Russian Entomol. J. 18(1): 25–46 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2009

New data on calling signals of Gomphocerinae grasshoppers (Orthoptera: Acrididae) from South Siberia and the Russian Far East

Íîâûå äàííûå î ïðèçûâíûõ ñèãíàëàõ ñàðàí÷îâûõ ïîäñåìåéñòâà Gomphocerinae (Orthoptera: Acrididae) Þæíîé Ñèáèðè è Äàëüíåãî Âîñòîêà Ðîññèè

D.Yu. Tishechkin & M.A. Bukhvalova Ä.Þ. Òèøå÷êèí, Ì.À. Áóõâàëîâà

Department of Entomology, Faculty of Biology, M.V. Lomonosov Moscow State University, Vorobyevy Gory, Moscow, 119991 Russia. E- mail: [email protected]. Êàôåäðà ýíòîìîëîãèè Áèîëîãè÷åñêîãî ôàêóëüòåòà Ìîñêîâñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà èì. Ì.Â. Ëîìîíîñîâà, Âîðîáü¸âû Ãîðû, Ìîñêâà 119991, Ðîññèÿ.

KEY WORDS: grasshoppers, Acrididae, Gomphocerinae, songs, signals, variability, bioacoustics, taxonomy, Russia. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: ñàðàí÷îâûå, Acrididae, Gomphocerinae, ñèãíàëû, èçìåí÷èâîñòü, áèîàêóñòèêà, ñèñòåìàòèêà, Ðîññèÿ.

ABSTRACT. Oscillograms and descriptions of sig- for elucidation of status of dubious forms. For this nals of 12 species of Gomphocerinae grasshoppers (Or- reason, in last decades extensive studies of sounds thoptera: Acrididae) from 12 localities in Southern Si- produced by Gomphocerinae grasshoppers were per- beria and the Russian Far East are presented. Within a formed by a number of specialists. subspecies or monotypical species no differences in At present, sounds of european representatives of temporal pattern of signals and acoustic behaviour be- the subfamily are described in literature quite adequate- tween different populations were revealed. Songs of ly. Comprehensive data on signals of West-European different subspecies sometimes differ clearly from each Gomphocerinae can be found in the monograph by other, however. Signals of Aeropedellus variegatus Ragge and Reynolds [1998]. Oscillograms of the songs minutus Mistshenko, 1951 are described for the first of the most part of species from European Russia were time, taxonomic status of this form is discussed. recently published by Bukhvalova and Vedenina [Bukh- valova, 1993b, 1998; Bukhvalova & Vedenina, 1998; ÐÅÇÞÌÅ. Ïðèâåäåíû îñöèëëîãðàììû è îïèñà- Vedenina & Bukhvalova, 2001] and Savitsky [Savitsky, íèÿ ñèãíàëîâ 12 âèäîâ ñàðàí÷îâûõ ïîäñåìåéñòâà 2000, 2002, 2005, 2007; Savitsky & Lekarev, 2007]. Gomphocerinae (Orthoptera: Acrididae) èç 12 ìåñòî- Information on signals of Gomphocerinae grasshop- íàõîæäåíèé â Þæíîé Ñèáèðè è íà Äàëüíåì Âîñòî- pers of Siberia and the Russian Far East is far less êå Ðîññèè.  ïðåäåëàõ ïîäâèäà èëè ìîíîòèïè÷åñêî- complete, however. Sounds of certain East-palaearctic ãî âèäà ðàçëè÷èé âî âðåìåííîì ðèñóíêå ñèãíàëîâ è forms and also, of a number of widespread species from àêóñòè÷åñêîì ïîâåäåíèè ìåæäó ðàçíûìè ïîïóëÿ- the siberian and the far-eastern populations were de- öèÿìè âûÿâëåíî íå áûëî.  òî æå âðåìÿ, ñèãíàëû scribed by Bukhvalova and Vedenina [Bukhvalova, ðàçíûõ ïîäâèäîâ èíîãäà ðàçëè÷àþòñÿ äîâîëüíî ÷¸ò- 1993a, b, 1998; Bukhvalova & Vedenina, 1998; Veden- êî. Âïåðâûå îïèñàíû ñèãíàëû Aeropedellus variegatus ina & Bukhvalova, 2001]. Oscillograms of signals of minutus Mistshenko, 1951; îáñóæäàåòñÿ òàêñîíîìè- siberian species of Chorthippus Fieber, 1852 (with the ÷åñêèé ñòàòóñ ýòîé ôîðìû. exception of species from Transbaikalia) are presented in Benediktov [2005]. Nowadays it is evident that investigation of acoustic Introduction signals of grasshoppers from different parts of the range allows revealing certain taxonomic inexactitudes, which Grasshoppers of the subfamily Gomphocerinae are are not so easy to discover relying on morphological well-known for their elaborate and species-specific call- characters only. Taxonomic changes in the Glyptoboth- ing signals (songs). Temporal structure of songs is rus biguttulus group [Bukhvalova, 1993b, 1998] and widely used as a taxonomic character for discrimination the establishment of species status of Chorthippus calig- of closely related morphologically similar species and inosus Mistshenko, 1951 [Vedenina & Bukhvalova, 26 D.Yu. Tishechkin & M.A. Bukhvalova

2001] can be mentioned as examples. For this reason, zones) and, in certain species, also the Russian Far East. publication of oscillograms of widespread species from For these species oscillograms of songs from only one populations not studied before seems to be useful for population were published previously. Signals of Aero- taxonomic studies. pedellus variegatus minutus Mistshenko, 1951 are de- In the present paper oscillograms and descriptions scribed for the first time. of signals of 12 species of grasshoppers are presented. Seven of them are widespread european-siberian or Matherial and methods even transpalaearctic forms, still, the songs of individu- als from eastern parts of their ranges were not studied Recordings of songs were made under natural condi- until now with the exception of Aeropedellus variega- tions from caged or freely-moving insects with micro- tus variegatus (Fisher von Waldheim, 1846). For four phone MD–382 (upper frequency limit 12.5 kHz) and species oscillograms of signals of individuals from eu- cassette recorder "Elektronika–302–1" (upper frequen- ropean populations are given for comparison. cy limit 10 kHz) or minidisk recorder "Sony Walkman The ranges of five other ones include only Siberia MZ-NH900" (sampling frequency 44.1 kHz). In all (mainly, within the limits of steppe and forest-steppe cases manual mode of recording level control was used.

Fig. 1. Map of localities, where the recordings of the songs of Gomphocerinae were made: 1 — Northern part of Saratov Area, environs of Khvalynsk, near Ulyanino Village; 2 — Saratov Area, Krasnokutskiy Distr., Dyakovka Village; 3 — South-east of Saratov Area, environs of Ozinki Town; 4 — Altai Mountains, northern end of Teletskoe Lake, environs of Yaylyu Village; 5 — Irkutsk Area, Uda River near the mouth of Uk, about 30 km north-west of Nizhneudinsk (approx. 450 km north-west of Irkutsk); 6 — Buryatia, the valley of Irkut River in the environs of Mondy Village (about 80 km west of Kyren); 7 — Irkutsk Area, steppes about 30 km north-east of Elantsy along the road Elantsy–Ol’khon Island (approx. 180 km north-east of Irkutsk); 8 — Buryatia, Barguzin Depression, Ina River 3–4 km west from Ina Village (about 40 km north-east from Barguzin Town); 9 — Buryatia, 10 km east of Onokhoy (about 60 km east of Ulan-Ude), the valley of Bryanka Riv; 10 — Buryatia, the valley of Selenga River 5 km north from Novoselenginsk (20 km south-south-east from Gusinoozersk); 11 — South- east of Chita Area, the valley of Onon River 5–6 km west of Nizhniy Tsasuchey village; 12 — South-east of Chita Area, Klichkinskiy Mtn. Ridge at the crossing with Urulyunguy River (15 km west of Klichka Town); 13 — South-west of Khabarovsk Province, about 5 km north of Obluchye Town; 14 — Southern Maritime Province, Khankaiskiy District, 3–4 km north from Novokachalinsk Village, bank of Khanka Lake; 15 — Southern Maritime Province, Khasan Region, environs of Andreevka Village (about 35 km south-west from Slavyanka). Ðèñ. 1. Ìåñòà ñáîðà ñàðàí÷îâûõ ïîäñåìåéñòâà Gomphocerinae äëÿ çàïèñè çâóêîâûõ ñèãíàëîâ: 1 — ñåâåð Ñàðàòîâñêîé îáë., îêðåñòíîñòè Õâàëûíñêà, áëèç äåðåâíè Óëüÿíèíî; 2 — Ñàðàòîâñêàÿ îáë., Êðàñíîêóòñêèé ðàéîí, ñåëî Äüÿêîâêà; 3 — þãî-âîñòîê Ñàðàòîâñêîé îáë., îêðåñòíîñòè ïîñåëêà Îçèíêè; 4 — Àëòàé, ñåâåðíàÿ îêîíå÷íîñòü Òåëåöêîãî îçåðà, îêðåñòíîñòè ïîñåëêà ßéëþ; 5 — Èðêóòñêàÿ îáë., ð. Óäà áëèç óñòüÿ ð. Óê, îêîëî 30 êì ÑÇ Íèæíåóäèíñêà (ïðèáëèçèòåëüíî 450 êì ÑÇ Èðêóòñêà); 6 — Áóðÿòèÿ, äîëèíà Èðêóòà â îêðåñòíîñòÿõ ñåëà Ìîíäû (îêîëî 80 êì Ç Êûðåíà); 7 — Èðêóòñêàÿ îáë., ñòåïè îêîëî 30 êì Ñ ïîñåëêà Åëàíöû ïî äîðîãå íà î. Îëüõîí (îêîëî 180 êì Ñ Èðêóòñêà); 8 — Áóðÿòèÿ, Áàðãóçèíñêàÿ êîòëîâèíà, ð. Èíà 3–4 êì Ç îäíîèìåííîãî ñåëà (îêîëî 40 êì ê Ñ îò ïîñåëêà Áàðãóçèí); 9 — Áóðÿòèÿ, 10 êì Ñ Îíîõîÿ (îêîëî 60 êì Ñ Óëàí-Óäý), äîëèíà ð. Áðÿíêè; 10 — Áóðÿòèÿ, äîëèíà Ñåëåíãè, 5 êì Ñ Íîâîñåëåíãèíñêà (20 êì ÞÞ Ãóñèíîîçåðñêà); 11 — Þãî-âîñòîê ×èòèíñêîé îáë., äîëèíà Îíîíà 5–6 êì Ç ñåëà Íèæíèé Öàñó÷åé; 12 — Þãî-âîñòîê ×èòèíñêîé îáë., Êëè÷êèíñêèé õðåáåò ïðè ïåðåñå÷åíèè ñ ð. Óðóëþíãóé (15 êì Ç ïîñåëêà Êëè÷êà); 13 — Þãî-çàïàä Õàáàðîâñêîãî êðàÿ, îêîëî 5 êì Ñ ïîñåëêà Îáëó÷üå; 14 — Þæíîå Ïðèìîðüå, Õàíêàéñêèé ðàéîí, áåðåã îçåðà Õàíêà 3–4 êì Ñ Íîâîêà÷àëèíñêà; 15 — Þæíîå Ïðèìîðüå, Õàñàíñêèé ðàéîí, îêðåñòíîñòè ñåëà Àíäðååâêà (îêîëî 35 êì ÞÇ Ñëàâÿíêè). Signals of Gomphocerinae from Siberia and the Russian Far East 27

Air temperature was measured during or immediately COMPARATIVE NOTES. In general, the song of this after recording on the place where the singing insect was species is similar with this of M. maculatus (Thunberg, 1815) sitting. [see Ragge & Reynolds, 1998; Bukhvalova & Vedenina, The specimens whose signals were recorded have 1998; Savitsky, 2005] and differs from it only in quantitative been taken for taxonomic identification. All the materi- parameters. In M. palpalis echemes are shorter than in M. maculatus. Pauses between echemes in the former species al studied is deposited in the collection of Zoological are equal to or longer than echemes whereas in the latter one Museum of Moscow State University. echemes exceed pauses in duration. Geographical points where the recordings were made are shown on Fig. 1. The numbers of localities in the text Omocestus haemorrhoidalis (Charpentier, 1825) (the first item in the description of signals of each MATERIAL. 1. Northern part of Saratov Area, environs of species) correspond with these on the map. Khvalynsk, near Ulyanino Village, 19.VII.2004. Signals of 1 # are recorded at the temperature 34–36°C. In most species of singing insects calling male adopts 7. Irkutsk Area, steppes about 30 km north-east of Elantsy along a specific calling site (a tree trunk, thin twig, grass stem, the road Elantsy — Ol’khon Island (approx. 180 km north-east of etc.) and as a rule demonstrates specific behaviour Irkutsk), 15.VII.2003. Signals of 1 # are recorded at the tempera- during singing [e.g. Boulard, 2006]. Gomphocerinae ture 31°C. 12. South-east of Chita Area, Klichkinskiy Mtn. Ridge at the are no exception. Data on acoustic behaviour for each crossing with Urulyunguy River (15 km west of Klichka Town), species are provided after description of the calling 22.VII.2003. Signals of 1 # are recorded at the temperature 30–31°C. song. 14. Southern Maritime Province, Khankaiskiy District, 3–4 km Song terminology used in the present paper is ac- north from Novokachalinsk Village, bank of Khanka Lake, cepted after Ragge [e.g. Ragge, 1987]. Also, it is de- 21.VII.2006. Signals of 2 ## are recorded at the temperature 32– 34°C. scribed in Bukhvalova and Vedenina [1998]. Data on DISTRIBUTION. Transpalaearctic. distribution are given mainly after Bey-Bienko and REFERENCES TO SONG. Ragge and Reynolds [1998]: re- Mistshenko [1951] and Storozhenko [1986]. For spe- cordings from Western Europe; Savitsky [2005]: recordings from cies whose songs were described by Ragge and Rey- the Lower Volga Region. nolds [1998] only references to the recent articles not SONG AND ACOUSTIC BEHAVIOUR. The calling song is an echeme lasting for about 2–5 s (Figs 14–25). It mentioned in this book are given. References to earlier begins quietly and gradually increases in amplitude. Syllable works can be found in the monograph cited above. The repetition period becomes longer towards the end of the song order of genera is accepted after Storozhenko [1986], and averages 30–75 ms for the echeme as a whole. the subdivision of Chorthippus sensu lato — after Stor- Male as a rule sings on the soil among sparse vegetation. ozhenko [2002]. Usually, it produces signals with irregular intervals from 0.5 up to several minutes. Occasionally, 4–5 and more echemes follow with short breaks lasting from 10–15 up to 40–50 s. Descriptions of songs COMPARATIVE NOTES. O. haemorrhoidalis is a wide- spread species, still until now only signals of individuals Myrmeleotettix palpalis (Zubowsky, 1899) from european populations were described in literature. The MATERIAL. 6. Buryatia, the valley of Irkut River in the envi- only exception is the article by Bukhvalova and Zhantiev rons of Mondy Village (about 80 km west of Kyren), 4.VII.2007. [1994], where the description of signals of this species from Signals of 3 ## are recorded at the temperature 30–35°C. Southern Tyva (Central Siberia) is given. 10. Buryatia, the valley of Selenga River 5 km north from Presently, only signals of nominotypical subspecies are Novoselenginsk (20 km south-south-east from Gusinoozersk), investigated. In general structure the songs of individuals ° 7.VII.2007. Signals of 1 # are recorded at the temperature 33 C. from european, siberian and the far-eastern populations are 12. South-east of Chita Area, Klichkinskiy Mtn. Ridge at the crossing with Urulyunguy River (15 km west of Klichka Town), almost indistinguishable (e.g. see Figs 14, 18, 22 and 15–17, 22.VII.2003. Signals of 1 # are recorded at the temperature 28–30°C. 19–21, 23–25). Temporal parameters of signals in our re- DISTRIBUTION. Southern Siberia from Altai to Transbaikalia, cordings are in good agreement with data of Savitsky [2005] Southwest of Amur Area, Mongolia. for populations from the Lower Volga Region. Unfortunate- REFERENCES TO SONG. The only oscillogram of the record- ly, it is impossible to compare our results with data in Ragge ing of poor quality from southern Tyva is presented in Bukhvalova and Reynolds [1998] for the reason of considerable differ- and Zhantiev [1994]. ence of temperatures during recordings (21–22 and 26°C for SONG AND ACOUSTIC BEHAVIOUR. The calling recordings from Western Europe). song is an echeme-sequence lasting for 7–12 s and consisting of 14–20 echemes (Figs 2–13). In the main part of the song Omocestus viridulus (Linnaeus, 1758) echemes are separated by intervals of about 0.2–0.3 s. Pauses MATERIAL. 4. Altai Mountains, northern end of Teletskoe between echemes in the end of signal are somewhat longer Lake, environs of Yaylyu Village, 5.VII.1999. Signals of 1 # are and average 0.4–0.6 s in our recordings. Echeme duration recorded at the temperature 27–30°C. averages 200–250 ms. The song begins quietly reaching 5. Irkutsk Area, Uda River near the mouth of Uk, about 30 km maximum intensity after 4th-7th echeme. Each echeme con- north-west of Nizhneudinsk (approx. 450 km north-west of Irkutsk), ° sists of about 20–30 pulses (Figs 11–13). Signals of individ- 4.VII.2003. Signals of 1 # are recorded at the temperature 30–31 C. DISTRIBUTION. Transpalaearctic. uals from different localities are quite similar. REFERENCES TO SONG. Ragge and Reynolds [1998]: re- Male sings on the open bare places or on the soil among cordings from Western Europe; Savitsky [2005]: recordings from sparse vegetation every time moving to another place after Western Caucasus. producing a signal. Usually, it produces up to 4–5 signals SONG AND ACOUSTIC BEHAVIOUR. The calling after which stops singing. Pauses between signals average song is a prolonged echeme varying in duration from 15–20 from 3–5 up to 15–20 s and more. up to 30–40 s in our recordings (Figs 26–32). Syllable 28 D.Yu. Tishechkin & M.A. Bukhvalova repetition period gradually increases towards the end of the in our recordings is the same as in individuals from european signal and changes from 55–90 ms in the first half of echeme populations. Temporal parameters (echeme duration and syl- to 90–110 ms in the end of a song. lable repetition period) are in general agreement with data for Male usually sings among dense vegetation sitting on the populations from Western Europe [Ragge, 1986] and North grass stem. Signals follow each other with rather long irreg- Caucasus (Adygeya) [Savitsky, 2005]. The range of variabil- ular intervals about 1–2 minutes. ity of parameters in individuals from the latter locality is COMPARATIVE NOTES. General structure of the song wider, which is evidently, a result of larger sample studied.

2 Buryatia, Irkut valley (6) 6 2 s

3 Buryatia, Irkut valley (6) 2 s 4 7 8 2 s Buryatia, Selenga valley (10)

5

9 10 2 s Chita Area, env. Klichka (12)

6

Buryatia, Irkut valley (6) 11 200 ms

7

Buryatia, Selenga valley (10) 200 ms 8

Buryatia, Selenga valley (10) 12 200 ms

9

Chita Area, env. Klichka (12) 13 200 ms

10

Chita Area, env. Klichka (12) 200 ms 11 Buryatia, Irkut valley (6) 100 ms 12 Buryatia, Selenga valley (10) 100 ms 13 Chita Area, env. Klichka (12) 100 ms

Figs 2–13. Oscillograms of calling signals of Myrmeleotettix palpalis (Zubowsky, 1899) from different localities (No. of the locality on the map is given in brackets). Faster oscillograms of the parts of songs indicated as “6–13” are given under the same numbers. Ðèñ. 2–13. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Myrmeleotettix palpalis (Zubowsky, 1899) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “6–13”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Signals of Gomphocerinae from Siberia and the Russian Far East 29

Stenobothrus nigromaculatus DISTRIBUTION. Europe, Southern part of European Russia, (Herrich-Schäffer, 1840) Caucasus, Kazakhstan, Kyrgyzstan, Southern Siberia. MATERIAL. 7. Irkutsk Area, steppes about 30 km north-east of REFERENCES TO SONG. Ragge and Reynolds [1998]: re- Elantsy along the road Elantsy — Ol’khon Island (approx. 180 km cordings from Western Europe; Vedenina and Bukhvalova [2001]: north-east of Irkutsk), 15.VII.2003. Signals of 3 ## are recorded at recordings from Ukraine, North Caucasus, European Russia (Ros- the temperature 33°C. tov Area) and South Urals (Orenburg Area).

14 Saratov Area, Khvalynsk (1) 2 s 18 15 Irkutsk Area, Elantsy (7) 2 s 19

16 Chita Area, env. Klichka (12) 2 s 20 17 Maritime Prov., Khanka (14) 2 s 21 18

Saratov Area, Khvalynsk (1) 22 200 ms 19

Irkutsk Area, Elantsy (7) 23 200 ms

20 Chita Area, env. Klichka (12) 24 200 ms

21

Maritime Prov., Khanka (14) 25 200 ms 22

Saratov Area, Khvalynsk (1) 100 ms 23

Irkutsk Area, Elantsy (7) 100 ms 24

Chita Area, env. Klichka (12) 100 ms 25

Maritime Prov., Khanka (14) 100 ms

Figs 14–25. Oscillograms of calling signals of Omocestus haemorrhoidalis (Charpentier, 1825) from different localities (No. of the locality on the map is given in brackets). Faster oscillograms of the parts of songs indicated as “18–25” are given under the same numbers. Ðèñ. 14–25. Îñöèëëîãðàììû ïðèçûâíûõ Omocestus haemorrhoidalis (Charpentier, 1825) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “18–25”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 30 D.Yu. Tishechkin & M.A. Bukhvalova

SONG AND ACOUSTIC BEHAVIOUR. The calling uals short additional echemes present between longer ones in song is a series of echemes following each other with inter- the second phase of courtship signal. In our recordings these vals about 1–3 s (Figs 33–35). The echeme lasts about 1–2 s. echemes are hardly distinguishable (Fig. 38, in the beginning It begins quietly and reaches maximum towards the end. of the oscillogram) or almost entirely reduced (Figs 39–40). Syllable repetition period averages 10–15 ms in our record- Temporal parameters of courtship signals in our recordings ings. Male sings sitting on the ground. Usually, it produces are similar with these given in Ragge [1987]. 2–5 echemes and then stops singing for several minutes. When courting female, male can sing ceaselessly for Stauroderus scalaris (Fischer von Waldheim, 1846) about ten minutes and more producing a signal of quite MATERIAL. 1. Northern part of Saratov Area, environs of another kind consisting of two different phases (Figs 36–43). Khvalynsk, near Ulyanino Village, 19.VII.2004. Signals of 1 # are The first phase of courtship song is a succession of low- recorded at the temperature 32–33°C. amplitude echemes having a duration about 200–400 ms and 5. Irkutsk Area, Uda River near the mouth of Uk, about 30 km repeating with a period 0.9–1.4 s (Fig. 37). Syllable repeti- north-west of Nizhneudinsk (approx. 450 km north-west of Irkutsk), tion period in the echemes is almost the same as in the calling 1.VII.2003. Signals of 1 # are recorded at the temperature 30–31°C. song. This part can last for several minutes. Then the male 9. Buryatia, 10 km east of Onokhoy (about 60 km east of Ulan- abruptly starts producing the second phase (Figs 38–43). It Ude), the valley of Bryanka Riv., 1.VII.2006. Signals of 1 # are includes 4–8 echemes consisting of two parts each. Syllable recorded at the temperature 36°C. repetition period in the first part averages 20–30 ms, whereas DISTRIBUTION. Europe, Caucasus, Kazakhstan, mountains of Middle Asia, Southern Siberia from Urals to Buryatia, Mongolia, in the second part it is half as long. Overall duration of Northern China. echeme is about 0.9–1.2 s. REFERENCES TO SONG. Ragge and Reynolds [1998]: re- COMPARATIVE NOTES. Only signals of nominotypi- cordings from Western Europe; Vedenina and Bukhvalova [2001]: cal subspecies were studied until now. Temporal pattern of recordings from Greece, North Caucasus, Southern Kazakhstan and calling song of individuals from Irkutsk Area is the same as in Altai Mountains. european populations [Ragge, 1987; Vedenina & Bukhvalo- SONG AND ACOUSTIC BEHAVIOUR. The main part va, 2001]. of the calling song is an echeme-sequence lasting for 10–30 s Courtship signals in our recordings differ slightly from (Figs 44–47, 50–52 and 55–57). It consists of 20–40 echemes these described by Ragge [1987]. In west-european individ- repeating with a period of 450–750 ms. Each echeme consists

26 4 s Irkutsk Area, Uda River (5) 27

27

Irkutsk Area, Uda River (5) 29 2 s

28

Altai Mountains (4) 30 2 s

29

Irkutsk Area, Uda River (5) 31 200 ms 30 Altai Mountains (4) 32 200 ms 31

Irkutsk Area, Uda River (5) 100 ms

32 Altai Mountains (4) 100 ms

Figs 26–32. Oscillograms of calling signals of Omocestus viridulus (Linnaeus, 1758) from different localities (No. of the locality on the map is given in brackets). Faster oscillograms of the parts of songs indicated as “27” and “29–32” are given under the same numbers. Ðèñ. 26–32. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Omocestus viridulus (Linnaeus, 1758) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “27” è “29–32”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Signals of Gomphocerinae from Siberia and the Russian Far East 31 of two parts including 8–14 longer syllables and 7–12 shorter COMPARATIVE NOTES. All populations studied be- ones respectively. long to nominotypical subspecies. Temporal parameters, the Occasionally, male produce a long train of short echemes pattern of calling song and acoustic behaviour in individuals preceding the main part of signal (Figs 44–45, 47). In certain from Eastern Siberia are the same as in populations from cases such a prelude lasts for 20–30 s and more. Echeme Europe (Figs 44–45, 48, 50, 53, 55 — male from Saratov repetition period in it varies greatly (Figs 44–45, 47–49), Area, Figs 46–47, 49, 51–52, 54, 56–57 — males from each echeme consists of 4–8 syllables (Figs 48–49, 53–54). Irkutsk Area and Transbaikalia). Male prefers places with rather high dense vegetation and Glyptobothrus maritimus (Mistshenko, 1951) sings sitting on the grass stem. Usually, after the end of the song it leaves the stem and flies for a distance of several MATERIAL. 2. Saratov Area, Krasnokutskiy District, Dyakov- meters. Quite often it starts singing anew immediately after ka Village, 18.VII.2004. Signals of 3 ## are recorded at the temperature 30°C. landing. In actively singing male pauses between signals 7. Irkutsk Area, steppes about 30 km north-east of Elantsy along average 10–20 s. the road Elantsy — Ol’khon Island (approx. 180 km north-east of

33 2 s 34 34 200 ms 35 35

100 ms 36 37 38 2 s 39 40 37 200 ms 38 41 200 ms

39 200 ms 42 40 200 ms 43 41 100 ms

42

100 ms

43

100 ms

Figs 33–43. Oscillograms of signals of Stenobothrus nigromaculatus (Herrich-Schäffer, 1840) from steppes about 30 km north-east of Elantsy, Irkutsk Area (No. 7 on the map). 33–35 — calling song, 36–43 — courtship song. Faster oscillograms of the parts of songs indicated as “34–35” and “37–43” are given under the same numbers. Ðèñ. 33–43. Îñöèëëîãðàììû ñèãíàëîâ Stenobothrus nigromaculatus (Herrich-Schäffer, 1840) èç ñòåïåé îêîëî 30 êì Ñ ïîñåëêà Åëàíöû, Èðêóòñêàÿ îáë. (òî÷êà ¹ 7 íà êàðòå). 33–35 — ïðèçûâíûé ñèãíàë, 36–43 — ñèãíàë óõàæèâàíèÿ. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “34–35” è “37–43”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 32 D.Yu. Tishechkin & M.A. Bukhvalova

44 4 s Saratov Area, Khvalynsk (1) 45 45

Saratov Area, Khvalynsk (1) 48 50 2 s

46

51 2 s Irkutsk Area, Uda River (5) 47

49 52 Buryatia, Onokhoy (9) 2 s 48 Saratov Area, Khvalynsk (1) 53 200 ms 49 54 200 ms Buryatia, Onokhoy (9) 50 Saratov Area, Khvalynsk (1) 55 200 ms 51 Irkutsk Area, Uda River (5) 56 200 ms

52 200 ms Buryatia, Onokhoy (9) 57 53 Saratov Area, Khvalynsk (1) 100 ms 54 Buryatia, Onokhoy (9) 100 ms 55

Saratov Area, Khvalynsk (1) 100 ms

56

Irkutsk Area, Uda River (5) 100 ms

57

Buryatia, Onokhoy (9) 100 ms

Figs 44–57. Oscillograms of calling signals of Stauroderus scalaris (Fischer-Waldheim, 1846) from different localities (No. of the locality on the map is given in brackets). Faster oscillograms of the parts of songs indicated as “45” and “48–57” are given under the same numbers. Ðèñ. 44–57. Îñöèëëîãðàììû ïðèçûâíûõ Stauroderus scalaris (Fischer-Waldheim, 1846) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “45” è “48–57”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Signals of Gomphocerinae from Siberia and the Russian Far East 33

Irkutsk), 15.VII.2003. Signals of 4 ## are recorded at the temper- Males from European Russia and Eastern Siberia are ature 31°C quite similar with representatives of nominotypical (far-east- 8. Buryatia, Barguzin Depression, Ina River 3–4 km west from ern) subspecies both in general pattern and in temporal Ina Village (about 40 km North-East from Barguzin Town), 17, parameters of signals (Figs 58, 63, 69 — male from Saratov 18.VII.2007. Signals of 3 ## are recorded at the temperature 20 Area and 59–62, 64–68, 70–74 — males from the eastern and 25–27°C. 12. South-east of Chita Area, Klichkinskiy Mtn. Ridge at the regions of Russia). crossing with Urulyunguy River (15 km west of Klichka Town), Chorthippus hammarstroemi (Miram, 1907) 22.VII.2003. Signals of 2 ## are recorded at the temperature 38–40°C. MATERIAL. 10. Buryatia, the valley of Selenga River 5 km 14. Southern Maritime Province, Khankaiskiy District, 3–4 km north from Novoselenginsk (20 km south-south-east from Gusi- north from Novokachalinsk Village, bank of Khanka Lake, 21.VII.2003. noozersk), 8, 9.VII.2007. Signals of 2 ## are recorded at the Signals of 3 ## are recorded at the temperature 33–34°C. temperature 26 and 31–32°C. DISTRIBUTION. Crimea, southern part of European Russia, 12. South-east of Chita Area, Klichkinskiy Mtn. Ridge at the North Caucasus up to subalpine zone (2000–2300 m above sea crossing with Urulyunguy River (15 km west of Klichka Town), level), Kazakhstan including North Tien-Shan, Turkmenistan (Ko- 22.VII.2003. Signals of 2 ## are recorded at the temperature 30–31°C. pet-Dagh Mtn. Ridge), Irkutsk Area, Transbaikalia, southern re- 14. Southern Maritime Province, Khankaiskiy District, 3–4 km gions of the Russian Far East [Bukhvalova, 1998]. Until now was north from Novokachalinsk Village, bank of Khanka Lake, not found in Western Siberia (Altai, Tyva). Western boundary of the 17.VII.2002 and 21.VII.2006. Signals of 3 ## are recorded at the range is obscure. temperature 23–25 and 27–28°C. REFERENCES TO SONG. Bukhvalova [1993b, 1998], Bukh- 15. Southern Maritime Province, Khasan Region, environs of valova and Zhantiev [1994]: recordings from Crimea, southern Andreevka Village (about 35 km south-west from Slavyanka), regions of European Russia, North Caucasus, southern Turkmeni- 14.VII.2006. Signals of 1 # are recorded at the temperature 23–25°C. stan, southern Kazakhstan, Amur Area and the Southern Maritime DISTRIBUTION. Southern Siberia from Altai to Transbaikalia Province; Benediktov [2005]: recording from Irkutsk Area; Sav- (northwards as far as Yakutia), southern part of the Russian Far itsky and Lekarev [2007]: recordings from the eastern parts of the East, Mongolia, south-east of China. Lower Volga Region. REFERENCES TO SONG. Benediktov [2005]: recordings from SONG AND ACOUSTIC BEHAVIOUR. Similarly with Tyva. other species of Glyptobothrus Chopard, 1951 from biguttu- SONG AND ACOUSTIC BEHAVIOUR. The calling lus-group, males of G. maritimus prefer to produce their song of a single individual is an echeme lasting for 10–20 s songs sitting on the open places or on the ground among grass (Figs 75–78). In actively singing male 5–10 and more echemes stems. This is one of the most actively-singing species of can follow each other with short breaks about 2–10 s. Sylla- Gomphocerinae. Males readily sing in reply to each other so ble repetition period averages 200–300 ms at the temperature that at times their songs sound as an unceasing chorus. In a 27–31°C and 300–500 ms at 23–25°C. Sometimes it becomes single male pauses between songs usually average 1–3 min- slightly longer towards the end of the echeme. The echeme utes and more. In males from the siberian and the far-eastern begins quietly reaching maximum intensity in 5–10 s. Each populations spontaneously produced signal as a rule consists syllable includes low- and high-amplitude parts (Figs 79– of initial echeme having duration about 3.5–8 s after which 102). In the syllables from the beginning of the song the 1–2 shorter ones (approximately 2.5–3 s each) follow (Figs former part is almost indistinguishable, whereas the latter one 58–62). As an exception signal consisting of seven echemes includes 5–7 short pulses separated by distinct gaps (Figs 79, once was registered in the male from Barguzin Depression. If 82, 85, 88, 91, 94, 97, 100). Towards the end of the signal the male produces his song in reply to another one, duration low-amplitude part gradually becomes more distinct, where- of the first echeme usually does not exceeds 3–4 s. as the gaps in the high-amplitude one disappear partly or Each syllable consists of one low-amplitude rather long entirely (Figs 80–81, 83–84, 86–87, 89–90, 92–93, 95–96, fragment followed by 3–5 high-amplitude ones usually sepa- 98–99, 101–102). As a rule, male sings sitting on the grass, rated by more or less distinct gaps (Figs 63–74). The structure usually preferring the place in the middle or in the lower half of syllables is rather variable and occasionally has distinct of the stem. differences even in the males from the same locality (Figs 67, Male courting female produce the same signal, but the 73 and 68, 74). Syllable repetition period averages 100–150 duration of echemes became more variable and averages from ms in our recordings. 5–8 up to 38–40 s (Figs 103–109). Syllable shape and repe- COMPARATIVE NOTES. Comparative investigation of tition period remain the same as in the calling song (Figs songs of this species from different localities was performed 110–112). Quite often the male sitting next to female sings by Bukhvalova [1998]. Oscillograms of signals of individu- ceaselessly for about 10 minutes and more. In this situation it als from Crimea, North Caucasus, southern part of European produces echemes more regularly with shorter intervals up to Russia, Turkmenistan, Kazakhstan and the Russian Far East 4–5 s (Figs 103–105). Then the male suddenly stops singing are given in her paper. Two oscillograms of the only signal and makes an attempt of copulation. It produces a succession from Irkutsk are presented in the article by Benediktov [2005], of syllables of variable structure at this moment (Figs 113– descriptions of songs of G. maritimus from Transbaikalia 116). Duration of syllables averages 200–250 ms and their were absent in literature. repetition period is about 250–700 ms in our recordings. As it was shown by Bukhvalova [1998], certain forms of Occasionally, these syllables alternate with short successions G. maritimus from North Caucasus (G. maritimus tsejensis of discrete pulses similar with these in syllables from the (Bukhvalova, 1993)), Turkmenistan (G. maritimus karakalen- beginning of calling (Figs 114–115). sis (Sytshev et Woznessenskij, 1996)) and Southern Kazakh- COMPARATIVE NOTES. Only oscillogram of one call- stan differ from european and the far-eastern ones. They ing signal of male from Tyva (Kyzyl) exists in literature usually produce single echemes of greater duration (10–20 [Benediktov, 2005]. Both general pattern and temporal pa- s); in males from Caucasus and Turkmenistan gaps in sylla- rameters of signals of males from different populations are bles are more distinct, than in individuals from steppes of quite similar. It should be noted, that all the material studied European Russia. belongs to nominotypical subspecies. 34 D.Yu. Tishechkin & M.A. Bukhvalova

58 2 s Saratov Area, Dyakovka (2) 63 59 Buryatia, Barguzin (8) 64 2 s 60 2 s Irkutsk Area, Elantsy (7) 65 61 Chita Area, env. Klichka (12) 66 2 s 62 Maritime Prov., Khanka (14) 2 s 67 63 200 ms Saratov Area, Dyakovka (2) 69 64 Buryatia, Barguzin (8) 70 200 ms 65

Irkutsk Area, Elantsy (7) 71 200 ms 66 200 ms Chita Area, env. Klichka (12) 72 67

Maritime Prov., Khanka (14) 73 200 ms 68 Maritime Prov., Khanka (14) 74 200 ms 69 Saratov Area, Dyakovka (2) 100 ms 70 Buryatia, Barguzin (8) 100 ms 71 Irkutsk Area, Elantsy (7) 100 ms 72 Chita Area, env. Klichka (12) 100 ms 73 Maritime Prov., Khanka (14) 100 ms 74 Maritime Prov., Khanka (14) 100 ms Figs 58–74. Oscillograms of calling signals of Glyptobothrus maritimus (Mistshenko, 1951) from different localities (No. of the locality on the map is given in brackets). Faster oscillograms of the parts of songs indicated as “63–67” and “69–74” are given under the same numbers. Ðèñ. 58–74. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Glyptobothrus maritimus (Mistshenko, 1951) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “63–67” è “69–74”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Signals of Gomphocerinae from Siberia and the Russian Far East 35

75 79 Buryatia, Selenga (10) 80 81 2 s 76 82 2 s Chita Area, env. Klichka (12) 83 84 77 Maritime Prov., Andreevka (15) 2 s 85 86 87 78

Maritime Prov., Khanka (14) 88 89 2 s 90 79 Buryatia, Selenga (10) 91 200 ms 80 Buryatia, Selenga (10) 92 200 ms 81 Buryatia, Selenga (10) 93 200 ms 82 Chita Area, env. Klichka (12) 200 ms 94 83 Chita Area, env. Klichka (12) 200 ms 95 84 200 ms Chita Area, env. Klichka (12) 96 85

Maritime Prov., Andreevka (15) 97 200 ms 86

Maritime Prov., Andreevka (15) 98 200 ms 87 200 ms Maritime Prov., Andreevka (15) 99 88 200 ms Maritime Prov., Khanka (14) 100 89 200 ms Maritime Prov., Khanka (14) 101 90 200 ms Maritime Prov., Khanka (14) 102 Figs 75–90. Oscillograms of calling signals of Chorthippus hammarstroemi (Miram, 1907) from different localities (No. of the locality on the map is given in brackets). Faster oscillograms of the parts of songs indicated as “79–102” are given under the same numbers. Ðèñ. 75–90. Îñöèëëîãðàììû ïðèçûâíûõ Chorthippus hammarstroemi (Miram, 1907) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “79–102”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 36 D.Yu. Tishechkin & M.A. Bukhvalova

Chorthippus caliginosus Mistshenko, 1951 each (Figs 117–121). If male is not disturbed, the signal can MATERIAL. 10. Buryatia, the valley of Selenga River 5 km north last up to 20–30 s and more. Occasionally, the amplitude of from Novoselenginsk (20 km south-south-east from Gusinoozersk), the song slightly increases towards its end. Fragments in it 8.VII.2007. Signals of 2 ## are recorded at the temperature 31–32°C. repeat with the period about 2–3 s, duration of each fragment DISTRIBUTION. Southern regions of Transbaikalia, Amur averages 0.8–1.0 s. Syllables repetition period in echemes is Area and Khabarovsk Region, also, south-east of China. about 10–15 ms. Usually (but not always), initial syllable in REFERENCES TO SONG. Vedenina and Bukhvalova [2001]: the echeme has higher amplitude than succeeding ones. recordings from two localities in Chita Area. Male sings sitting on the grass among dense vegetation. SONG AND ACOUSTIC BEHAVIOUR. Calling song is Sometimes, two or three neighbouring individuals sing in a succession of short fragments consisting of two echemes turn, so that their echemes alternate.

91 Buryatia, Selenga (10) 100 ms 92 Buryatia, Selenga (10) 100 ms

93 Buryatia, Selenga (10) 100 ms 94 Chita Area, env. Klichka (12) 100 ms

95 Chita Area, env. Klichka (12) 100 ms 96

Chita Area, env. Klichka (12) 100 ms 97

Maritime Prov., Andreevka (15) 100 ms

98 Maritime Prov., Andreevka (15) 100 ms

99

Maritime Prov., Andreevka (15) 100 ms 100 Maritime Prov., Khanka (14) 100 ms

101

Maritime Prov., Khanka (14) 100 ms

102

Maritime Prov., Khanka (14) 100 ms

Figs 91–102. Oscillograms of the parts of calling signals of Chorthippus hammarstroemi (Miram, 1907) from different localities (also, see Figs 75–90, No. of the locality on the map is given in brackets). Ðèñ. 91–102. Îñöèëëîãðàììû ôðàãìåíòîâ ïðèçûâíûõ ñèãíàëîâ Chorthippus hammarstroemi (Miram, 1907) èç ðàçíûõ ãåîãðàôè- ÷åñêèõ òî÷åê (ñì. òàêæå ðèñ. 75–90, â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Signals of Gomphocerinae from Siberia and the Russian Far East 37

103

4 s Buryatia, Selenga (10) 104 104

Buryatia, Selenga (10) 2 s 105

Chita Area, env. Klichka (12) 2 s

106 Maritime Prov., Khanka (14) 4 s 107 107 110 112 Maritime Prov., Khanka (14) 111 2 s 108 2 s Maritime Prov., Khanka (14) 109

Maritime Prov., Khanka (14) 2 s 110 Maritime Prov., Khanka (14) 200 ms

111 Maritime Prov., Khanka (14) 200 ms 112 Maritime Prov., Khanka (14) 200 ms

113 Maritime Prov., Khanka (14) 114 115 2 s

114

Maritime Prov., Khanka (14) 116 200 ms 115 Maritime Prov., Khanka (14) 200 ms 116

Maritime Prov., Khanka (14) 100 ms

Figs 103–116. Oscillograms of signals of Chorthippus hammarstroemi (Miram, 1907) from different localities (No. of the locality on the map is given in brackets). 103–112 — courtship song, 113–116 — precopulatory song. Faster oscillograms of the parts of songs indicated as “104”, “107”, “110–112” and “114–116” are given under the same numbers. Ðèñ. 103–116. Îñöèëëîãðàììû ñèãíàëîâ Chorthippus hammarstroemi (Miram, 1907) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). 103–112 — ñèãíàë óõàæèâàíèÿ, 113–116 — ïðåêîïóëÿöèîííûé ñèãíàë. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “104”, “107”, “110–112” è “114–116”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 38 D.Yu. Tishechkin & M.A. Bukhvalova

COMPARATIVE NOTES. Only signals of Ch. caligino- Gusinoozersk), 7.VII.2007. Signals of 1 # are recorded at sus from two localities in Chita Area were described in the temperature 30–32°C. literature [Vedenina & Bukhvalova, 2001]. Songs of males DISTRIBUTION. Tyva, Southern Transbaikalia, south- from Buryatia and Chita Area have no significant differences. ern regions of the Russian Far East, Korea. REFERENCES TO SONG. Bukhvalova and Vedenina Schmidtiacris schmidti (Ikonnikov, 1913) [1998]: recording from Amur Area (as Chorthippus schmidti). MATERIAL. 10. Buryatia, the valley of Selenga River 5 SONG AND ACOUSTIC BEHAVIOUR. Calling song km north from Novoselenginsk (20 km south-south-east from consists of single or irregularly repeated short echemes hav-

117 118 2 s 118

120 121 200 ms 119 200 ms 120 100 ms

121

100 ms

122

124 2 s 123 2 s 125 126 124 200 ms 127 125 200 ms 126 200 ms 128 127 100 ms 128 100 ms

Figs 117–128. Oscillograms of calling signals of Chorthippus caliginosus Mistshenko, 1951 (117–121) and Schmidtiacris schmidti (Ikonnikov, 1913) (122–128) from Selenga valley, Buryatia (No. 10 on the map). Faster oscillograms of the parts of songs indicated as “118”, “120–121” and “124–128” are given under the same numbers. Ðèñ. 117–128. Îñöèëëîãðàììû Chorthippus caliginosus Mistshenko, 1951 (117–121) è Schmidtiacris schmidti (Ikonnikov, 1913) (122–128) èç äîëèíû Ñåëåíãè, Áóðÿòèÿ (òî÷êà ¹ 10 íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “118”, “120–121” è “124– 128”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Signals of Gomphocerinae from Siberia and the Russian Far East 39 ing duration about 0.6–1 s and including 5–7 syllables each the same male (on Figs 144–146 three parts of one song are (Figs 122–128). Syllable repetition period averages 120–150 presented). Syllable repetition rate is almost the same as in ms in our recordings. Ae. variegatus variegatus and averages 60–70 syllables/s at Usually, singing males form aggregations in wet places the temperature 27°C and 80 syllables/s at 33°C (Figs 153– with high dense grass (e.g. on gramineous vegetation on river 162). Occasionally, syllables are grouped in pairs and sepa- banks). Calling males sit on the tall grass stems about 0.5–1 rated by distinct gaps, but their repetition rate remains the m above the ground. Neighbouring individuals produce sig- same (Figs 157, 162). We emphasize that such signal was nals with irregular intervals from 4–6 s up to one minute and registered from the intact male with two hind legs. more so that echemes of different males alternate. Singing male usually sits on the ground and walks from COMPARATIVE NOTES. Presently, oscillograms of one place to another during pauses between echemes. If it the only signal of individual from Amur Area were published. starts singing when moving, amplitude distortions present in Our recordings are quite similar with this presented in Bukh- the beginning of the echeme (e.g. Fig. 147, the last echeme). valova and Vedenina [1998]. In the region of our observations Ae. variegatus minutus Schmidtiacris Storozhenko, 2002 differs from Chorthip- inhabited steppes, preferring wet moderately salted meadows pus s. str. both in morphological and karyological characters with gramineous vegetation on the shores of shallow salted [Sergeev & Bugrov, 1988; Storozhenko, 2002]. Nonetheless, lakes. temporal pattern of syllables in this species is quite similar COMPARATIVE NOTES. There is no consensus of with this in certain species from other genera, including opinion among authors as to the taxonomic status of certain Chorthippus and Glyptobothrus (Figs 127–128 and 69–74, siberian forms in the genus Aeropedellus Hebard, 1935. The 87). This fact is in a good agreement with the opinion of other form with swelled fore tibiae from Minusinsk Depression and authors [Ragge & Reynolds, 1998; Savitsky, 2005] that Irkutsk Area was described almost contemporaneously by bioacoustic characters in taxonomy of Gomphocerinae are of Miram [1906–1907] as Ae. reuteri (Miram, 1907) and Ikon- little or no value on superspecies level. nikov [1911] under the name Ae. simillimus (Ikonnikov, 1911). Later it was treated as a synonym of Ae. variegatus by Aeropedellus variegatus variegatus Tarbinskiy [1931]. Mistshenko subdivided Ae. variegatus s. (Fisher von Waldheim, 1846) str. into five subspecies, among them Ae. variegatus minutus MATERIAL. 3. East of Saratov Region, 10 km east of Ozinki from steppes of Irkutsk Area [Bey-Bienko & Mistshenko town, 25.VI.1996. Signals of 3 ## are recorded at the temperature 1951]. Also, he considered Ae. reuteri as a separate species; 27–30°C. this opinion was accepted by Ivanova [1967] and Sergeev 11. South-east of Chita Area, the valley of Onon River 5–6 km [1986]. On the other hand, Berezhkov [1956] in his compre- west of Nizhniy Tsasuchey village, 19.VI.1995. Signals of 3 ## hensive work on grasshoppers of Western Siberia points out are recorded at the temperature 29°C. DISTRIBUTION. Mountains of Western Europe, European that he has never collected this form in the region under Russia, North Caucasus, South-eastern Kazakhstan, Siberia (north- investigation. In other articles on grasshoppers of Siberia for wards as far as Yakutia). the most part the name Ae. variegatus is used without speci- REFERENCES TO SONG. Bukhvalova and Vedenina [1998]: fying the subspecies. Ae. variegatus minutus was only men- recordings from two localities listed above. tioned in the catalogue of Orthoptera of Northern Asia in the SONG AND ACOUSTIC BEHAVIOUR. The calling monograph by Sergeev [1986]. song is a series of several echemes (2–6 in our recordings) It should be pointed out that morphological characters in repeated with intervals of 2.5–5 s (Figs 129–133). The echeme Aeropedellus are rather variable. For this reason differences lasts for 1.8–3 s (mean value 2.4±0.10 s) reaching maximum between certain forms sometimes are obscure. Fore tibiae in in the second half or in the end of its duration. The echeme our specimens from Irkutsk Area (Fig. 180) are slightly wider consists of short simple syllables following each other at a than in Ae. variegatus variegatus from Saratov (Figs 173– rate 70–80/s (recording at 27°C from Saratov Area) or 85–90/ 174) and Chita Areas (Figs 176–177), but are not so swelled s (recording at 29°C from Chita Area) almost without gaps as in Ae. reuteri (Figs 183–184; topotypes of Ae. simillimus (Figs 134–143). with the label “Minusinsk, P.P. Sushkin, 10.VI.1902”). In Male usually produce his song sitting on the soil or on the the form of lateral carinae of pronotum and in width ratio of grass close to the ground. claw and arolium males from Irkutsk Area fall into Ae. COMPARATIVE NOTES. A description of calling song variegatus minutus (Fig. 181–182), however among Ae. var- of this species illustrated by three oscillograms of the record- iegatus variegatus specimens with wide arolium also can be ing from Chita Area is given in Bukhvalova and Vedenina found (Fig. 178). [1998]. More comprehensive description is provided here for The specimens from Chita Area are somewhat larger than comparison with the song of the next subspecies. the ones from Saratov Area, still, according to the key in Bey- Bienko and Mistshenko [1951], the material from both local- Aeropedellus variegatus minutus Mistshenko, 1951 ities belongs to nominotypical subspecies. It is worth noting MATERIAL. 7. Irkutsk Area, steppes about 30 km north-east of that Ae. variegatus variegatus was recorded from South- Elantsy along the road Elantsy — Ol’khon Island (approx. 180 km Eastern Transbaikalia by Popov [1964]. north-east of Irkutsk), 15.VII.2003. Signals of 4 ## are recorded at Echemes in Ae. variegatus minutus are about twice as the temperature 27 and 33°C. long as in Ae. variegatus variegatus, moreover, the ranges of DISTRIBUTION. Steppes of Irkutsk Area. variability of this parameter do not overlap. Also, in the REFERENCES TO SONG. Unknown. echemes of Ae. variegatus minutus wave-shaped changes of SONG AND ACOUSTIC BEHAVIOUR. In general struc- amplitude usually (but not always!) present, whereas for ture the calling song is similar with this of nominotypical signals of Ae. variegatus variegatus as a rule rather monoto- subspecies, but differs from it in longer echemes averaging nous increasing of amplitude along echeme is intrinsic. On 3.6–6.5 s (mean value 4.7±0.15 s) (Figs 144–152). Usually, the other hand, both temporal pattern and repetition rate of wave-like changes of amplitude present in the echemes (Figs syllables in two subspecies are almost identical. Thus, these 146, 149–152), but this character can vary even in the song of two forms differ distinctly from each other only in echeme 40 D.Yu. Tishechkin & M.A. Bukhvalova

129 Saratov Area, Ozinki (3) 2 s 134 130 Saratov Area, Ozinki (3) 135 2 s 131 Saratov Area, 2 s Ozinki (3) 136 132 Chita Area, Onon Riv. (11) 2 s 137 133 2 s Chita Area, Onon Riv. (11) 138 134 Saratov Area, Ozinki (3) 200 ms 139 135 Saratov Area, Ozinki (3) 140 200 ms 136 Saratov Area, Ozinki (3) 141 200 ms 137 Chita Area, Onon Riv. (11) 142 200 ms 138

Chita Area, Onon Riv. (11) 143 200 ms 139

Saratov Area, Ozinki (3) 100 ms 140

Saratov Area, Ozinki (3) 100 ms 141 Saratov Area, Ozinki (3) 100 ms 142 Chita Area, Onon Riv. (11) 100 ms 143 Chita Area, Onon Riv. (11) 100 ms

Figs 129–143. Oscillograms of calling signals of Aeropedellus variegatus variegatus (Fisher-Waldheim, 1846) from different localities (No. of the locality on the map is given in brackets). Faster oscillograms of the parts of songs indicated as “134–143” are given under the same numbers. 129, 134 and 139 — male No.1, 130, 135 and 140– male No.2, 131, 136 and 141 — male No.3, 132, 137 and 142 — male No.4, 133, 138 and 143 — male No.5. Ðèñ. 129–143. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Aeropedellus variegatus variegatus (Fisher-Waldheim, 1846) èç ðàçíûõ ãåîãðàôè÷åñêèõ òî÷åê (â ñêîáêàõ óêàçàí íîìåð òî÷êè íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “134–143”, ïðåäñòàâ- ëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 129, 134 è 139 — ñàìåö ¹ 1, 130, 135 è 140– ñàìåö ¹ 2, 131, 136 è 141 — ñàìåö ¹ 3, 132, 137 è 142 — ñàìåö ¹ 4, 133, 138 è 143 — ñàìåö ¹ 5. Signals of Gomphocerinae from Siberia and the Russian Far East 41

144 147 6 s 145

148 6 s

146

149 6 s

147

153 2 s

148

154 2 s 149 2 s 155 150 156 2 s 151 2 s 157 152 2 s

153

158 200 ms

154

159 200 ms 155 160 200 ms 156

161 200 ms 157 200 ms 162 Figs 144–157. Oscillograms of calling signals of Aeropedellus variegatus minutus Mistshenko, 1951 from steppes about 30 km north- east of Elantsy, Irkutsk Area (No. 7 on the map). Faster oscillograms of the parts of songs indicated as “147–149” and “153–162” are given under the same numbers. 144–149 and 153–155 — male No.1, 150 and 156– male No.2, 151 and 157 — male No.3, 152 — male No.4. Ðèñ. 144–157. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Aeropedellus variegatus minutus Mistshenko, 1951 èç ñòåïåé îêîëî 30 êì Ñ ïîñåëêà Åëàíöû, Èðêóòñêàÿ îáë. (òî÷êà ¹ 7 íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “147–149” è “153–162”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 144–149 è 153–155 — ñàìåö ¹ 1, 150 è 156 — ñàìåö ¹ 2, 151 è 157 — ñàìåö ¹ 3, 152 — ñàìåö ¹ 4. 42 D.Yu. Tishechkin & M.A. Bukhvalova

158 100 ms 159 100 ms

160 100 ms 161 100 ms

162 100 ms 163 2 s 165 166 164 2 s 167 168 165 200 ms 169 166

170 200 ms

167 171 200 ms 168

172 200 ms 169 100 ms

170 100 ms 171 100 ms 172 100 ms Figs 158–172. 158–162 — oscillograms of the parts of calling signals of Aeropedellus variegatus minutus Mistshenko, 1951 from steppes about 30 km north-east of Elantsy, Irkutsk Area, No. 7 on the map (also, see Figs 144–157). 163–172 — oscillograms of calling signals of Dasyhippus barbipes (Fischer-Waldheim, 1846) from the environs of Onokhoy, Buryatia, No. 9 on the map. Faster oscillograms of the parts of songs indicated as “165–172” are given under the same numbers. Ðèñ. 158–172. 158–162 — îñöèëëîãðàììû ôðàãìåíòîâ ïðèçûâíûõ ñèãíàëîâ Aeropedellus variegatus minutus Mistshenko, 1951 èç ñòåïåé îêîëî 30 êì Ñ ïîñåëêà Åëàíöû, Èðêóòñêàÿ îáë., òî÷êà ¹ 7 íà êàðòå (ñì. òàêæå Ðèñ. 144–157). 163–172 — îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Dasyhippus barbipes (Fischer-Waldheim, 1846) èç îêðåñòíîñòåé Îíîõîÿ, Áóðÿòèÿ, òî÷êà ¹ 9 íà êàðòå. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “165–172”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Signals of Gomphocerinae from Siberia and the Russian Far East 43

175 178 179

173 176

177 174

181

185

182

184 180 183

Figs 173–185. Fore leg (173–174, 176–177, 180, 183–184) and claws and arolium of middle tarsus (175, 178–179, 181–182, 185) of Aeropedellus spp.: 173–175 — Ae. variegatus variegatus from Ozinki, Saratov Area (No. 3 on the map); 176–179 — same from Chita Area (No. 11 on the map); 180–182 — Ae. variegatus minutus Mistshenko, 1951 from Irkutsk Area (No. 7 on the map); 183–185 — Ae. reuteri (Miram, 1907) from Minusinsk. Ðèñ. 172–184. Ïåðåäíÿÿ íîãà (173–174, 176–177, 180, 183–184) è êîãîòêè è ïðèñîñêà ñðåäíèõ ëàïîê (175, 178–179, 181–182, 185) âèäîâ Aeropedellus: 173–175 — Ae. variegatus variegatus èç Îçèíîê Ñàðàòîâñêîé îáë. (òî÷êà ¹ 3 íà êàðòå); 176–179 — òî æå èç ×èòèíñêîé îáë. (òî÷êà ¹ 11 íà êàðòå); 180–182 — Ae. variegatus minutus Mistshenko, 1951 èç Èðêóòñêîé îáë. (òî÷êà ¹ 7 íà êàðòå); 183–185 — Ae. reuteri (Miram, 1907) èç Ìèíóñèíñêà. duration. On the contrary, populations of Ae. variegatus REFERENCES TO SONG. Bukhvalova and Vedenina [1998]: variegatus from Saratov and Chita Areas do not differ signif- recordings from Chita Area. icantly in this character (probability of identity P=0.38 by SONG AND ACOUSTIC BEHAVIOUR. The calling Wilcoxon test), consequently, this is a very constant parame- song is an echeme lasting for about 4.5–7.5 s (Figs 163–164). ter of the signal within a subspecies. This give good reason to Its main part consists of 4–6 complex fragments each includ- believe that Ae. variegatus minutus is a separate subspecies. ing 15–30 short discrete pulses followed by 4–6 syllables Differences of the same level (different duration of echemes, (Figs 165, 167, 169, 171). Pulses repeat at a rate 80–110/s, but identical syllable structure and repetition rate) were also syllable repetition period averages approximately 80–130 described between subspecies of Mongolotettix japonicus (I. ms. The main part lasts for about 2.8–4.5 s in our recordings. Bolivar, 1898) and G. maritimus [Bukhvalova, 1998; Veden- The song ends with a succession of 15–30 syllables of the ina & Bukhvalova, 2001]. same structure as in the main part (Figs 166, 168, 170, 172). Male produces signals with prolonged irregular intervals Dasyhippus barbipes (Fischer-Waldheim, 1846) averaging several minutes. It chooses for singing the places MATERIAL. 9. Buryatia, 10 km east of Onokhoy (about 60 km with bare ground among sparse vegetation. east of Ulan-Ude), the valley of Bryanka Riv., 1.VII.2006. Signals of COMPARATIVE NOTES. Signals of individuals from 1 # are recorded at the temperature 36°C. Buryatia and Chita Area are quite similar both in temporal DISTRIBUTION. Transbaikalia, Mongolia, northern China. pattern and in quantitative parameters. 44 D.Yu. Tishechkin & M.A. Bukhvalova

Chrysochraon dispar major Uvarov, 1925 individual variability. Male produce signals sitting on the MATERIAL. 13. South-west of Khabarovsk Province, about 5 stem among dense vegetation. If not disturbed, it can sing km north of Obluchye Town, 5–6.VII.2002. Signals of 3 ## are unceasingly for several minutes. recorded at the temperature 29–30 and 34–35°C. COMPARATIVE NOTES. Calling songs of Ch. dispar DISTRIBUTION. Eastern part of North Caucasus (Dagestan), dispar (Germar, 1831) from many localities in Western Eu- Georgia, South-eastern Kazakhstan, Kyrgyzstan, Eastern and North- rope, Ukraine and European Russia were described in litera- eastern Uzbekistan, Transbaikalia, Amur Area, south of Khabarovsk ture. Temporal pattern of signals of individuals from differ- Province, Maritime Province, Western China [Mistshenko, 1986]. ent populations are quite similar and do not differ from these REFERENCES TO SONG. Ch. dispar dispar (Germar, 1831): of Ch. dispar major. Temporal parameters of songs within Ragge and Reynolds [1998] (recordings from Western Europe); each subspecies as well as in different subspecies overlap to Vedenina and Bukhvalova [2001] (recordings from Ukraine and European Russia). a large extent (Table). Ch. dispar major: Vedenina and Bukhvalova [2001] (record- ings from southern Kazakhstan; subdivision into subspecies is not given in the paper). Discussion SONG AND ACOUSTIC BEHAVIOUR. The calling song is a sequence of echemes, each lasting from 1.2–1.5 s Presently, a great body of information on acoustic (includes 18–20 syllables; recording at 34–35°C, Figs 187, signals of palaearctic Gomphocerinae exists in litera- 189–190, 192–193) up to 3–3.5 s (includes 23–26 syllables; ture. For the most part of species descriptions of calling recording at 29–30°C, Figs 186, 188, 191). Syllables repeti- songs from different populations were studied. Howev- tion period in these two cases averages 65–85 and 110–130 er, such investigations usually were conducted within ms respectively. Evidently, such differences result not only from different temperatures during recording, but also from the boundaries of Western Europe or European Russia. The only exception is the article by Vedenina and

186 2 s 188 187

189 2 s

188

191 200 ms

189 200 ms 192 190 200 ms 193 191 100 ms

192

100 ms

193

100 ms Figs 186–193. Oscillograms of calling signals of Chrysochraon dispar major Uvarov, 1925 from the environs of Obluchye, Khabarovsk Province (No. 13 on the map). Faster oscillograms of the parts of songs indicated as “188–189” and “191–193” are given under the same numbers. Ðèñ. 186–193. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Chrysochraon dispar major Uvarov, 1925 èç îêðåñòíîñòåé ïîñåëêà Îáëó÷üå, Õàáàðîâñêèé êðàé (òî÷êà ¹ 13 íà êàðòå). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “188–189” è “191–193”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Signals of Gomphocerinae from Siberia and the Russian Far East 45

Table. Temporal parameters of calling songs of two subspecies of Chrysochraon dispar Òàáëèöà. Âðåìåííûå ïàðàìåòðû ïðèçûâíîãî ñèãíàëà äâóõ ïîäâèäîâ Chrysochraon dispar

Bukhvalova [2001], where the oscillograms of songs of This fact gives good reason to believe that in the case widespread species from different localities in Ukraine, under consideration these differences provide a reliable European Russia, Southern Siberia and the Russian Far taxonomic character, and the forms under investigation East are published. In the present paper the results of actually have subspecies status. farther investigations of geographical variability of the As it was shown by Bukhvalova [2006], partitioning songs of Gomphocerinae are provided. of acoustic transmission channels in grasshopper com- As is seen from the oscillograms presented, in Gom- munities is based on differences in syllable repetition phocerinae grasshoppers, temporal pattern of calling period to a great extent. For this reason, the results of signals is one of the most constant taxonomic charac- more detailed investigation of the variability of this ters. Within a subspecies or monotypical species it parameter will be considered in another paper. retains all its parameters over many hundreds or even The preferences of calling site and position are no thousands of kilometres of the range. As a rule, no less constant ethological characters than signal struc- significant differences in the structure of calling songs ture. Data on acoustic behaviour of widespread species between populations studied can be found. The songs of in european populations [Ragge & Reynolds, 1998; O. haemorrhoidalis (Figs 14–25), S. scalaris (Figs 44– Savitsky, 2005; Savitsky & Lekarev, 2007] are in good 57), G. maritimus (Figs 58–74) and Ch. hammarstro- agreement with our observations in Siberia and the emi (Figs 75–102) can be mentioned as examples. Only Russian Far East. Possibly, ethological characters in between courtship signals of S. nigromaculatus from certain cases also can be used for solving of taxonomic Irkutsk Area and european populations certain minor problems in Gomphocerinae. differences were found. Songs of different subspecies sometimes differ clearly ACKNOWLEDGEMENTS. The first author is greatly from each other. In Chorthippus apricarius apricarius indebted to Dr. K.A. Kolesnichenko (Botanical Garden of (Linnaeus, 1758) and Ch. apricarius major (Pylnov, M.V. Lomonosov Moscow State University) for his invalu- able help in expeditions to Southern Siberia and the Russian 1914), Mongolotettix japonicus japonicus (I. Bolivar, Far East. The study was supported by a grant of a State 1898) and M. japonicus vittatus (Uvarov, 1914) and in Program “Development of Scientific Potential of Higher different subspecies of G. maritimus this is the case School” (project “Biological Diversity: Structure, Stability, [Bukhvalova, 1998; Vedenina & Bukhvalova, 2001]. Evolution”) and Russian Foundation for Basic Research (# On the other hand, no differences in song pattern were 07–04–00349–a). found between Chorthippus macrocerus purpuratus (Vorontsovsky, 1928) and Ch. macrocerus ponticus References (Mistshenko, 1951) [Vedenina & Bukhvalova, 2001]. Similar situation takes place in two subspecies of Chryso- Bey-Bienko G. Ya. & Mistshenko L.L. 1951. [Grasshoppers of chraon dispar. Signals of Ch. dispar major do not differ the fauna of USSR and adjacent countries. Part II.] // from these of nominotypical form neither in general Opredeliteli po faune SSSR, izdavaemye Zoologicheskim pattern (Figs 186–193) nor in quantitative parameters Institutom Akademii Nauk. T.40. Moscow-Leningrad: (Table). Izdatel’stvo Akademii Nauk SSSR. P.381–667 [in Russian]. Comparative analysis of songs from different geo- Benediktov A.A. 2005. [Fauna and acoustic signals of grasshoppers of the genus Chorthippus Fieb. (Orthoptera, Acrididae) from graphical points provides a way of estimating the range Southern Siberia] // Trudy Russkogo Entomologicheskogo Ob- of intraspecific variability of signal pattern and, conse- shchestva. Vol.76. P.118–130 [in Russian, with English sum- quently, of correct interpretation of differences ob- mary]. served. Berezhkov R.P. 1956. [Grasshoppers of Western Siberia]. Tomsk: Izdatel’stvo Tomskogo Universiteta. 175 pp. [in Russian]. Thus, differences between signals of Ae. variegatus Boulard M. 2006. Acoustic signals, diversity and behaviour of variegatus and Ae. variegatus minutus are not very cicadas (Cicadidae, Hemiptera) // Insect Sounds and Commu- great (Figs 129–143 and 144–162). They are most nication. Physiology, Behaviour, Ecology and Evolution. S. stable, however, because signals of the former subspe- Drosopoulos & M.F. Claridge (eds.). Boca Raton, London, cies from two localities situated about 4500 km apart New York: CRC Press, Taylor and Francis Group. P.331– 349. from each other are indistinguishable (Figs 129–131, Bukhvalova M.A. 1993b. [Comparative analysis of acoustic signals 134–136, 139–141 and 132–133, 137–138, 142–143). of Arcyptera fusca fusca (Pall.) and A. fusca albogeniculata 46 D.Yu. Tishechkin & M.A. Bukhvalova

Ikonn. (Orthoptera, Acrididae)] // Vestnik Moskovskogo Savitsky V.Yu. 2000. [Acoustic signals, ecological features and Universiteta. Ser.16. Biologiya. No.1. P.46–49 [in Russian, reproductive isolation of grasshoppers of the genus Docios- with English summary]. taurus (Orthoptera, Acrididae) in semidesert] // Zoolog- Bukhvalova M.A. 1993b. [Acoustic signals and morphological icheskiy Zhurnal. Vol.79. No.10. P.1168–1184 [in Russian, characters of certain species of grasshoppers of the genus with English summary]. Chorthippus from the Ch. biguttulus group (Orthoptera, Acri- Savitsky V.Yu. 2002. [Acoustic communication, distribution pattern, didae) of Russia and adjacent territories] // Zoologicheskiy and ecology of grasshoppers of the genus Ramburiella (Ortho- Zhurnal. Vol.72. No.5. P.55–65 [in Russian, with English ptera, Acrididae) in Russia and Transcaucasia and some problems summary]. of taxonomy of the tribe Arcypterini] // Zoologicheskiy Zhurnal. Bukhvalova M.A. 1998. [New data on taxonomy of Chorthippus Vol.81. No.1. P.13–28 [in Russian, with English summary]. biguttulus group (Orthoptera, Acrididae) from Russia and adja- Savitsky V.Yu. 2005. [New data on acoustic communication of cent countries] // Zoologicheskiy Zhurnal. Vol.77. No.10. grasshoppers of the genera Omocestus Bol. and Myrmeleotettix P.1128–1136 [in Russian, with English summary]. Bol. (Orthoptera, Acrididae) from Southern European Russia Bukhvalova M.A. 2006. Partitioning of acoustic transmission chan- and their taxonomic importance] // Trudy Russkogo Entomolog- nels in grasshopper communities // Insect Sounds and Commu- icheskogo Obshchestva. Vol.76. P.92–117 [in Russian, with nication. Physiology, Behaviour, Ecology and Evolution. Eds.: English summary]. S. Drosopoulos, M.F. Claridge. London, New York: CRC Savitsky V.Yu. 2007. [New data on acoustic communication and Press, Boca Raton: Taylor and Francis Group. P.199–205. ecology of grasshoppers of the genera Eremippus and Docios- Bukhvalova M.A. & Zhantiev R.D. 1994. Acoustic signals in taurus (Orthoptera: Acrididae) and notes on the use of bioacous- grasshopper communities (Orthoptera, Acrididae, Gomphocer- tic data in supraspecific taxonomy of the subfamily Gomphocer- inae) // Entomological Review. Vol.73. No.2. P.121–136. inae] // Zoologicheskiy Zhurnal. Vol.86. No.7. P.813–830 [in Bukhvalova M.A. & Vedenina V.Yu. 1998. Contributions to the Russian, with English summary]. study of acoustic signals of grasshoppers (Orthoptera: Acrid- Savitsky V.Yu. & Lekarev A.Yu. 2007. [New data on acoustic idae: Gomphocerinae) from Russia and adjacent countries. I. communication and sexual behaviour of grasshoppers (Ortho- New recordings of the calling songs of grasshoppers from Russia ptera: Acrididae) from semi-deserts and deserts of Russia and and adjacent countries // Russian Entomol. J. Vol.7. No.3–4. adjacent countries] // Russian Entomol. J. 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