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Russian Entomol. J. 16(3): 265-272

ї RUSSIAN EN TOMOLOGICAL J OUR NAL, 20 07

Similar calling signals in different species of leafhoppers (Homoptera: Cicadellidae): an example of Paralimnini Сходные призывные сигналы у разных видов цикадок (Homoptera: Cicadellidae): исследование на примере Paralimnini D.Yu. Tishechkin Д.Ю. Тишечкин
Depar tment of Entomology, Faculty of Biology, M.V. Lomonosov Moscow Sta te Univer sity, Vor obyevy Gory, Moscow 1 199 92, Ru ssia. Ema il: macr opsis@yandex.r u. , , . .. , , , 119 992 .

KEY WORDS: leafhoppers, taxonomy, dia gnostic characters, vibr ator y signals, Para limni ni, Cicadellidae. : , , , , Paralimnini, Cicadellidae. ABSTRACT . Examples of simi larit y of call ing signal s in different species of Pa rali mnini (Hom optera: Cicadell idae) are discussed. Species producing sign als wit h identical st ruct ure n ever come int o acoustic int eract ions with each other due to differ ences in h ost specia lisa tion, habi tat preferences or geograph ical dist ribut ion. Ther efore, usi ng of bioacoustic chara cters in systema tics with out considerin g the dat a on biol ogy and distribution can lead to wr ong taxon omic conclusions, because the simi larit y or even absolute ident ity of structure of signa ls in differen t forms i s not always unequivocal evidence of their synonymy. . Paralimnini (Homoptera: Cicadellida e). , , , . , , . It i s wel l known t ha t t hese a re di fferen ces i n t empor al pat tern of ca lli ng si gn a ls, whi ch provi de r eproduct ive i sol at ion i n ma ny speci es of i n sects using a coust ic com muni cati on . For th is rea son, an a lysi s of sign al s i s successfull y em pl oyed i n t axonomy for discri mi n ati on between cr ypt ic speci es an d for est abl ish in g of sta tus of cl osel y r elat ed form s showing smal l mor ph ological di fferen ces. Sim ila rit y of si gnal patt er ns in di fferen t for ms does n ot al wa ys i ndica tes t ha t t hey bel ong to t h e sam e species, however. For exam ple, i n cer t ai n gra ssh oppers, e.g. , in Eur opea n E uc horthi ppus a nd in a num ber of Omoce stus species (Or t hopter a: Acri dida e) t emporal pa tt ern of call in g sign al s i s quit e sim ila r a nd somet im es is alm ost i den ti cal [Ragge & Reynol ds, 1984; Ragge, 1986]. Recent in vesti gati on of ca ll in g songs i n gra ssh opper comm unit ies dem onst rat ed t h at th e com plex of t em por al pa ram et ers of si gn al det er mi n es for each species i ts own pla ce in th e a cousti c en vir onm en t of t he comm unit y, so-ca ll ed a coust ic ni che, wh ich i s a par t of t he ecol ogi cal n i ch e a s a wh ol e [Bukhval ova, 2006]. Si gn a ls of species belongi ng to t h e sam e comm un i ty di ffer fr om each oth er, i . e. occupy differ ent a cousti c ni ch es, whereas i n m em bers of different comm un i ti es th ey som eti mes a re al most i ndist in gui sh a ble i n t em por al pat tern . T h is i s n o ba rr ier t o successful comm un ica ti on because such species n ever com e i nto a coust ic in ter act ions wit h ea ch oth er. Simi la r sit uat ion ta kes pla ce i n Psylli nea (Hom opt er a). T emporal patt ern of call in g songs i n t his group for t h e m ost par t i s species-specifi c. Nonet hel ess, in a num ber of species sign al s a re quit e si mi lar a nd ca nn ot be tol d a pa rt wit h cer tai nt y [Ti shech kin, 2006]. Occa sion all y, such speci es ar e forma ll y sympat r ic, i .e. l ive i n t h e sam e biotope. Sti ll , i t shoul d be r em ember ed, th at psylli ds use n ot ai rborn e soun ds, but subst rat e-born e vi brat ion s for t h ei r comm un i cat ion. Such si gnal s ca n not be t ra n sm itt ed fr om one pl ant to an oth er wit hout physical con tact. Con sequen tl y, form s wi th na rr ow h ost special iz ati on dwell in g on different pla nt species as a rule can not hea r th e si gn als of ea ch oth er. Sma ll Auchenor rh yncha (Hom opt era : Cica di nea wit h t he excepti on of si n gi ng ci cadas, Ci ca di dae) a lso


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D.Yu. Ti shech kin used. An a lysis of r ecor di n gs wa s per for m ed on PC provided wit h an a log/digi tal con ver t er a n d a ppropria te softwa re. Data for recor di n gs of si gn a ls of leafh opper s used i n t he pa per a re gi ven i n t h e Ta ble. It is appar en t , t ha t i n a ll opa tr i c speci es di ffer en ces i n st r uct ur e of com mun ica ti on si gna ls pla ys n o r ole in r epr oduct ive i sol ati on . Con sequen tl y, such for ms can produce simi la r si gn al s a n d sti ll exi st as sepa r ate speci es. In certa in Pa ral im ni n i t ha t i s t he ca se. Sorhoanus medi us (Mulsant et Rey, 1855) and S. hilaris (Melich ar, 1900) are common and abundant steppe-dwell ing species feeding on grasses (Poacea e). They in habit sim ilar biotopes, but the range of the for mer one i ncludes E urope, Ka zakhstan and south ern par t of West ern Siberi a, wh erea s the latt er occurs in steppes of Eastern Siberia (Irkutsk Area, Tran sbaikalia, Yakutia) and in Centra l and Eastern Mongolia. In both species, call ing signal s con sist of single or r egula rly repeatin g short fragments with variable shape (Figs 1- 10 and 11-16). Occasion ally, fra gment s of quite different shape present in t he si gnals of speci mens from the sam e populat ion (Fig. 1). Inner structur e of signa ls a lso vari es grea tly. Si gnal consists either of sine waves (Figs

produce substr a te-born e vi bra tory si gn al s. Alt hough a great num ber of r epresen t at ives of ma n y order s of i nsect s possess well -developed acousti c comm un i cat ion , si gnal s of Ci cadi nea sta n ds out beca use of m ost ela bor at e t empora l str uct ur e. E ven i n closel y r ela ted speci es t he pa tt ern of ca ll in g songs som eti mes i s complet el y differ en t . T hus, in ma n y t axa of Cicadi nea a coust ic an a lysis pr ovi des useful ch ar a ct ers for discri mi n ati on between cr ypt ic speci es [for exam ple, T ish echki n, 1999a , b, 2002]. By con t ra st , i n cer t ain tribes of leafhoppers (Cicadellidae) and in some Fulgoroidea th e st ructur e of si gnal s is rat her si mple and uni for m. Di fferen ces bet ween sign al s of closel y r el ated speci es i n such groups ar e i ndi sti nct or a bsent. T herefor e, si gnal a nal ysi s for ta xonom ic purposes m ust be used wit h ca ut ion i n t hi s si tuati on. Cer tai n exam ples of sim il ar it y of cal li n g si gn a ls in the representatives of the tribe Paralimnini (Homoptera: Cicadel li da e: Del toceph al in ae) wi ll be di scussed here. Recordin gs of vi brat or y sign al s wer e m ade by m ea n s of pi ezo-el ect ri c crysta l gram ophon e car tr i dge conn ected t o t h e mi crophone in put of ca ssett e r ecorder "El ekt ron ika -302-1" or mi n idisk r ecor der Sony Wal kma n MZ-NH900 via t he mat ch i ng am pli fi er . In a ll ca ses ma nual mode of recor di n g l evel con tr ol was

Da ta for rec ordings of callin g signa ls o f the stud ied spe cie s of Pa ralimnini Да нные о запи сях пр изывных сигнал ов и зуч енны х видо в Pa ralimnini


Sim ilar sign als in di fferent species of leafhoppers

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1 2 2 3 4 5 5 6 7 8 9 9
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3

4

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Figs 1-1 6. Osc illogr ams of ca lling sig nals o f m ale s: 1-1 0 -- So rho anus medius; 1-5 -- fro m S ara tov Are a; 6-9 -- fro m M osc ow Ar ea; 10 -- fro m Altai; 1 1-1 6 -- S. hilaris; 11-15 -- from Burya tia; 16 -- fro m Chit a Area . P art s o f signals ind ica ted as 2-5, 9 a nd 12 ar e g ive n o n o scillo gra ms und er the sa me num ber s. Рис. 1-1 6. Осцилл огр аммы призывных с игнало в с амцов: 1-10 -- S orh oan us medius; 1-5 -- из Сар ато вск ой обл .; 6-9 -- из Мо ско вск ой обл .; 10 -- с Ал тая ; 1 1-1 6 -- S. hilaris; 11-15 -- из Буря тии; 1 6 -- из Читинско й о бл. Фр агмент ы с игнало в, об означе нные цифр ами 2-5, 9 и 12 , пред ста вле ны на осцилл огр аммах под та кими ж е номе рами.


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D.Yu. Ti shech kin

17 18 18 19 20 21 22 23 24 25 26 26 27 28 29 30 31 31 32
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20

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200 ms 200 ms 200 ms 200 ms 200 ms 200 ms 200 ms 2s

29

2s 200 ms

32

33

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Figs 17-33. Osc illogr ams of ca lling sig nals: 1 7-2 1 -- Mo cuellu s c ollinus; 22-2 4 -- M. an gustiarum; 25-2 7 -- Psamm otettix atrop idicola; 28-3 3 -- P. kaszabi. Par ts of sig nals indicat ed as 18, 20, 2 6, 29 and 31 -33 ar e g ive n o n o scillo gra ms und er the sa me num be rs. Рис. 17-33. Осцилл огр аммы приз ывных сигнало в: 17-21 --Mo cuellu s c ollinus; 22-2 4 -- M. an gustiarum; 25-2 7 -- Psamm otettix atrop idicola; 28-3 3 -- P. kaszabi. Фра гме нты сигна лов , о боз нач енные цифрами 1 8, 20, 26 , 2 9 и 31 -33 , пред ста вле ны на ос цил лограммах по д та кими ж е номе рами.


Sim ilar sign als in di fferent species of leafhoppers 4-5), or of vibra tions with more complex and l ess regular shape (Figs 8-9). Evidently, variability of waves sha pe is a r esult of different transmission proper ties of the part s of twigs or stems and of differ ences in relat ive position of singi ng in sect and vibrot ransducer detecting vibrational signa l. So, as is seen from the oscil logra ms on Figs 1-16, n o clear-cut di stin ction exi sts bet ween calli ng si gnals of these two speci es. Mocuellus collinus (Boheman, 1850) and M. angustiarum Ti shetshkin, 1994 is an other pai r of species indistin guishable in struct ure of acousti c si gnals. M. col linus is a tr anspa laea rctic species dwelli ng on gra sses in va rious open habitats. M. angustiarum wa s found only in highly peculi ar mountain st eppes in arid depression bet ween the ridges of North Ossetia (Nort h Caucasus) an d, apparen tly, is endem ic of thi s region. Sign als of both species a re syllabl es, i.e. succession s of more or less distinct pul ses with t otal durat ion approximat ely from 0.8 up to 1. 5 s (Figs 17-21 and 22-24). They are sim ilar both in temporal pattern an d in dura tion of ser ies. On a verage, syllables of M. angustiarum a re som ewhat shor ter; still , signals of t wo species over lap alm ost completely in th is pa ramet er. Call in g sign al s of t he m ost pa r t of th e studied species of Psammot ett ix ha ve th e sam e sch eme of t em por al pa tt ern . T hey consist of shor t syl la bles foll owi ng ea ch ot h er wi th more or less regula r i nt er vals [Ti shech kin , 1999c, 2000]. Signal s of di ffer en t species differ fr om each oth er m ostl y i n t h e l en gt h of syll ables. However, in certa in ca ses t hey over la p in t hi s cha r acter a s well . Psammot et tix k aszabi Dl abol a, 1961 a nd P. at ropidic ola Em elja nov, 1962 ar e an exam pl e. Syl la bles r epetit ion per i od in t h ei r si gn al s is r at her va ri abl e a nd h a s sim ila r val ues i n bot h species (Fi gs 25 a nd 28, 30) as wel l a s th e dur at ion of syll ables (Fi gs 26-27, 29, 48). Cer tai n differ en ces i n t h e st r uct ur e of pul ses is of n o con cern , because t he sh ape of waves i n substr at e-t ra nsmi tt ed vibrat ions depen ds t o a l ar ge ext en t on t he ph ysi cal pr oper t ies of a substr a te, a s i t wa s m ent ion ed above. As a r esult , si gn al s of i nsect s sin gin g i n differ en t poin t s on th e sa me pl ant a re somewhat differ ent (Figs 30-33). Bot h species dwel l on sal in e la nds i n sout hwest ern par t of E ur opea n Russia a n d i n Kaz akh sta n a nd often can be foun d i n t he sa me ha bit at , e. g. , i n t he Lower Vol ga r egion. On t he oth er h a nd, th ei r host specia li sat ion i s quit e differ en t . P. kaszabi feeds on sa lt wormwoods (A rte mi sia subg. Seri phidi um), wherea s P. atropidi cola dwell s on Puc ci ne ll ia spp. (Poa ceae), th us, t hey a voi d acoust ic i n ter feren ce. Simi la r sit uat ion i s observed i n th e species of Dipl oc ole nus s. str . I fa il ed t o fi n d a ny di fferen ces bet ween t he si gn a ls of t h ree species st udied (D. bohemani (Zett erstedt, 1840), D. frauenfe ldi (Fi eber, 1869) a nd D. sutt hol li Vi lbast e, 1980) basin g on a vai la ble m at eri al . As i n t he ca ses di scussed above, t he sh ape of waves in a syll able (Fi gs 35-36 a n d 37-38, si gn al s of

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t he sa me mal e ar e pr esen t ed on oscil logr a ms) a s well a s t he outli ne of sylla bl es an d t h ei r dur a ti on (Fi gs 34- 35, 37 a n d 39-46) pr ovi de no relia bl e di agnostic cha ra ct er s. Al l speci es a re pa rt ly sympat ri c. Th e ra nges of D. bohemani an d D. f raue nfel di in cl ude E urope, Kaza kh sta n a nd South ern Si ber ia (possibl y, wi th the except ion of E ast er n T ra n sbai kali a). D. sutt hol li inhabi ts steppes of Mongolia and Sout hern Si beria (T yva, Bur yat ia ). Th ese speci es were n ever foun d in th e same biot ope, however. For i nst an ce, i n Buryati a D. sutt hol li wa s coll ected on ly on t h e mea dows wit h steppe vegeta ti on on t he ri ver ba nks, wherea s D. fraue nfe ldi dwell ed on th e gl ades in dr y pin e for est on t h e slopes of t he h i ll s surr oun di ng th e vall ey (T abl e). T her efor e, t he sh ape of waves i n a sign al , a s well a s t he outli ne of syll abl e (sl ope of t he lea di ng and t ra ili ng edges, etc.) ca n var y gr ea tl y even wi th i n the sam e recordin g, especia ll y i f th e si ngin g in sect cha nges it s posi ti on from t im e t o t im e (Fi gs 34-38). Con sequentl y, di fferen ces of t hi s kin d r eveal ed bet ween two or thr ee selected osci ll ogr am s do n ot gi ve gr oun ds for a ny ta xon omi c con clusi ons. Dur a ti on of syl la bles i s a lso r at h er va r ia ble. T his par a meter ca n var y by a fa ct or of about 1. 3-1.5 and m or e a t con sta n t t em per at ure (Figs 47-48). In a ddit ion , it sh oul d be t aken i nt o a ccoun t t hat un der n at ural con dit ion s t em per at ure va ri es from pla ce t o pla ce, even on a very sm all scal e. As it wa s dem onst ra ted by de Vr i jer [1984], fema les of pla n th opper s r espond r em ar kabl y wel l t o pla yba ck of ma le ca lli ng si gn a ls at t em per at ures differ i ng by 5њC from t h e mal e call ing t em per at ure. T hus, i t is bel ieved t h at compa ri son of sign al s r ecorded at tem perat ur es differ in g wi th in the r an ge of 5њC is qui te corr ect. Alt h ough th e r an ges of va ria bil it y of syl la bles dur a ti on in di fferen t speci es over la p con sidera bl y or even com plet el y, sta ti sti cal di fferences bet ween samples exi st alm ost a lwa ys. In t he pr esent st udy Wi lcoxon t est was used for esti mat ion of probabili ty of i dent it y of t he sam pl es pr esen ted on h ist ogr am s on Figs 47-48. It was found t h at al l sam ples i ncl udin g r ecordi ngs of th e sam e speci es fr om di fferent local it ies differ sign ifica ntl y fr om each oth er. Consequen t ly, i n t his case i t i s i mpossibl e t o di scr i mi n at e bet ween species and intraspecific forms using statistical methods. Moreover , as a r ul e r ecepti ve femal e sta r ts si ngi ng in r eply t o cal li n g m al e a lm ost im mediat el y. So, even if it is gra n ted t hat femal e possesses i n t he ner vous system some mechan ism ca pable of sta tistic an alysis, i t should h ea r con sider a bl e n um ber of si gn a ls t o obta in a sufficient sam pl e pri or to pr oduci n g r eply. In certa in ca ses species pr oduci ng si milar sign als differ disti nctly from each oth er in the shape of geni talia . The studied speci es of Moc uell us (Figs 49-50 and 51-52) an d Psammot ettix [E melya nov, 1964] can be men tioned as an example. D. bohemani al so differs from all con gener ic species in these char acters. Morphological di fferences between t wo ot her species of the gen us an d also, between Sorhoanus medi us an d S.


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D.Yu. Ti shech kin sol ute i denti ty of str uctur e of calli ng signal s is not always unequivoca l evidence of synonymy of th e forms under investigat ion.
AC KNOWLEDGEME NTS. The study was supported by a grant of a State Program "Development of Scientific Potential of Higher School" (project "Biological Diversity: Structure, Stability, Evolution") and R ussian Foun dation for Basic Research (No 07-0 4-0 039a).

hilaris ar e not so clear (Figs 53-56 an d 57-60). In such cases th e use of acoustic a nalysis wi thout con sider ing the data on biology an d distribution can lead to wr ong taxonomic conclusions. Species not involved into acoustic interact ions in na ture due to di fferences in geogra phica l di stribution , host special isat ion or ecologi cal preference can produce signals indi stin guish able in tem poral patt ern. Therefore, the simi larit y or even ab-

34 35 35 36 36
20 ms 200 ms 2s

37 38 39
200 ms 20 ms 200 ms

40 41 42 43 44 44 45 46 41

2s

200 ms 200 ms 2s

200 ms 200 ms 200 ms

Figs 34-46. Oscillograms of calling signals of males: 34-39 -- Diplocolenus frauenfeldi; 34-38 -- from Saratov Area; 39 -- fr om Burya tia ; 4 0-4 2 -- D. su tth olli; 43-4 6 -- D. bo hemani, 43-44 -- from Altai, 4 5-4 6 -- fr om Mosc ow Are a. Par ts of sig nals indicat ed as 35 -36 , 3 8, 41 and 44 ar e g ive n o n o scillo gra ms und er the sa me num ber s. Рис. 34-46. Ос цил лограммы призыв ных сигна лов са мцо в: 34-39 --Diplo colenu s frau enfeldi; 34-38 -- из Сар ато вск ой обл ., 39 -- из Бур ятии; 4 0-4 2 -- D. su tth olli; 43-4 6 -- D. bo hemani; 43-44 -- с Ал тая , 4 5-4 6 -- из Мос ков ско й о бл. Фр агмент ы с игнало в, об означе нные цифр ами 35 -36 , 3 8, 41 и 4 4 пред ста вле ны на осцилл огр аммах под та кими ж е номе рами.


Sim ilar sign als in di fferent species of leafhoppers

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47
Figs atrop idic Рис. atrop idic

48
47-48. Syllab les durat ion in ca lling signals: 4 7 -- So rho anus medius a nd S. hilaris; 48 -- Psamm otettix kaszabi and P. o la. Ea ch point on histog ram s d eno tes durat ion of one syllable. 47 -48. Дл ите льност ь с ерий в пр изывных с игнала х: 47 --So rho anus medius и S. hilaris; 48 -- Psamm otettix kaszabi и P. ola. Кажд ая точ ка на гис тограммах со отве тст вуе т д лит ельнос ти одной сер ии

49

50

53

54

57

58

51

52

55

56

59

60

Figs 49-60. Penis: 49-50 -- Mocuellus collinus; 51-52 -- M. angustiarum; 53-54 -- Sorhoanus medius; 55-56 -- S. hilaris; 57-58 -- Diplocolenus frauenfeldi; 59-60 -- D. suttholli; 49, 51, 53, 55, 57, 59 -- lateral view; 50, 52, 54, 56, 58, 60 -- back view. Рис. 49-60. Пе нис : 4 9-5 0 -- Mo cuellu s c ollinus; 51-5 2 -- M. an gustiarum; 53-5 4 -- So rho anus medius; 55-5 6 -- S. hilaris; 5 7- 58 -- Diplo colenu s frau enfeldi; 59-6 0 -- D. su tth olli; 49, 51 , 5 3, 55, 57 , 5 9 -- сб оку; 50, 5 2, 54, 56 , 5 8, 60 -- с зад и.


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sinae ) from the t err ito ry of Russia with not es on other Pa lae arc tic sp ecies of the ge nus // Ib id. Vo l.8 . N o.4 . P .23 9- 252. Tishechkin D.Yu. 1999c. [Variability of acoustic signals and some morphological characters in Psammotettix striatus (Homoptera, Cicadellidae) from Russia and adjacent countries] // Zoologicheskiy Zhurnal. Vol.78. No.11. P.1298-1305 [in Russian with English summary]. English translation: Entomological Review. 1999. Vol.79. No.9. P.1263-1270. Tishec hkin D. Yu. 2 000. Vib ra tio na l c omm unicat io n in Ap hr od inae lea fhopp e r s ( De lt o c ep ha linae a uct. , H omop t e ra: Cica d e llid ae ) a nd r e late d g r oup s wit h no t e s o n c la ssificat io n o f hig he r ta x a // Russia n E nt omo lo g ica l J . Vo l. 9 . No . 1. P . 1-66. Tishe chkin D.Yu. 2 002 . Review of the sp ecie s o f t he genus Ma cro psis L ewis, 183 4 (Hom opt era , Cic ade llidae , M acr opsinae) fro m E uro pea n Russia a nd adjace nt ter rit ories // Russian Ent omo log ica l J . Vol. 11. No .2. P. 123 -18 4. Tishe chkin D.Yu. 2 006 . Vibr ato ry com munica tion in Psylloid ea (H omo pte ra) // Insec ts sounds and c omm unicat ion. P hysiology , b eha vio ur, ec olo gy and ev olutio n. Eds.: S. Dro sop oulos & M.F. Clar idg e. CRC Pr ess, Taylor and Fr anc is Gro up, Bo ca Ra ton, L ond on, New Yo rk. P. 357 -36 3. Vrije r P.W. F. de. 19 84. Va ria bility in ca lling sig nals o f t he plant hop per Ja vesella pellu cida ( F.) (H omo pte ra , D elp hac idae) in relation to tem per ature, and c onsequenc es for sp ecies re cog nit ion during d ist ant co mmunic ation // Net her lands J. Zo ol. Vo l.3 4. No .3. P. 388 -406.

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