Anadolua schwarzi Ramme, 1939

Şirin, Deniz, Taylan, Mehmet Sait, Sevgili, Hasan, Akyildiz, Dilan Hevra & Güngör, Oğuzhan, 2024, A revision of Anadolua (Orthoptera: Tettigoniidae: Tettigoniinae) based on morphological and bioacoustic characters: Newly confirmed data suggesting a new synonym and two new species, Zootaxa 5529 (2), pp. 201-244 : 223-227

Anadolua schwarzi Ramme, 1939

https://orthoptera.speciesfile.org/otus/804972/overview

Anadolua schwarzi : Ramme 1939: p.68; Ramme 1951, p.362; Karabağ 1952, p.143; Burr 1952; Karabağ 1958, p.52; Önder et al. 1999, p.244; Sevgili et al. 2011, p.637; Tazegül & Önder 2012, p.115; Ünal 2018, p.14 .

Syn.n: Anadolua burri Karabağ, 1952 ; Karabağ 1952, p.142; Anadolua burri Karabağ, 1958 ; Karabağ 1958, p.52.

Type locality: Turkey: Muğla: S.W. Anatolia, Sandras-Mountain, Köyceğiz

Holotype: Male

Distribution: A. schwarzi is an Anatolian endemic species and is mainly distributed in the Aegean region and Toros Mountain range ( Fig. 17A ) ( Ramme 1939; Ramme 1951; Karabağ 1952; Karabağ 1958; Önder et al. 1999; Sevgili et al. 2011; Tazegül & Önder 2012; Ünal 2018).

Previous records include Anadolu schwarzi ( Ramme 1939; Ramme 1951; Karabağ 1952; Karabağ 1958; Önder et al. 1999; Sevgili et al. 2011; Tazegül & Önder 2012; Ünal 2018).

Turkey: SW Anatolien, Sandras Dagh, Köycegis , 2600 m, vii.1938, 1 male (leg.: Schwarz) (Holotype) ( MfN) ; Sandras Dagh , 2 males, 2 females (leg.: M. Burr) ( Karabağ 1952) ; Denizli Province, Babadağ , 1500–1900 m, 23.viii.1950, 1 male (leg.: P.H. Davis) ( NHMUK) ; İzmir province, Oedemish, Bozdag , 1300–1600 m, 16.viii.1950, 2 male, 1 female (leg.: P.H. Davis) ( NHMUK) ; Kütahya province: Gediz Dist. Saphane dagh 1800m. 29.viii.1950, 1 female (leg.: P.H. Davis) ( NHMUK) ; İzmir, ÖdemiŞ, Bozdağ Mountain , 12.viii.1992, 3 females, 26.viii.1992, 4 males, 5 females, 9.ix.1992, 5 males, 3 females, 1 female nymph, 7.x.1992, 2 males, 4 females, 26.vi.1993, 1 male (leg.: E. Tazegül) ; Kütahya, Gediz, Akdağ Mountain , 2070 m, 10.viii.2007, 2 males, 2 females, 1860–1910 m, 10.viii.2007, 5 males, 5 females, 2074 m, 7.ix.2016, 2 males, plus 1 male in alcohol, 1700 m, 7.ix.2016, 1 female- - Kütahya, Murat Mountain , 2030–2050 m, 11.viii.2007, 2 females- - Afyonkarahisar, Çay, Gelincikana hill, 2085 m, 1.ix.2015, 5 males, 6 females, plus 2 males, 4 females, 1 male nymph, 1 female nymph in alcohol- - Antalya, Elmalı, Akdağ, Gömbe , road to SubaŞı Plateau , 2025 m, 15.vii.2011, 1 male, 2 females- - Antalya, Gömbe, Akdağ Mountain , 2035 m, 20.ix.2011, 14 males, 3 females (all leg.: M. Ünal) ( AİBÜEM) ; Balıkesir, Edremit, Kazdağı Mountain , 25.viii.2004 (leg.: H. Sevgili, Y. DurmuŞ) ( HUZOM) .

Previous records as Anadolu burri ( Karabağ 1952; Karabağ 1958; Önder et al. 2011; Tazegül and Önder 2012; Ünal 2018).

Turkey: Anatolia, Denizli Prov. (Caria), Babadağ (Cadmus), 1700–1900 m, 24.viii.1950, 1 male (Holotype), 2 females (Paratypes) (leg.: P.H. Davis) ( NHMUK); Adana, Pozantı, Bulgar Dag, 2700 m, 2.ix.1949, 2 males (leg.: P.H. Davis) (Paratypes) ( NHMUK); Anatolien, Akchehir (Aksehir)-Göl, 1.x.1934, 1 female (leg.: Fuss) ( Ünal (2018) indicated this specimen is a paratype of A. schwarzi and holotype of A. rammei ) ( MfN); Konya, SeydiŞehir, TaraŞcı, Rezebeli, 1960 m, 2.ix.2015, 2 males, plus 1 male in alcohol, 1960–1990 m, 3.ix.2015, 2 males, 2 females, plus 1 male, 1 female in alcohol-- Karaman, Oyuklu Mountain, 2100 m, 7.ix.2015, 3 males, plus 1 male in alcohol- - Sivas, Gürün, Ziyaret Pass, 1900 m, 9.ix.2013, 1 male, 2 females, in alcohol, 1900–2000 m, 13.ix.2016, 1 female-- Antalya, Gömbe, Akdağ Mountain, 2035 m, 20.ix.2011, 4 males, 3 females-- Antalya, GündoğmuŞ, Geyik Mountains, Namaras-Karabul Plateau, 2120–2200 m, 20.ix.2017, 4 males, 2 females, plus 2 males, 3 females in alcohol-- Antalya, GündoğmuŞ, Geyik Mountains, Susambeli, 2300 m, 20.ix.2017, 3 males, 2 females, plus 4 males, 3 females in alcohol-- Antalya, GündoğmuŞ, Geyik Mountains, Namaras-Susambeli, 2200–2330 m, 14.vii.2017, 4 males nymph (in alcohol)--İçel, Anamur, TaŞeli Plateau, Koçpazarı, 2200 m, 22.ix.2017, 2 males, 2 females, plus 2 males, 2 females in alcohol-- Niğde, Darboğaz, Bolkar Mountains, Kapıgöl Lake, 2820 m, 24.ix.2017, 6 males, 1 female, plus 3 males, 2 females in alcohol (leg.: M. Ünal) ( AİBÜEM).

The examined material. Sandras Dagh, Köycegis, 2600 m, vii.1938, 1 male (leg.: Schwarz) (Holotype) ( MfN); Aksehir-Göl, Anatolien, 1.x.1934, 1 female (leg.: Fuss) (paratype of A. schwarzi and holotype of A. rammei ) ( MfN); Denizli Prov. (Caria), Baba dağ (Cadmus), 1700–1900 m, 24.viii.1950, 1 male (Holotype of A. burri ), 1 female (Paratype of A. burri ) (leg.: P.H. Davis) ( NHMUK) (detailed and scaled photo of these two samples were send to us for this research by scientist Beulah Garner from NHMUK); Antalya, Tahtalıdağ Mountain, N 36.536934, E 30.418848, 1828 m, 21.vii.2017, 1 male, (leg.: D. Sirin, A. Mol & M. S. Taylan); Antalya, Gömbe-I, Akdağ Mountain, the lower part of İkizgöl Lake, N 36.579563, E 29.584550, 2320 m N.N., 28. vii. 2021, 3 male, 4 female (leg.: D. Sirin, H. Sevgili, M.S. Taylan & A. Mol); Antalya, Gömbe-II, ridges of SubaŞı Plateau, N 36.573398, E 29.590977, 2181 m N.N., 28. vii. 2021, 4 male, 6 female (leg.: D. Sirin, H. Sevgili, M.S. Taylan & A. Mol); UŞak, Şaphane Mountain, N 39.060325, E 29.254171, 1954 m N.N., 09.viii.2022, 6 male, 4 female (leg.: D. Sirin, H. Sevgili & M.S. Taylan); Kütahya, Simav, Civanağa Mountain, Martlı location, N 39.256486, E 28.816930, 1932 m N.N., 10.viii.2022, 4 male, 3 female (leg.: D. Sirin, H. Sevgili & M.S. Taylan); Muğla, Sandras Mountain, 1 km to Kartal Lake, N 37.099695, E 28.844111, 1935 m N.N., 14.viii.2022, 1 male (leg.: D. Sirin, H. Sevgili & M.S. Taylan); Muğla, Sandras Mountain, Beyağaç Lake, picnic area, N 37.059972, E 28.805842, 1768 m N.N., 14.viii.2022, 4 male, 3 female (leg.: D. Sirin, H. Sevgili & M.S. Taylan); Denizli, Babadağ Mountain, end of the WPP’s (wind power plants), N 37.751041, E 28.865162, 1932 m N.N., 29.vii.2021, 4 male, 2 female leg.: D. Sirin, H. Sevgili, M.S. Taylan & A. Mol, 15.viii.2022, 6 male, 4 female (leg.: D. Sirin, H. Sevgili & M.S. Taylan); Denizli, Çameli, Yaylacık Mountain, N 36.978522, E 29.306078, 1993 m N.N., 16.viii.2022, 5 male, 5 female (leg.: D. Sirin & M.S. Taylan); İzmir, ÖdemiŞ, Bozdağ Mountain, Millik location, N 38.358381, E 28.096575, 1560 m N.N., 12/ 13.viii.2022, 5 male, 8 female (leg.: D. Sirin, H. Sevgili & M.S. Taylan).

Redescription. Head equal width with pronotum in both sexes. Fastigium slightly wider than scapus in both sexes. Maximum width of fastigium / Maximum width of scapus 1.0– 1.46 in male, 1.05–1.44 in female. A black band between the eyes is sometimes seen. The dorsal pronotum has an hourglass shape and is weakly flattened in the prozona but distinctly flattened in the metazona, and has a deep transverse depression behind the middle, and its posterior edge is broadly rounded. Maximum distance between lateral carinae / minimum distance between lateral carinae 2.53–3.58 in male, 2.22–2.97 in female. The median carinae is absent or slightly distinct, and the lateral carinae are prominent as round in the second part of the metazona. Hind femur long, its length 4.50–5.45 times of its maximum width in male 4.29–6.28 in female. Males micropterous and tegmen mostly reaches the end of second abdominal tergite, sometimes even surpassing it; the female tegmina is the squamipterous with wings which do not touch each other and only slight extend beyond the posterior margin of the pronotum. The speculum is in quadriform (generally trapezoidal), and usually no side vein development occurs. Costal vein absent or very weak; subcostal vein reaches the apex, generally parallel to radial vein, and, rarely, they may merge with a small vein near the apex. Medial vein quite sinuate at the apex and usually has a dichotomy at ¾ from the base. Cercus bearing an internal tooth; length is between 1.6 and 1.9 mm, shorter than half of anterior femur, with a strong tooth near between 1/3 and 1/4 from the base ( Fig. 9A–I ); last tergite of male with a broad excision, which nearly divides the plate in two lobes ( Fig. 5A–I ); Subgenital plate with a deep deltoideus excision ( Fig. 7A–I ). Female subgenital plate large, with a distinct angled excision, and a diamond like depression at the base of plate ( Fig. 8A–H ). The basal arms of the titillator are spineless, while the apical arms bear a small number of spines near the tip. The number of spines can differ not only between individuals but also between the two arms of an individual ( Fig. 11A–I ). Ovipositor longer than 18 mm and weakly structured, straight or slightly curved upwards. Males and females mostly blackish brown, dark orange or dark yellowish dorsally and brownish or yellowish ventrally in their general appearance. Male and female individuals generally have a pattern of brownish and/or blackish spots on their bodies on a grey background. Some individuals with yellowish and/or dirty yellowish body colour may not have spot patterns. The tegmina from base to apex is dark in colour up to nearly 1/2 part of it and the rest of it is yellowish or brownish ( Fig. 4A–I ). The whole leg colour matches the general body colour and pattern.

Coloration: Individuals of the species have a wide range of colours both within and between populations, including light yellow, dark yellow, light orange, grey, light brown, dark brown and black (Appendix 1–6).

Description of song: A total number of 74 calling song recordings from 20 male individuals have been examined ( Table 6 ). The calling songs have been recorded from males at 25–32°C in room conditions (by D. Şirin and M.S. Taylan). The male calling song consists of quite variable phrases in duration ( Fig. 17B ). The phrase structure is more similar to that of the A. moli sp. nov. than to the song type of A. bergeri sp. nov. Phrase duration ranges between 2.93 s and 111.95 s (19.09 ± 16.57 s). Each phrase contains variable number (16–644) triple syllables group. The phrase usually starts with a low amplitude syllable group series ranging from 1 to 11 and continues and lasts with a similar amplitude syllable group series. In a triple syllables group, syllable durations are quite similar, and the amplitude of the calling song varies (low-high-high amplitude and/or low-high-medium amplitude in the first, second and third syllables, respectively). Oscillographic analyses show that the period durations of the triple syllables group vary between 125.6 and 272.9 ms (172.4 ± 27.5) ( Table 6 ). The durations of the first, second and third syllable in the triple syllables group vary between 21.9–39.2 ms (27.6 ± 4.41), 19.6–40.5 ms (28.5 ± 5.25), and 20.4–46.6 ms (30.9 ± 5.22), respectively. Each syllable shows a standard structure, formulated as a short and rather indistinct soft opening hemisyllable + loud closing hemisyllable ( Fig. 17C–D ). Each closing hemisyllable begins with a low amplitude, reaches the highest level at approximately the midpoint, and lasts by decreasing in amplitude after this point. The spectral analyses show that Delta and Peak frequencies of the first low amplitude syllable group vary between 18.68 and 37.36 kHz (29.28 ± 3.01) for Delta and between 15.75 and 28.88 kHz (21.03 ± 3.17) for Peak. Delta and Peak frequencies of the Phrase syllable group (except first one) vary between 26.29 and 39.33 kHz (31.66 ± 2.30) for Delta and between 16.88 and 27.75 kHz (21.44 ± 3.40) for Peak ( Table 7 ).

Remarks. Ramme (1939) described Anadolua schwarzi by using colour variations of the characters (general body, face, fastigium, antennae, pronotum, elytra and tergites) and defined genus characters. Karabağ (1952) studied the British Museum material collected by Dr. M. Burr and Mr. P.H. Davis, and he described A. burri , A. rammei , and A. davisi . Karabağ (1952) separated it from A. burri by a number of characters, which are the following: (i) male cercus length (2 mm for A. schwarzi and 1.6 mm for A. burri ); (ii) tooth position on the cercus (approximately on the middle for A. burri and between base and the middle for A. schwarzi ); (iii) titillator shape and teeth number (basal branch very narrow and slender, apical branch strong and parallel sided, and serrate part with 1–3 teeth for A. burri and basal branch very long and wide, apical branch strong gradually narrowed, and serrate part with 3–4 teeth for A. schwarzi ); (iv) last tergite characters (with a deep excision which is narrowed in the first half and widened in the second half for A. burri and with a shallow and wide excision, which divides the plate in two parts for A. schwarzi ); (v) subgenital characters for male (with a rounded obtusangular excision A. burri and with a deep rounded excision for A. schwarzi ); (vi) subgenital characters for female (large and convex, with a obtusangular excision and a depression on the middle part for A. burri and large, with a distinct obtusangular excision, and a shallow depression which is widened at the base and apex and strongly constricted in the middle for A. schwarzi ); (vii) the character of the ending part of the female’s seventh sternite (with a very shallowly rounded obtusangular excision for A. burri and with a deep obtusangular excision for A. schwarzi ); and (viii) ovipositor characters (ovipositor short and almost straight for A. burri and ovipositor very long and straight for A. schwarzi ). The present study has examined the descriptive characters used in Karabağ’s (1952) key to separate the species. The characters have been found to be highly variable, in terms of both morphology and body size, across a wide range of populations (including type localities) and individuals. ( Tables 4 , 5 , Figs. 2–12 ). As a result, it is suggested here that A. burri should be synonymized with this species.

IUCN status proposal. This species has not been assessed for the IUCN Red List (2023-1). It is primarily threatened by agricultural activity by local people and also by the loss of habitat through conversion of suitable areas to cultivated land. Uncontrolled nomadic grazing and touristic activities are other important problems for the species. The species should be considered as a Vulnerable (B2b (i, iii) + c(iv)) status based on the extent of occurrence criteria of IUCN (https://www.iucnredlist.org/).

Figures

FIGURE 2. Dorsal wiev of male Pronotum. A–I, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, Antalya (Tahtalıdağ); G, UŞak (Şaphane); H, Kütahya (Civanaga); I, Denizli (Çameli); J–K, A. bergeri sp. nov.: J, Konya (Rezebeli); K, Antalya (GündoğmuŞ); L, A. moli sp. nov.: L, Konya (Sultandağ).

FIGURE 3. Dorsal wiev of female Pronotum.A–H, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, UŞak (Şaphane); G, Kütahya (Civanaga); H, Denizli (Çameli); I, A. bergeri sp. nov.: I, Konya (Rezebeli); J, A. moli sp. nov.: J, Konya (Sultandağ).

FIGURE 4. Tegmina of the males. A–I, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, Antalya (Tahtalıdağ); G, UŞak (Şaphane); H, Kütahya (Civanaga); I, Denizli (Çameli); J–K, A. bergeri sp. nov.: J, Konya (Rezebeli); K, Antalya (GündoğmuŞ); L, A. moli sp. nov.: L, Konya (Sultandağ).

FIGURE 5. Dorsal view of male last tergite. A–I, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, Antalya (Tahtalıdağ); G, UŞak (Şaphane); H, Kütahya (Civanaga); I, Denizli (Çameli); J–K, A. bergeri sp. nov.: J, Konya (Rezebeli); K, Antalya (GündoğmuŞ); L, A. moli sp. nov.: L, Konya (Sultandağ).

FIGURE 6. Dorsal view of female last tergite.A–H, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, UŞak (Şaphane); G, Kütahya (Civanaga); H, Denizli (Çameli); I, A. bergeri sp. nov.: I, Konya (Rezebeli); J, A. moli sp. nov.: J, Konya (Sultandağ).

FIGURE 7. Subgenital plate of the males. A–I, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, Antalya (Tahtalıdağ); G, UŞak (Şaphane); H, Kütahya (Civanaga); I, Denizli (Çameli); J–K, A. bergeri sp. nov.: J, Konya (Rezebeli); K, Antalya (GündoğmuŞ); L, A. moli sp. nov.: L, Konya (Sultandağ).

FIGURE 8. Subgenital plate of the females. A–H, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, UŞak (Şaphane); G, Kütahya (Civanaga); H, Denizli (Çameli); I, A. bergeri sp. nov.: I, Konya (Rezebeli); J, A. moli sp. nov.: J, Konya (Sultandağ).

FIGURE 9. Cerci of the males. A–I, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, Antalya (Tahtalıdağ); G, UŞak (Şaphane); H, Kütahya (Civanaga); I, Denizli (Çameli); J–K, A. bergeri sp. nov.: J, Konya (Rezebeli); K, Antalya (GündoğmuŞ); L, A. moli sp. nov.: L, Konya (Sultandağ).

FIGURE 10. Cerci of the females.A–H, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, UŞak (Şaphane); G, Kütahya (Civanaga); H, Denizli (Çameli); I, A. bergeri sp. nov.: I, Konya (Rezebeli); J, A. moli sp. nov.: J, Konya (Sultandağ).

FIGURE 11. Titillators of the males. A–I, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, Antalya (Tahtalıdağ); G, UŞak (Şaphane); H, Kütahya (Civanaga); I, Denizli (Çameli); J–K, A. bergeri sp. nov.: J, Konya (Rezebeli); K, Antalya (GündoğmuŞ); L, A. moli sp. nov.: L, Konya (Sultandağ).

FIGURE 12. Lateral view of Ovipositor. A–H, A. schwarzi: A, Muğla (Sandras); B, Denizli (Babadağ); C, İzmir (Bozdağ); D, Antalya (Gömbe-I); E, Antalya (Gömbe-II); F, UŞak (Şaphane); G, Kütahya (Civanaga); H, Denizli (Çameli); I, A. bergeri sp. nov.: I, Konya (Rezebeli); J, A. moli sp. nov.: J, Konya (Sultandağ).

FIGURE 17. Distribution map (A) and male calling song of Anadolua schwarzi (B—one complete phrase, C—a group of syllables series and D—six syllables, black arrow is the hypothetic opening hemisyllable movement; grey arrow is the hypothetic closing hemisyllable movement).