mol). Kazdağı. Material examined: 1♂, 2♀♀, Balıkesir: Kazdağı, Stenobothrus fi scheri fi scheri (Eversmann, 1848) Sarıkız zirve, 1730 m, 20.07.2005 (H. Sevgili). Oedipoda fi scheri: Eversmann, 1848. Additamenta Remark: Th is recently described species (Figure quaedam levia ad Fischeri de Waldheim Orthoptera 22) is known only from the type locality. Rossica 11.

645 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Material examined: 2♂♂, 2♀♀, Balıkesir: Oedipoda caerulescens caerulescens (Linnaeus, Kazdağı, Sarıkız zirve, 1730 m, 20.07.2005; 3♂♂, 1758) 6♀♀, same locality, 23.07.2004 (Y. Durmuş & H. Gryllus (Locusta) caerulescens: Linnaeus, 1758. Sevgili). Systema Naturae per Regna tria naturae (10th ed.) 1: Remark: Like other Stenobothrus, the species lives 432. in montane to alpine meadows. We collected it from Material examined: 9♂♂, 2♀♀, Balıkesir: the highlands of Kazdağı. Kazdağı, camping area, 800 m, 19.07.2005 (Y. Remark: Th is cold-tolerant species inhabits Durmuş & H. Sevgili). highlands, and males called in open areas within Remarks: O. caerulescens was found to cohabit some alpine steppe vegetation in Kazdağı during the oft en with O. miniata and O. aurea in vegetation on day. rocks and bare slopes near the forest. However, O. Oedipodinae aurea was quite rare in the study area and preferred Acrotylus insubricus insubricus (Scopoli, 1786) midle altitudes. Gryllus insubricus: Scopoli, 1786. Delicidae forae Oedipoda miniata (Pallas, 1771) et faunae insubricae: 64. Gryllus miniatus: Pallas, 1771. Reisen durch Material examined: 1♀, Balıkesir: Edremit, verschiedene Provinzen des Russischen Reiches: 467. Kazdağı, camping area, 800 m, 04.06.2005 (H. Sevgili Material examined: 4♂♂, 5♀♀, Balıkesir: & Y. Durmuş); 1♂, 1♀ (in alcohol), Kazdağı, Zeytinli Kazdağı, camping area, 800 m, 19.07.2005 (H. Sevgili Çayı, 17.08.2005 (H. Sevgili); 1♂, 1♀ (in alcohol), & Y. Durmuş). ♂♂ ♂ Kazdağı, Tozlu altı, 23.07.2004; 2 (1 nymph), Tetrigoidea 3♀♀ (1♀ nymph) (in alcohol), Kazdağı, camping area-Ayı Deresi yolu, 25.08.2004 (H. Sevgili & Y. Tetrigidae Durmuş). Tetriginae Remark: In the summer, A. insubricus insubricus Uvarovitettix depressus (Brisout de Barneville, could be found in various sites of the national park, 1848) all located in the intertidal zone at the water’s edge, Tetrix depressa: Brisout de Barneville, 1848. Ann. from June to October. Soc. Ent. France (n.s.) (2)6: 424. Aiolopus strepens strepens (Latreille, 1804) Material examined: 1♂, 3♀♀ (in alcohol), Acrydium strepens: Latreille, 1804 [1802]. Hist. Balıkesir: Edremit, Kazdağı, Tozlu altı, Dere kenarı, Nat. Crust. Ins. 3(12): 154. 1300 m, 23.07.2005 (H. Sevgili & Y. Durmuş); 1♀, same locality, 1330 m, 19.07.2005 (Y. Durmuş & H. Material examined: 1♀ (in alcohol), Balıkesir: Sevgili). Edremit, Kazdağı, Zeytinli Çayı, 17.08.2005 (H. Sevgili); 1♀ (in alcohol), Kazdağı, camping area-Ayı Remark: Typically, this pygmy grasshopper is Deresi yolu, 25.08.2004 (H. Sevgili). found along streams and ponds. Th e species is, like all Tetrigidae species, hygrophilous and thermophilous. Remarks: During the study period, it was only found in sites with wet, fi ne soil. MANTODEA Oedipoda aurea (Uvarov, 1923) Oedipoda germanica aurea: B.P. Uvarov, 1923. Mantinae Ent. Monthly Mag. 3(9): 32. religiosa religosa (Linnaeus, 1758) Material examined: 4♂♂, Balıkesir: Kazdağı, Gryllus (Mantis) religiosa: Linnaeus, 1758. Syst. camping area, 800 m, 19.07.2005 (Y. Durmuş & H. Nat. (10th ed.) 1: 426. Sevgili). Material examined: 2♂♂ (in alcohol), Balıkesir:

646 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

Remark: Th is beautiful species was recorded from Savaştepe (Balıkesir) by Demirsoy (1977a).

Discussion Fauna In Kazdağı National Park, 34 species/subspecies belonging to 27 genera within 5 families were Figure 23. Rivetina fasciata (female). confi rmed to occur. In addition, 3 praying belonging to 3 genera were collected from Kazdağı Kazdağı, camping area-Ayı Deresi yolu, 800 m, (Table 2). 29.08.2004 (H. Sevgili & Y. Durmuş). Endemic taxa Miomantinae Isophya cania, Poecilimon luschani egrigozi, Rivetina fasciata Th unberg, 1815 Anadolua schwarzi, Rhacocleis acutangula, Paranocaracris rubripes burri, and Chorthippus Rivetina fasciata: Th unberg, 1815. Mem. Acad. (Glyptobothrus) kazdaghensis are endemic to the Sci. St. Petersb. 5: 292. region. Of these species, Isophya cania is quite Material examined: 1♂ (Figure 23) (in alcohol), remarkable since its distribution range is limited Balıkesir: Kazdağı, camping area, 29.08.2004 (A. only to the Balıkesir, Gönen, and Kazdağı areas. Demirsoy). Metrioptera (Broughtonia) arnoldi was previously Tarachodidae known from the Balkans. In this study, it was collected Tarachodinae from high meadows in August from Kazdağı. Anadolua schwarzi, which was known from western Iris oratoria (Linnaeus, 1758) Anatolia, was also found in the study area. One of Gryllus (Mantis) oratorius: Linnaeus, 1758. Syst. the interesting species from Kazdağı is Chorthippus Nat. (10th ed.) 1: 426. (Glyptobothrus) kazdaghensis, belonging to the C. Material examined: 1♀ (in alcohol), Balıkesir: demokidovi group, which occurs in open areas within Kazdağı, Zeytinli Çayı, 17.08.2004 (H. Sevgili & Y. forests (about 1400 m) and alpine zones at high Durmuş). altitudes.

Table 2. Number of species of Orthoptera and Mantodea in Kazdağı National Park.

Order Family Number of species Endemic

Orthoptera Tettigoniidae 16 4

ʺ Oecanthidae 1 --

ʺ Tetrigidae 1 --

ʺ Pamphagidae 2 1

ʺ Acrididae 14 1

Mantodea Mantidae 3 --

Total 37 6

647 Orthoptera and Mantodea fauna of Kazdağı (Ida) National Park with data on the calling songs of some bush-crickets

Altitudinal preferences and faunal connection nymph. Adults of the species of genera Metrioptera, Altitudinal distribution and habitat preferences Anadolua, Rhacocleis, and Chorthippus could be seen of Orthoptera are variable among species whose all the way into autumn within the study region. specifi city can be directly related to the availability When the Orthoptera fauna of Kazdağı is of food plants and climatic conditions. Our compared to the Orthoptera fauna of Greece and the observations showed that the species can be grouped Balkans (Willemse, 1984; Heller et al., 1998; Popov into 3 altitudinal zones (Figure 24). Th e Uvarovitettix, and Chobanov, 2004; Popov, 2007), similarities Oedipoda, Platycleis, Bucephaloptera, Saga, at both the generic and species level are evident Poecilimon pergamicus, P. anatolicus, Chorthippus, (with Bulgaria, for detail, see Popov, 2007). Th e Tylopsis, and Rhacocleis species were predominant Parapholidoptera castaneoviridis, Bucephaloptera in habitats within forest gaps and at water edges at bucephala, Saga natoliae, Platycleis spp., Tylopsis medium altitudes (800-1400 m) within Kazdağı, while lilifolia, Acrometopa syriaca, Isophya, Poecilimon, Isophya, Poecilimon luschani, Anadolua, Metrioptera, Metrioptera arnoldi, Paranocaracris, Myrmeleotettix Paranocaracris, Stenobothrus, Myrmeleotettix, maculatus, Chorthippus spp., Stenobothrus spp., and Chorthippus kazdaghensis preferred mostly and Oedipoda spp. species within Kazdağı are also high elevation habitats (1400-1700 m). Among distributed within the Th race region and Bulgaria. the recorded genera, Mantis, Iris, Acrometopa, Biogeography Anacridium, and Orchamus were observed at low to medium elevations (<800 m) within the mountains Koç (2007) stated that neotectonic movements, of Kazdağı National Park. As a typical Mediterranean especially from the Oligocene (about 30 mya) species, Orchamus davisi prefers habitats underneath and Miocene (about 16 mya) to the Pleistocene pines and along roadsides at low altitudes within the (about 1.8 mya), were the determinant factor in the national park. geomorphological features of Kazdağı. In the earlier periods of this time scale, Kazdağı was a lowland area, Within the national park, the fi rst species partly covered by lakes and seas. Climate and land from which adults are recorded aft er winter was morphology changed due to tectonic processes and Paranocaracris rubripes, which winters as a last instar the Mediterranean climate dominated the Kazdağı region during the Pliocene period (Fairbridge et al., 1997). Both geomorphological evolution and 25 climate change aff ected the distribution patterns of >1400 m numerous terrestrial fauna. Th e climatological shift s in the past triggered the distribution of many taxa 20 6 800-1400 m into numerous allopatrically isolated small habitats. A number of recent studies have documented that 4 the glacial cycles within the Pleistocene presented 15 <800 m 7 opportunities for dispersal and admixture, while the 3 warmer interglacial periods resulted in vicariance and 10 isolation for high altitude fauna such as those found within Kazdağı (e.g. Hewitt, 1996, 2000; Demirsoy, 2002; Çıplak, 2003b, 2008). It is possible that Kazdağı 11 5 10 off ered a single homogenous and continuous 1 periglacial and refugial mountain throughout the 1 1 1 Pleistocene. Some short-winged grasshoppers, bush- 0 e e a ae crickets, and toad grasshoppers have restricted ranges ida d id i d on rigi i g t due to their limited mobility. Most of them might ti Te Acr et T Oecanthidae Pamphagidae have evolved vicariantly, subsequent to the dispersal Figure 24. Th e number of the species and their distribution of their ancestors. For example, I. cania and I. thracica depends on the altitudes in Kazdağı National Park. are evaluated in the same subgroup phylogenetically

648 H. SEVGİLİ, A. İ. DEMİRSOY, Y. DURMUŞ

(Sevgili, 2004a), but their distribution does not as subspecies on many geographically isolated high overlap geographically due to their divergence from mountains throughout Anatolia, which is thought the ancestral taxa. While I. thracica has a northern to be the radiation center of the genus (Demirsoy, distribution and prefers suitable habitats at low 1973, 2002). Th ey prefer stony and rocky habitats and altitudes within Th race, I. cania has a restricted relatively poor alpine steppes, like P. rubripes burri, distribution within Balıkesir Province but occurs which inhabits mountain tops in Kazdağı. everywhere around Kazdağı, in glades and on high Most of the species from the area are mainly mountain tops, at elevations from 1000 to 1700 m. Mediterranean elements that occur all over the Another interesting example is Metrioptera arnoldi, Mediterranean zones, but in diff erent habitats (e.g. which has a disjunct distribution from the Balkan Tylopsis, Acrometopa, Bucephaloptera, Platycleis, region to Kazdağı. Present data show that both the Rhacocleis, Oecanthus, Orchamus, Calliptamus, Balkan (Bulgaria and Serbia) and the Anatolian Acrotylus). populations of M. arnoldi prefer mainly alpine zones within the mountains. Th ere are no high mountains In conclusion, this study provides baseline data between these populations and no faunistic record of the Orthoptera and Mantodea fauna of Kazdağı in this intermediate region. Th e distribution of Natural Park for future conservation eff orts. Th ese several taxa demonstrates the very close faunal mountains host several endemic species and some connections between the Balkan Peninsula and Asia isolated local populations belonging to highland- Minor (Kenyeres et al., 2009), i e. Parapholidoptera inhabiting species of Orthoptera. Understanding castaneoviridis and Poecilimon anatolicus. the exact biogeographical implications and faunal exchange between the Balkans and Anatolia must All species of the Anatolian endemic genus wait until additional data from the entire Biga Anadolua were distributed allopatrically on diff erent Peninsula are gathered. isolated mountains (e.g. Bozdağ, Sandras Dağı, Honaz Dağı) in the western part of Turkey (Karabağ, 1952, 1958), and the genus was represented by one Acknowledgements mountain-top species in Kazdağı, which covers the Th is study was supported by the Scientifi c northernmost distribution of the genus. In support and Technological Research Council of Turkey of this pattern, the distribution of Poecilimon (TÜBİTAK, YDABAG-103Y110). We are grateful luschani, Paranocaracris, and some short-winged to the National Park Department of Edremit and species of Chorthippus (the C. demokidovi group) H. Basri Avcı for help during the fi eld studies. Our shows similar distributional patterns within Anatolia thanks go to M. Ohl for kind help in working with (many genera biogeographically revised by Çıplak, the collection of Orthoptera at the Museum für 2003b, 2008). Most of them probably derived from a Naturkunde, Berlin (ZMHB). We are much indebted single ancestral species by habitat fragmentation and to D. Chobanov, who kindly lent us specimens of M. geographical isolation. For example, Paranocaracris arnoldi from Bulgaria, and İ.K. Sağlam, who corrected includes 23 taxa, known from Greece, Bulgaria, and the English. We thank 4 anonymous reviewers for the Anatolia. More than half of these taxa are distributed critical reading of the manuscript.

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