Bulletin of the British Museum (Natural History) The songs of the western European grasshoppers of the genus Stenobothrus in relation to their taxonomy (Orthoptera: Acrididae) D. R. Ragge Entomology series Vol55 No2 24 September 1987 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) © British Museum (Natural History), 1987 The Entomology series is produced under the editorship of the Keeper of Entomology: Laurence A. Mound Publications Manager (Entomology): W. Gerald Tremewan ISBN 0 565 06032 5 ISSN 0524-6431 Entomology series Vol 55 No 2 pp 393-424 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 24 September 1987 The songs of the western European grasshoppers of the genus Stenobothrus in relation to their taxonomy (Orthoptera: Acrididae) D. R. Ragge Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD Contents PERI N FY SUR oe iyi so ets ctrrache senting eke Aaa MAA sate Re Mahe wiynwsaudvleavenivedinndndavanenniisan 393 BEYETEMCAUCILORN Soca Sane coterie tear adeaaen eacie.s diee en aaah Anions onis cl calsidorawseas'a cabin in aninyscanied v/ansia ter 393 PA CKMOW I COPEMEM (Stems onrme tate me ee ntee iors etter entre cea tania cs <rdcsertesmeseaessnaiesi 394 PX DOLE VIA TONS OL CC POSIOGIES caarersiicn: states roe eosseber sess sedeasecdeesatervosesadedonsetescese se 394 SARI RERIGR A testa er, 2o J nccPe Nc gute est ace sce ever sT AREY « cunadee Jan un vende due sin stlia en aabuavad deawsvati« 395 RECOLGIN gan dlanalySiie tHe SOMPSs oat ns-ceeee see het adaceewasns apkieua= Wa nateccnoanenctestenes 395 SON CMETOMMOlO CViternne ee Nen cere nce actetac te tease ane eca re et ne eetnanttinsione eeting awake eansehey 397 Rea Ms ANON roe crn's duty tein as seh ecideicla at nF <-ig ss AN deaets Sniaithas Zara cello «Shak Mh dade a disawadiesdesiea 397 SUDESON rasa TAN ONOMIC CHALACTEN 5 ss. cevac aoadatine det aeraedes eye «clewe atsaleatendiatideee cra visiee ston’ 398 BP Lene MENTE CNEL MINS st sac cue oi meen cscie vimev'nnechigesifhanenwesnssantnndnvabamainddnceenedesan 399 RRP RRIENE SR RINE MER Mae esto coh on xn vw 2c Aaidy athitan Sener asad Gam cnmpewadnemeniien'nhinnns 399 Keys to the principal western European species of Stenobothrus ...............0cc0cceeeeeee 400 Key based pronarily ommorpholopical ChatacCters:..........0.c-se.scerssscccavcucaseeravenses 400 PREY ASE OM SOME AR ARL CES rien coca scuaviansccecesternedswosenwasheseseasewaseoaracavennecersanse 402 The songs of the westerm European StenODOMris «6s... cccssseccedseseccenensaesenecsscsecees 404 LNGHSTIRETT TSS Gor aneuioc dene dncoocee RO GusBagE CEA nUCUPRTP aoa cepcoCadnee Te ntaaene CaCC ee EE BHP eT CEE RE anE EEE 422 NCEE CORN tes eis ecg at Wh sspint one eens Maar lesb aOR nae cath Uacasmcwdsiiiadnonvadedvemecoseradsas 424 Synopsis The male calling songs of nine species of the Gomphocerine genus Stenobothrus are described and fully illustrated with oscillograms at several different speeds. The courtship song is also described for the seven species in which such a song is known. The songs of three species (festivus, bolivarii and apenninus) are described for the first time. For each species a list is provided of references to any past published work on the songs, classified according to the kind of information given. Notes on recognition, using both morphology and song, are given for each species, and two identification keys are provided, one based mainly on morphology and the other on song. The value of the song as a taxonomic character is discussed. Introduction Stenobothrus is one of the better known genera of Gomphocerinae, a group that includes most of the common European grasshoppers of open grassland and moorland. They are well known for their characteristic songs and are widely used in bio-acoustic, ecological and cytogenetic research. The subfamily includes about 150 European species, of which over 20 are pests in parts of their range, seven of them being regarded as major ones (Tsyplenkov, 1970). Stenobothrus is a widespread Palaearctic genus, including over 30 species living in habitats ranging from lush grassland to steppe and high mountains. Eleven species have been recorded from western Europe, of which three, /ineatus, nigromaculatus and fischeri, become pests when they occur in sufficient numbers, causing damage to cereal grasses, hayfields and alfalfa (Bei-Bienko & Mishchenko, 1951; Tsyplenkov, 1970). It is now well known that the songs of Gomphocerine grasshoppers are of great taxonomic value. Their role as a mate recognition system makes them particularly valuable in resolving Bull. Br. Mus. nat. Hist. (Ent.) 55 (2): 393-424 Issued 24 September 1987 394 D. R. RAGGE complexes of sibling species and in establishing the status of local populations showing small morphological differences. There has so far been no comprehensive account of these songs, presented group by group and illustrated with oscillograms. The main aim of the present paper, and of similar papers already published (Ragge, 1986) or in preparation, is to provide such an account for the western European species, arranged systematically by group and fully illustrated with oscillograms at several different speeds. No information has previously been published on the songs of three of the nine species included in this study (festivus, bolivarii and apenninus) and no oscillograms have previously been published for a further two (fischeri and grammicus). For the species in which published information on the song is already available, references to the sources are given, classified according to the kind of information published. Acknowledgements I am much indebted to Drs A. Gorochov and A. Kaltenbach, who have kindly lent me type-specimens or other material from their respective institutions. I should also like to thank Mr C. F. Dewhurst and Miss T. M. Howard, who kindly brought me live males of S. stigmaticus from the Isle of Man, and Mr E. C. M. Haes, who was kind enough to give me a live male of S. rubicundulus that he had collected in the Italian Alps. I am particularly grateful to my colleague Mr W. J. Reynolds, who made the studio recordings used in this study, helped me in reviewing the past literature on the songs, tested the identification key based on morphological characters and provided valuable comments at various stages in the preparation of the paper. Finally I wish to thank my wife, who has given me invaluable help in all my field-work. Abbreviations of depositories BMNH _ British Museum (Natural History), London MNCN Museo Nacional de Ciencias Naturales, Madrid NM Naturhistorisches Museum, Vienna ZI Zoological Institute of the Academy of Sciences of the U.S.S.R., Leningrad Methods Recording and analysing the songs All the field recordings of the songs used in this study were made in full sunshine (except recording 184/1 — see p. 405) using a Uher 4000, 4200 or 4200IC tape recorder and AKG D202 microphone; the tape speed was always 19 cm/s. The subjects were approached stealthily so that the microphone could be held (or rested on the ground) about 10 cm from them without causing any disturbance. The studio recordings were made in the BMNH Acoustic Laboratory using a Kudelski Nagra IV tape recorder and Sennheiser MKH405 microphone or (recordings 119 and 121 only) Kudelski Nagra III tape recorder and AKG C61 microphone; the tape speed was 19 or 38 cm/s. In every case a bench lamp was used to provide light and radiant heat. Further data are given in Tables 1 and 2 for the recordings used for the oscillograms reproduced in Figs 9-116. However, the song descriptions given for each species are in most cases based on many more recordings of numerous songs, all of which were analysed oscillo- graphically. The oscillograms reproduced in the figures were chosen as being typical of the species concerned and, in some cases, showing the extent of intraspecific variation. Although I have given in Tables 1 and 2 the ambient air temperatures at the time of recording, it is really the body temperature of the singing insect that is important, and this is more dependent on radiant heat than on the air temperature; as the insects were always receiving radiant heat from the sun or a bench lamp, the conditions under which the recordings were made were more uniform than the different air temperatures might suggest. All the oscillograms were made from recorded songs with a Mingograf 34T ink-jet recorder. SONGS OF THE WESTERN EUROPEAN Stenobothrus 395 Table 1 Data for the field recordings of male songs of Stenobothrus used for oscillograms reproduced in this study. All these recordings were made from different males. Recordists’ names are abbreviated as follows: DRR = D. R. Ragge; WJR = W. J. Reynolds. Date recorded and collected Shade air Recording Species Locality Recordist temperature No. lineatus nigromaculatus fischeri festivus grammicus bolivarii stigmaticus FRANCE: Seine- Maritime, near Gamaches Iva.y: Lazio, Monti Reatini, foot of chair-lift to Monte Terminilletto, 1700 m FRANCE: Pyrénées- Orientales, near Saillagouse, Val d’Eyne FRANCE: Lozére, Causse Méjean, near Crosgarnon Same locality Itay: Abruzzi, Gran Sasso d'Italia, Fossa di Paganica, 1680 m ITta.y: Marches, Monti Sibillini, Forca di Presta, 1540 m FRANCE: Vaucluse, Mont Ventoux, road from Bédoin, 1350 m Same locality FRANCE: Alpes- Maritimes, near Col de Vence, 930 m SPAIN: Madrid, near Guadarrama, Embalse de la Jarosa FRANCE: Vaucluse, Mont Ventoux, near Pavillon de Rolland, 800 m FRANCE: Vaucluse, Mont Ventoux, road from Bédoin, 1100 m SPAIN: Madrid, near Navacerrada, Embalse de Navalmedio, 1300 m FRANCE: Haute-Vienne, near St Yrieix-la-Perche SPAIN: Madrid, near Navacerrada, Valle de Barranca 11. viii. 1983 11.ix.1985 22.1x.1976 31.vii. 1982 31.vii.1982 31. viii. 1985 14.ix.1985 15. viii. 1977 15. viii. 1977 25.vili.1977 19.vi.1984 11.viii.1977 12.vili.1977 26. vii. 1983 30.vili.1978 27.vii. 1983 DRR DRR WJR DRR DRR DRR DRR DRR DRR DRR DRR DRR DRR DRR WJR DRR 24°C 18°C 16°C 16°C 16°C 23°C 18°C 2a 2a°€ 23 © 22 C 2356 Lee 26°C 20°C 22°C 511/2 608/4 184/1 482/2 482/3 603/4 609/10 237/1 237/3 241/2 531/6 233/4 234/1 501/8 268/2 503/6 396 D. R. RAGGE Table 1 — cont. Data recorded Shade air Recording Species Locality and collected Recordist temperature No. apenninus Iraty: Abruzzi, Gran Sasso d’Italia, Fossa di Paganica, 1680 m 31.viii.1985 DRR 24°C 603/2 Same locality 31.vili.1985 DRR 19°@ 603/3 ITvAcy: Lazio, Monti Reatini, foot of chair-lift to Monte Terminilletto, 1700 m 11.ix.1985 DRR 18°C 608/1 rubicundulus FRANCE: Alpes- Maritimes, near St Dalmas de Tende, La Miniére de Vallaure, 1500 m 24. vili.1977 DRR 23°C 240/4 ITALY: Trentino-Alto Adige, near Riva del Garda, Tremalzo, 1200 m 27. viii. 1985 DRR 13°C 601/5 Table2 Data for the studio recordings of male songs of Stenobothrus used for oscillograms reproduced in this study. All these recordings were made from different males except for 369 and 371, which were made on different days from the same male of rubicundulus. The recordist was W. J. Reynolds in every case. Date Date Air Recording Species Locality collected recorded temperature No. lineatus ENGLAND: Kent, near Wrotham, Trottiscliffe 10. viii. 1978 11.vili.1978 PEAS 260 Same locality 12.viii.1978 13.vili.1978 29°C 321/1 Same locality 12.viii.1978 13.vili.1978 28°C 322/2 nigromaculatus FRANCE: Pyrénées- Orientales, near Saillagouse, Val d’Eyne 22.ix.1976 29.ix.1976 Zi-€ 176 festivus SPAIN: Madrid, near Guadarrama, Embalse de la Jarosa 19. vi.1984 4.vii.1984 — S27 stigmaticus ISLE OF MAN: Langness Peninsula 1-6.viii.1974 8.viti.1974 24°C 119 Same locality 1-6.vili.1974 = 9. vii. 1974 24°C 121 Same locality 30. viii. 1983 6.i1x.1983 23°C 497 rubicundulus ITALY: Belluno, Pordoi Pass, below cable line to Pordoi Joch 4.1x.1980 16.ix.1980 26.6 369 Same locality 4.ix.1980 18.1x.1980 25°C 371 SONGS OF THE WESTERN EUROPEAN Stfenobothrus 397 Song terminology The bio-acoustic terms used are defined as follows. Calling song. The song produced by an isolated male. Courtship song. The special song produced by a male when close to a female. Syllable. The sound produced by one complete up and down movement of the hind legs (Fig. 1). Echeme. A first-order assemblage of syllables (Fig. 1). Echeme-sequence. A first-order assemblage of echemes (Fig. 1). Momentary breaks in the sound (of at least 1-25 ms) during the course of a syllable are referred to as ‘gaps’ (Fig. 1). Presentation The presentation is identical to that used in my recent paper on Omocestus Bolivar (Ragge, 1986: 215-219). As in that study I have relied mainly on oscillograms in presenting information on the songs and have kept the verbal descriptions as brief as possible. The references to past accounts of the songs are not exhaustive but are intended to include all sources that the reader might find useful to refer to for additional or confirmatory information. The speeds of the oscillograms are (after reduction for printing) 10 mm/s, 80 mm/s, 320 mm/s and 1280 mm/s, although the higher speeds are given only when considered to be useful. For three species (/ineatus, festivus and stigmaticus) I have included oscillograms taken from the calling songs of males with only one hind leg; such oscillograms, at the higher speeds, show the individual peg-impacts with particular clarity (see Figs 30, 61, 93). The song as a taxonomic character In my paper on Omocestus I outlined the history of the use of the songs of Orthoptera in taxonomy (Ragge, 1986: 219-220) and more recently (Ragge, in press) I have emphasized the particular importance of the songs in the taxonomy and biogeography of several groups of western European Orthoptera. The taxonomic value of the male calling songs of the western European species of Stenobothrus can be seen at a glance from Figs 9-17. The song-patterns show a wide variety from the long single echeme of lineatus, lasting more than 10s and with a syllable repetition rate of less than 2/s, to the rapid series of short echemes of grammicus and bolivarii, in which each echeme lasts less than 0-2 s and the syllable repetition rate is more than 50/s. S. rubicundulus stands apart from the other species included in the present study in producing a loud wing-buzz, often during hovering flight, in addition to the more usual femoro-alary stridulation. The wing-buzz clearly forms part of this species’ communicative behaviour and often follows immediately after a femoro-alary echeme (Fig. 17). According to Dr C. Walter, whose observations are quoted by Harz (1975: 777), S. cotticus also produces a wing-buzz; I have not yet been able to study the acoustic behaviour of this species, which is known only from a small area of the French Alps. The calling song of /ineatus is unique among all known Gomphocerine songs in having such long syllables, each usually lasting 0-6—0-8 s. This species also has a uniquely dense stridulatory file, so that, in spite of the slow leg movements, the peg-impact rate (for one hind leg) is still of the order of 500/s in the middle of the downstroke (see Fig. 30). S. /ineatus is also unique among the western European species of Stenobothrus in producing an echeme lasting more than 10s. Among the other species in which the calling song normally consists of a single echeme (fischeri, festivus, stigmaticus and apenninus), the song conforms to the same basic pattern of a simple series of syllables, beginning quietly but soon reaching maximum intensity (see Figs 11, 12, 15, 16). The number and repetition rate of the syllables enable fischeri and festivus to be distinguished from the other two species and also from each other. However, stigmaticus and apenninus are extremely similar to each other in both calling and courtship songs; although the echemes of stigmaticus are on average longer than those of apenninus and are composed of more 398 D. R. RAGGE ECHEME ae a Lier seal re SYLLABLE GAPS 0-05 s J Stenobothrus stigmaticus ECHEME-SEQUENCE . eS) LOls 4 SYLLABLES ECHEME Loos sea; Stenobothrus grammicus Fig1 Oscillograms of the male calling songs of two species of Stenobothrus, showing the terminology used in this paper. SONGS OF THE WESTERN EUROPEAN Stenobothrus 399 syllables, there is an overlap in both these song characters and I have not been able to find a constant difference between the two songs. As these two species are completely allopatric I think it likely that apenninus is a recent derivative from stigmaticus that has diverged from it morphologically as a result of isolation in the Apennine mountain range. It would be most interesting to establish experimentally whether there is any ethological barrier to intercopula- tion between these two species. The remaining three species, nigromaculatus, grammicus and bolivarii, have calling songs consisting of a sequence of echemes; they also differ from the other six species in having syllable repetition rates of more than 50/s. As can be seen from Figs 10, 13 and 14, the calling song of nigromaculatus differs markedly from those of grammicus and bolivarii in that the constituent echemes are far fewer in number (usually 2—6) and much longer (usually lasting 1-2 s). The very short, rapidly repeated echemes of grammicus and bolivarii show a marked contrast with the other seven species, but sound similar to each other to the unaided ear, differing mainly in the syllable structure within the echemes. These two species are also similar morphologically and may well have evolved from common stock quite recently. As I have recently emphasized (Ragge, 1986: 220), the value of the songs as indicators of phylogeny is very limited. Their taxonomic importance, as would be expected of a mate recognition system, is almost entirely at the specific and subspecific levels. This can be exemplified by some fairly close intergeneric resemblances in song between certain species of Stenobothrus and Omocestus. The male calling song of S. apenninus, for example, is quite similar to that of O. raymondi (Yersin) (Ragge, 1986: figs 64-72), and there is also quite a close resemblance between the male calling songs of S. fischeri and O. broelemanni (Azam) (Ragge, 1986: figs 90-98). The members of each of these pairs are allopatric and so they have no need for differing recognition signals (although in each case there is a marked difference in courtship song). The calling song of all four of these species is a simple echeme and has presumably not been subject to the selective pressures needed for further differentiation. Echemes of this basic type also occur in several species of Chorthippus Fieber. New lectotype designations Gomphocerus (Stenobothrus) Bolivarii Brunner, in Bolivar, 1876: 327. Dr A. Kaltenbach of the NM, Vienna has kindly sent me a number of syntypes of this species, and I have selected and labelled a male as lectotype. I have labelled two other males and six females as paralectotypes. Stenobothrus apenninus Ebner, 1915: 551. Dr Kaltenbach has also kindly lent me six syntypes of this species, and I have selected and labelled an adult male as lectotype; this specimen was already labelled ‘typus’ by Ebner. I have labelled the remaining five specimens (two adult males, one adult female and two female nymphs, all labelled ‘paratypus’) as paralectotypes. STENOBOTHRUS Fischer Stenobothrus Fischer, 1853: 313. Type-species: Gryllus lineatus Panzer, by subsequent designation (Kirby, 1910: 160). Stenobothrodes Tarbinskii, 1948: 116 [as subgenus of Stenobothrus Fischer]. Type-species: Stenobothrus eurasius Zubovskii, by monotypy. Syn. n. Crotalacris Chopard, 1952: 280 [as subgenus of Stenobothrus Fisher]. Type-species: ‘Stenobothrus rubicun- dus (Germar)’ [= Stenobothrus rubicundulus Kruseman & Jeekel (see p. 422)|, by monotypy. [Synony- mized with Stenobothrodes Tarbinskii by La Greca, 1965: 70.] Diacnosis. O'. Head of typical Gomphocerine shape, foveolae well developed. Antennae usually filiform or slightly thickened towards tip, exceptionally clubbed (males of S. clavatus Willemse). Pronotal lateral carinae varying from sharply incurved to almost straight. Brachypterous to macropterous. Fore wings without bulge on anterior margin of precostal area; medial area usually widened, often conspicuous- ly so, with Cu, and Cuz sometimes partly or even completely fused. Hind wings varying from transparent to strongly smoky, sometimes with enlarged medial area. Ovipositor with lateral tooth on each valve. Tympanal aperture slit-like. 400 D. R. RAGGE Discussion. Stenobothrus can be recognized reliably only by the ovipositor, which has a lateral tooth on all four valves. Males can sometimes be confused with Omocestus, but in most cases can be recognized by the widened medial area of the fore wings. Both sexes of most species have somewhat broader pale bands along the pronotal lateral carinae than are shown by Omocestus and other related genera. The subgenera Stenobothrodes and Crotalacris are certainly not distinct from each other at the subgeneric level, and even considered together they do not in my opinion justify recognition as a distinct subgenus. On the other hand, I have followed Harz (1975) and other recent workers on the European fauna in regarding Omocestus and Myrmeleotettix Bolivar as distinct genera rather than subgenera of Stenobothrus; Stauroderus Bolivar and the Nearctic genus Bruneria McNeill, neither of which has a toothed ovipositor, are similarly excluded. As in the other large Gomphocerine genera Omocestus and Chorthippus, the male calling songs show a wide range of patterns in Stenobothrus, varying from the long single echeme of lineatus to the rapid sequence of short echemes of grammicus and bolivarii, and the remarkable wing-buzz of rubicundulus. DistriBuTION. All Europe, the larger Mediterranean islands, North Africa and temperate Asia as far as Mongolia. INCLUDED SPECIES. S. amoenus (Brisout), S. apenninus Ebner, S. berberus Uvarov, S. bolivarii (Brunner), S. bulgaricus Ramme, S. burri Karabag, S. carbonarius (Eversmann), S. caucasicus Dovnar-Zapolskii, S. clavatus Willemse, S. cobresianus Bei-Bienko, S. cotticus Kruseman & Jeekel, S. crassipes (Charpentier), S. croaticus Ramme, S. eurasius Zubovskii, S. festivus Bolivar, S. fischeri (Eversmann), S. graecus Ramme, S. grammicus Cazurro, S. kirgizorum Ikonnikoy, S. lineatus (Panzer), S. maroccanus Uvarov, S. miramae Dirsh, S. newskii Zubovskii, S. nigromaculatus (Herrich-Schaffer), S. palpalis Uvarov, S. posthumus Ramme, S. rubicundulus Kruseman & Jeekel, S. stigmaticus (Rambur), S. sviridenkoi Ramme, S. tadzhicus Mishchenko, S. ursulae Nadig, S. werneri Adelung, S. zubowskyi Bolivar. Keys to the principal western European species of Stenobothrus The only western European species not included in these keys are very local forms, sometimes of doubtful status, in which the song is unknown to me. Two keys are provided here, one based on morphological characters but with notes on song differences added where appropriate, and one based entirely on the male calling songs. Some of the species are difficult to separate morphologically and, in the absence of song information, some specimens may not be correctly identified using the morphological key. Reference should in any case always be made to the notes given on recognition in the accounts of each species. Key based primarily on morphological characters This key is for sexually mature adults, either freshly collected or showing good colour preservation. The terminology used for wing characters is illustrated in Fig. 3. 1 Hind wings vestigial in both sexes; fore wings reduced to about the same length as the pronotunim 9 (Apennime mountains)! .2..5. 26: sons-ce-ceew cetwecsecsutenacehetecbenestk apenninus(p. 418) = Both pairsof wings welldeveloped : :22. 25.521. csecBegossanthateerert cs oases clan co ceaceseeenoeee eee 2 2 Palpsibiackiondankebrownat theytipieessssisecseer acer a eee eseeeee ete tee tee ee eee grammicus(p. 410) —) ‘Palpsnotblackordark browmnlatithe tip e-n-ccasesce-co-es eee ereee eee e een eer eee cent eee eee ee eeeee 3 3 Mad ei ic2 cog atau dee ostninaateon wenaareetionictic shisha cisioeces Goa iove ee cet ae ete i ale ee eee eee Oe ee 4 = (BOM ale: oo acces ste ne ceiete ene oe dercactee ao emcl eee ete ae eee ee ont ee iceland, ae 10 4 Last segment of the maxillary palps bulbous and usually coloured orange-pink, red-brown or Vellow DIOWily 3.....505 caakacncuiscee: Serueaas sets RES ee ae are eee erent ne pene eae bolivarii(p. 413) — Last segment of the maxillary palps not bulbous and not coloured pink or brown ................... 5 5 Pronotal lateral carinae clearly angled in the prozona and interrupted at the angle by an additional'transverse sulcuS(Pig. 2) sc c.asateye see eet eeteat area eer eee ee festivus(p. 410) — Pronotal lateral carinae not or hardly angled in the prozona and not interrupted by an additional'sulcus (mot as tm Fig.2)). che atts woe aeak she cae cee ae ase ence eee eee ee nee ee 6 6 Stigmawithin3 mnofthe fore wingtip’ ios eaccsoes. fos os seeee dees eae (eae oe es oe ee a =» Stigma further than 3 mmifrom the fore: wing=tipyec..1.ce-se- oo, see ee ee eee oa eee ee ee 8 7 Medial area of the fore wing enlarged, as in Fig. 5, Cu, becoming fused with Cuz within the proximal half of the fore wing. Total length more than 14 mm. (Song a sequence of two or moreechemes, as im Pigs 50-34) en -aae- eenece eee ee eee hae ee eae nigromaculatus(p. 405) SONGS OF THE WESTERN EUROPEAN Stenobothrus 401 medial area stigma medial area M fes I 5 mm j so 2 ae ee Pett tt 2 = i ay aS sauce fay ris we SUVSESONS rrp MH ITA — 4 lin LAN Swe sone = = DTT - oR AS 8 5 nig ERS > — SWS OY | ASSL) TA Ss Ss — UT Figs 2-8 Pronotum and wings of species of Stenobothrus. 2. Dorsal view of the pronotum of S. festivus. 3. The right male fore and hind wings of S. rubicundulus, showing the principal veins and the wing terminology used in this paper. 4—7. The right male fore wing of (4) S. lineatus, (5) S. nigromaculatus, (6) S. fischeri, (7) S. stigmaticus. 8. The right male fore and hind wings of S. rubicundulus. 402 18 D. R. RAGGE Medial area of the fore wing not enlarged, as in Fig. 7, Cu, remaining separate from Cu until the distal half of the fore wing. Total length less than 14 mm. (Song almost always a single echeme, asin Hie; 15 ex 25. ta asie to haba eeee cae eater a aa ee el ae stigmaticus(p. 413) Cu, fused with Cuz at or near the base of the fore wing, as in Fig. 4. Cerci conical. (Song with syllable repetition cate less than 2)S).-33i5.o-.-02es.e2te- ahaa besser meee eee ee lineatus(p. 404) Cu, remaining distinct from Cuz in much of the proximal half of the fore wing (although the two veins often touch in places), not as in Fig. 4. Cerci laterally compressed towards the tip. (Song with syllable. repetition rate more that 4/s)\\ 22.2 - <2. 2d-cne)-2222eaeee Saeco sor ei'> sees 9 Fore wings very broad, as in Fig. 8. Hind wings as in Fig. 8, with the more distal part of the anterior margin strongly sclerotized and a much enlarged medial area........ rubicundulus(p. 422) Fore wings much narrower, asin Fig. 6. Hind wings not asin Fig. 8 .................06.06 fischeri(p. 405) Cu, fused or continguous with Cuz in at least part of the proximal half of the fore wing............. 11 Cu, remaining separate from Cu, until the distal half of the fore wing .........................00e0eeeee 14 Stigma within 4 mnrof the fore WiING-lIp ... 6 2<1b.90n0<n neuen eine rmmalonnvacionne nigromaculatus(p. 405) Stigma furthenthan 4 mim fromthe fore wWithp-tip 5... 2<.0c.is»caasaceseaenvoareqe-sateveesneenaes teen 12 Cu, fused with Cuz at or near the base of the fore wing ..................cseeeeeeeeeeeeeenees lineatus(p. 404) Cu, remaining distinct from Cu in much of the proximal half of the fore wing (although the two veins sometumes touch or fuse in places). 6 x..sacddacece sent ccete sone os ere cee acne eae ee 13 Hind wings as in Fig. 8, with the more distal part of the anterior margin strongly sclerotized and amuch enlarged medial area card ies.... cece devoceee ve eeeseeeenc sence eee eee rubicundulus(p. 422) Hind wingsnotas in Fig® 2 shee at hoc at aes ait bcm at eee eee fischeri(p. 405) enethofthetorewines lessithant lima e.-eeeeecs-eeceee eee tee ee eee eee eeeee eee ree stigmaticus(p. 413) Length of the fore wings more tham 11) mmo... 0.5 vee cans ceive saewnou the Ceaerenaceeseh eae eee 15 Hind wings as in Fig. 8, with the more distal part of the anterior margin strongly sclerotized and amuch enlarged mediallarea. [22.5 .cis0.<2 cateoeceesns andes .eee eee eee eeee ene rubicundulus(p. 422) Hind wings not. as i Pig. 8 «0. esas «desictgs adap ties as onin os tile sents oltre prec ae Soe eee ee 16 Eengthofithe tore wines more thane Spano ieeeen a eeeceeeteeeaeeeee ener eee eee eee ee bolivarii(p. 413) Ienpth ofthe fore: wingsilessthamilspmunis sasnesee o-- cr cee aes eeneee nee eeetee ae eee ee 17 Pronotal lateral carinae clearly angled in the prozona and interrupted at the angle by an additional transverse sulcus (Fig. 2). Cu; and Cuz diverging in the proximal half of the fore WHS. aaah wines taomew saints Dales ebro cae Seats es Pea ee SD festivus(p. 410) Pronotal lateral carinae not or hardly angled in the prozona and not normally interrupted by an additional sulcus (not as in Fig. 2). Cu, and Cu converging or running parallel in the proximal half of the fone: Wing 0x sos chess oS 0 «cine wince so doneslo cob one one REN see eee 18 Stigma within 4mm of the fore WiRB IP o.5..S.. 0.5- donmanaas ann ocwanniensivdninna cannes nigromaculatus(p. 405) Stigma further than 4mm from the fore WINg-tip .....6522.-..cs0scesercosermccecersuncdeastes fischeri(p. 405) Key based on song characters (see especially Figs 9-17) This key is based on the calling songs produced by isolated males in warm, sunny conditions. Many of the song differences can be easily detected by the human ear assisted by the second hand (or digital count) of a watch, but a few of the couplets require oscillographic analysis, or at least the ability to play back the recorded song at a slower tape speed. I have been unable to find a consistent difference between the calling songs of stigmaticus and apenninus (couplet 9), but these species are completely allopatric and easily separated morphologically. See p. 397 for definitions of the terms ‘echeme’, ‘syllable’ and ‘gap’. 1 Song consisting, at least partly, of aloud buzz produced by wing-vibration, either on the ground orin flight (Figs 109046) cosas as2.cwecSante teach cee eee eee eee teen eee ee rubicundulus(p. 422) Song produced entirely by femoro-alary StrithulahOm 222 ooo coc cwnaem ass susioms Jovenen ao sepaaeanee 2 Song asequence of echemes in rapid succession (at least U/S) <<<. cceces eer oelsveee scent sacnatepen se Qen 3 Song a single echeme or a sequence of echemes repeated at the rate of one every 2 s or more SLOWLY, cdosesave nsec dauctnge suena comes de tenatlplent tae neeenistaee tease me geinns tan eines oskioeing sceioelsa sets eee eae eee “4 Each echeme lasting less than 80 ms, loudest towards the end (Figs 62-72) ....... grammicus(p. 410) Each echeme lasting more than 80 ms, loudest at the beginning (Figs 73—76) ........ bolivarii(p. 413) Song consisting of a single echeme lasting more than 10 s and produced by very slow leg movements (less than 2/s); syllables lasting 0-5—1-0s (Figs 18-30) .................2.. lineatus(p. 404) Song consisting of one or more echemes, each lasting less than 10s and produced by more rapid leg movements (at least 3/s); syllables lasting less than 0-2§ ................ceeeceeeeeeeeeeeeeeeneenes 3 Echeme usually lasting more than 5 s, with the syllables, at least in the early part of the echeme, 403 SONGS OF THE WESTERN EUROPEAN Stenobothrus (96 ‘Soe “dd) Z pur | safqey, Woy eyep [[Ny dy} UIeIGO 0} pasn aq URD pue spell d10M suIeISOTIIOSO 94} YOIYM WoIZ SsUIPIOI1 JY} 0} JOJOI sJoquuNu [ews oY], “snayjoqouals Jo saidads Jo sduos Suryjeo afew eo1dAy Jo swesZo]]IOsSQ _L[-6 SBIq : Sig 1 S09 snjnpunsigqn4 Zl z/€09 snuiuuadp ol 6lL snoyowBiys Gl ggg $$ B/loS DAIJOg Al p/EET = sNJIWWDIB €l 8 FE SNES SMAHS®S ZT] eve ay rsiy |] si c/eey = snyojn2owosBiu Oy Wize snyauly 6 404 D. R. RAGGE composed of clearly separated upstroke and downstroke sounds of equal amplitude, as in Figs £03 N06. 0220 fae nsve wacdegeunsiaes oo teak tate oe ee ee Sg eee IR a eae rubicundulus(p. 422) — Echeme lasting less than 5 s, with the syllables composed mainly of downstroke sounds (not as IM FIgs VOS—NOB). ek sad sees nee seca d awan eee eee eeeolabe acerca meek Re 6 6 Song consisting of at least 2 echemes; syllable repetition rate more than 50/s (Figs 31-44) nigromaculatus(p. 405) — Song almost always consisting of a single echeme; syllable repetition rate less than 20/s ........... fj 7 Echeme composed of more than 40 syllables and usually lasting more than 3 s (Figs 45-53) fischeri(p. 405) — Echeme composed of fewer than 40 syllables and usually lasting less than3s ..................-...05. 8 8 Syllable repetition rate less than 12/s; early part of echeme with about 7-8 gaps in each syllable (Bigs SA Gil aoe: iy to ah oa oa gamete wale ees coh pe eee Soeree oene cae Mette eee festivus(p. 410) — Syllable repetition rate usually more than 12/s; early part of echeme with fewer than 5 gaps in each syllable (E108) 7 F102) oo ssit dost cases Suece sedel nok «ebaiiesetuncaciebamsesnaseeoeesaes Ree eae aman 9 ‘Occurring oniyin.the Apennine mountains <1... 5. ois 32a ner~ oo eee erercomeea nde eeone apenninus(p. 418) — Notoccurringinthe Apennine mountams ... 05-2. .0.. c-ctosceeencene a sense scone seater stigmaticus(p. 413) The songs of the western European Stenobothrus Stenobothrus lineatus (Panzer) (Figs 4, 18-30) Gryllus lineatus Panzer, 1796: 9. Type(s): GERMANY (lost); the two ‘neotypes’ designated by Harz (1975: 759) are clearly invalid. REFERENCES TO SONG. Oscillogram: Dumortier, 1963; Elsner, 1974a, 1975; Elsner & Popov, 1978; Grein, 1984; Haskell, 1957, 1958. Diagram: Bellmann, 1985a; Duijm & Kruseman, 1983; Haskell, 1957; Jacobs, 1950, 1953; Luquet, 1978; Ragge, 1965. Frequency information: Dumortier, 1963; Haskell, 1957, 1958. Musical notation: Yersin, 1854. Verbal description only: Baier, 1930; Broughton, 1972; Chopard, 1922, 1952; Faber, 1928, 1953; Harz, 1957; Haskell, 1955. Disc recording: Andrieu & Dumortier, 1963; Grein, 1984; Ragge, Burton & Wade, 1965. Cassette recording: Bellmann, 1985b. RECOGNITION. Males of this species may be distinguished from the rather similar species nigromaculatus and fischeri by the cerci, which are simply conical (laterally compressed towards the tip in the other two species). In both sexes the veins Cu; and Cuz are more completely fused together in the fore wings than in nigromaculatus and fischeri, and the hind wings are more strongly smoky. In the field males may be recognized at once by their highly characteristic calling song, in which the syllable repetition rate (under 2/s) is lower than in any other European grasshopper. SonG (Figs 18-30). The calling song of this species is unique among the western European Gomphocerinae in both the slowness of the leg movements and the quality of the sound produced. It consists of an echeme lasting about 10—25 s and composed of about 15-35 syllables repeated at the rate of about 1-2—1-5/s. The echeme usually begins quietly but reaches maximum intensity within a few syllables. The sound is usually continuous, with a highly characteristic, rather wheezy quality, and showing slight fluctuations in intensity corresponding with the leg movements — though partly masked by the fact that one hind leg always moves slightly in advance of the other. In the presence of a female the male usually produces longer echemes (with a more gradual crescendo at the beginning), sometimes lasting a minute or more but otherwise similar to the calling song. Between these echemes the male produces a quite different ‘ticking’ sound (Figs 21, 22, 26, 27) by moving the hind legs through a much smaller angle than in the calling song and at a much higher rate (usually S—7 syllables/s). This ticking sound lasts typically for about half a minute before another echeme of the calling song type is produced, but it may be greatly prolonged, sometimes lasting for even longer than an hour. The ticking sound and the ‘calling song’ echemes usually alternate several times before the male attempts to copulate with the female. The courtship song of this species and the leg movements that produce it have been described in great detail by Elsner (1974a, 1975). DISTRIBUTION. Widespread in Europe, but not reaching further north than southern Sweden and limited to mountains in the southern peninsulas. The range extends eastwards across central Asia to Mongolia. SONGS OF THE WESTERN EUROPEAN Stenobothrus 405 Stenobothrus nigromaculatus (Herrich-Schaffer) (Figs 5, 31-44) Acridium nigromaculatum Herrich-Schaffer, 1840: 10. Type(s) unknown; the two ‘neotypes’ designated by Harz (1975: 761) are clearly invalid. REFERENCES TO SONG. Oscillogram: Grein, 1984; Schmidt & Schach, 1978. Diagram: Bellmann, 1985a; Duijm & Kruseman, 1983; Jacobs, 1950, 1953. Verbal description only: Faber, 1928, 1953; Harz, 1957. Disc recording: Grein, 1984. Cassette recording: Bellmann, 1985b. RECOGNITION. Males of this species may be distinguished from lineatus by the cerci, which are laterally compressed towards the tip (simply conical in Jineatus). In the fore wings of both sexes the veins Cu, and Cuz are less completely fused together than in /ineatus, and the stigma is nearer the wing-tip than in fischeri (cf. Figs 5, 6). In the field males may be recognized quite easily by their calling song, in which the syllable repetition rate (about 70—110/s) is much higher than in /ineatus (about 1-2—1-5/s) and fischeri (about 10—15/s). Sone (Figs 31-44). The calling song consists of a series of 2-6 echemes (each beginning quietly but showing a rapid crescendo), separated by intervals of 1-3 s. Each echeme lasts about 0-8—2-0 s (rarely up to 2-5 s) and consists of about 90—200 syllables (rarely up to 250) repeated at the rate of about 70—110/s. In the presence of a female the male produces a quite complicated courtship song, consisting of two alternating phases (Figs 35, 36, 41-44). The first phase consists of a series of fairly quiet echemes separated by intervals of about 0-6—0-9 s; during these intervals a few even quieter sounds are often produced. Each echeme lasts for about 0-3—0-9 s and is composed of about 40-80 syllables repeated at the rate of about 80—100/s. This phase lasts for an indefinite period, usually more than 30 s and often several minutes. The second phase, which follows abruptly, consists of a series of about 3-7 louder and longer echemes, each of which is preceded by a much shorter echeme. The shorter echemes last for less than half a second, and in fact after the first longer echeme they are often reduced to little more than a single syllable. The longer echemes usually last for about 0-8—2-2 s and are subdivided into two parts, one with slower syllables (about 40-—60/s) and one with faster syllables (about 70—120/s). At the end of this second phase the male either attempts to copulate with the female or, after a short pause, begins the first phase again. Note that the difference in temporal pattern between the oscillograms taken from recording 482/3 (Figs 35, 41, 42) and those taken from recording 184/1 (Figs 36, 43, 44) may have been caused, at least partly, by the fact that recording 184/1, unlike any other field recording used in this study, was made in conditions of hazy sunshine, so that the subject was receiving less radiant heat than it would have in full sunshine. DIsTRIBUTION. Southern Europe and central Asia; absent from the Iberian Peninsula except in the vicinity of the Pyrenees, and confined to uplands in the Italian and Balkan Peninsulas. Stenobothrus fischeri (Eversmann) (Figs 6, 45-53) Oedipoda fischeri Eversmann, 1848: 11. Syntypes of both sexes, U.S.S.R.: Ural Mountains (ZI, Lenin- grad) [1 o’ , 1 2 syntypes examined]. REFERENCES TO SONG. Diagram: Luquet, 1978. Verbal description only: Faber, 1953; Harz, 1957. RECOGNITION. Males of this species may be distinguished from /ineatus by the cerci, which are laterally compressed towards the tip (simply conical in /ineatus). In the fore wings of both sexes the veins Cu, and Cuz are less completely fused together than in Jineatus and the stigma is further from the wing-tip than in nigromaculatus (cf. Figs 5, 6). In the field the single echeme of the male calling song, which lasts for 3—5 s and is composed of more than 40 syllables, enables this species to be distinguished from Jineatus, nigromaculatus, grammicus and bolivarii. The calling songs of festivus and stigmaticus are more similar, but in both these species the echeme is shorter, with fewer than 40 syllables. D. R. RAGGE 406 (96€ “Soe “dd) Z pue | safqvy, wo. yep [[Ny ay) UIeIGO 0} pasn 9q URS PUR SPRL SIOM SUILIBOT[IOSO DY} YOIYM WO] SSUIPIOIOI IY} O} JOJO SIOQUINU [[PUIS BY, “s8uOs diys}in0d OM} Jo Weg ‘ZZ ‘[Z *39] puly ouo (0Z) pur sso] purty OM) (6T ‘8T) YIM soyeUl Jo ssuOs SUTT[eD ‘QZ—-I “Snivauy snaysOqouafg JO Sa|eUI DAY JO SBUOS dy} JO SWILIZOT[IOSO + 7Z—-SI SBlq ss 097 Co 7/@ZE 0Z Sz ‘84 I/LZE él yz 814 Z/LLS gl €z ‘Big 407 ‘39] pury suo ATUO YIM [eu B WOIy Ie YE pu CZ SSL “ZZ—I SSL Ul UMOYS sMJvaUT] SN4yJOgoUaTS JO SBUOS 94} JO Syed pajeoIpUI OY} JO SWILISOTIIOSO J9\se.y OE-EZ SBI t SLO ] 0} 3 SONGS OF THE WESTERN EUROPEAN Stenobothrus Cr Utes were BOP YA dean Afi, PAH D. R. RAGGE 408 "(96€ ‘Soe “dd) Z puke | sojqey, Woy eyep [[Ny Oy) UTe}GO 0} pasn dq URS pUP SpE d1OM SUILITOT[IOSO dy} YIYM WO] SSuIP1OIEI 9Y} 0} JoJo1 SJaquiNu [BUS BY, “(COP “d UO syIeW9I Oy} 90S) s8uos diys}in0s OM} Jo WIRY “O¢ ‘SE “SBUOS BUILD “pE-TE “SNID[NIDWLOASIU SNAYJOGOUAIS JO S2|BU XIS JO SBUOS dy} JO SWRIDOT[ISQ 9E-TE SBA 1c \/rsl 9€ e/eey «= CS | be [({——_— ty 614 ty ‘614 ¥/E£09 VE [Eee EEE | Or 614 “ €€ 6e 614 01/609 cE ge “614 7/78V Le Ze ‘64 409 SONGS OF THE WESTERN EUROPEAN Stenobothrus ‘O€-TE SBLJ UI UMOYS SnynINIDUUIOLS1U SNAYIOGOUAIS JO SBUOS dy} Jo syed PO}LSIPUT 9Y} JO SWIRIZOT[INSO I9}sey pp—-Le¢ sBIq $¢:0 ! a4 Ze 410 D. R. RAGGE SonG (Figs 45-53). The calling song is a single echeme lasting about 3-5 s and consisting of about 40-55 syllables repeated at the rate of about 12—13/s. The echeme begins quietly, usually reaching maximum intensity about halfway through its duration. Oscillographic analysis shows that there are gaps in each syllable. Previously published accounts of the calling song of this species have been based almost entirely on Faber (1953), who gave lower figures for both the duration of the echeme (‘2 oder etwas mehr Sekunden’) and the syllable repetition rate (8-914/s), although suggesting that these figures applied to males singing in full sunshine. Faber’s observations were made in the laboratory on males collected for him from Oberweiden, east of Vienna in Lower Austria, whereas mine were made in the field in southern France; in France I would expect such a low syllable repetition rate only in dull weather or at unusually low air temperatures. In the presence of a female the male produces a long and complicated courtship song (Figs 47, 50, 53), which has been fully described by Faber (1953: Beilage 4). DISTRIBUTION. In western Europe this species seems to be confined to two isolated areas: southern France and central Spain. Further east it occurs from Austria eastwards across eastern Europe (including the Balkan Peninsula) and central Asia to Mongolia. Stenobothrus festivus Bolivar (Figs 2, 54-61) Stenobothrus festivus Bolivar, 1887: 94. Syntypes of both sexes, Spain: Uclés, Aranjuez, San Ildefonso, Escorial, Gredos, Navacerrada (7MNCN, Madrid). REFERENCES TO SONG. No published work known to me. RECOGNITION. This species is perhaps most likely to be confused with grammicus or bolivarii, which share with festivus the strongly inflexed pronotal lateral carinae. However, males of festivus lack the modified or characteristically coloured last palpal segments shown by the other two species, and in both sexes the pronotal lateral carinae are more sharply angled in the prozona, where they are clearly interrupted by an additional transverse sulcus on each side of the pronotum (Fig. 2). In the field the single echeme of the male calling song enables this species to be distinguished easily from grammicus and bolivarii, which both produce a rapid sequence of echemes. The male calling songs of fischeri and stigmaticus are more similar, but fischeri has a longer echeme, with more than 40 syllables (usually fewer than 30 in festivus) and in stigmaticus the syllable repetition rate is usually more than 12/s (less than 12/s in festivus). Sonc (Figs 54—61). The calling song is a single echeme lasting about 2—3 s and consisting of about 20-30 syllables repeated at the rate of about 7—11/s. The echeme usually begins quietly, reaching maximum intensity about halfway through its duration or a little earlier, but sometimes it begins more abruptly and sometimes there are one or more louder syllables before the quiet beginning. Oscillographic analysis shows that there is a characteristic pattern of gaps in the syllables: at the beginning of the echeme the syllables are completely broken up by gaps, but as the echeme progresses the gaps gradually disappear so that there are usually no gaps in the last few syllables. In a song produced by only one hind leg the gaps often persist until rather later in the echeme (Fig. 57), since there is no tendency for the gaps in the syllables produced by one hind leg to be obscured by sounds produced by the other. DISTRIBUTION. Widespread in the Iberian Peninsula and occurring locally in southern France. Stenobothrus grammicus Cazurro (Figs 62-72) Stenobothrus grammicus Cazurro, 1888: 457. Syntypes of both sexes, SPAIN: Navacerrada, Penalara, Escorial, San Ildefonso, Ofia (7MNCN, Madrid). REFERENCES TO SONG. Diagram: Luquet, 1978. RECOGNITION. Both sexes of this species may be recognized easily by the palps, which are black or dark 411 SONGS OF THE WESTERN EUROPEAN Stenobothrus ‘(s6¢ “d) T e1qe], Woy eep [NJ 94} UTe}GO O} posn dq URS PUP Ope 919M SWIPISOT[IOSO 94} YSIYM WOIJ SSUIPIOIOI 9} O} JOJoI sIOquINU |[eUIs oY] “SuOs diysjino|g ‘¢¢ ‘OS ‘Lp ‘S8uos BUTT[eD “ZS ‘TS “6b ‘8h ‘OP ‘Sp ‘Mayasy snayjoqouals JO Sa|eU 9914} JO SSUOS ay} JO Spoads yUaIaTIp 9914} Je SUILIBOT[IOSE. © ES—Sp SBIg S10 es eez J Uae OY ewe OY —Fiiiine —_ 00 —§$ 6y “B14 Sy 614 D. R. RAGGE 412 ‘(96¢ ‘S6¢ dd) Z puke {| sajqe, Woy eyep [[NJ ay} UIe}qGO 0} pasn aq URS PUL Spe SI9M SUILIZOT[IOSO DY} YOIYM WO] SSUIPIOIOI OY} O} J9JOI SIOQUINU |[BUUS oY], “Soy PUTY BUO YIM (T9 ‘6S “LS ‘S¢) JOYI0 oY} puke sso| pUurY OM} YIIM (C9 ‘8g ‘9g ‘pS) QUO ‘snaysaf sn4ysoqouaIg JO Sa;BW OM} JO SdUOS SUTT[ed dy} JO Spoads jUSIAIJIP INOJ ye SUIBIBOT[IOSE. T19-pS SBI PS100 | 9 09 ib « ‘al i ret F 6S L al ey t9 “B14 gc SS eee 09 ‘614 f $¢-0 ZS ! ! i | i EE EEE SE ees 6s ‘84 96 (a Se ee ee ae gs 514 i s¢ HE GC les 69S Gases SSSI) Zs ‘614 9¢ 614 SONGS OF THE WESTERN EUROPEAN Stenobothrus 413 brown at the tip. In addition the male antennae are curved outwards and noticeably thickened and darkened towards the tip. In the field the distinctive calling song, consisting of about 10-30 very brief and rapidly repeated echemes, enables males to be easily distinguished from most other species of Stenobothrus; however, oscillographic analysis is necessary to show clearly the difference from the calling song of bolivarii, which has longer echemes with a rather different syllable structure (see below). Sonc (Figs 62—72). The calling song consists of a sequence of about 10-30 echemes lasting about 5-15 s; the echemes are thus repeated at the rate of about 2/s. Each echeme lasts about 40-80 ms and is composed of a series of 4—8 syllables. The only other species of Stenobothrus known at present to have a rather similar calling song is bolivarii, but oscillographic analysis shows that species to have longer echemes, lasting about 80-110 ms, with a rather different syllable structure: in grammicus the loudest syllables are towards the end of each echeme, whereas in bolivarii the loudest syllable is at the beginning (cf. Figs 71, 76). Luquet (1978: 427) gives the rather lower number of 6—8 echemes in each sequence of the calling song, based on observations in the Mont Ventoux area of Vaucluse in SE. France. My own studies of four males at two localities, also in the region of Mont Ventoux, suggest that this number is unusually low. In the presence of a female the male produces a series of quieter and much longer echemes (Figs 64, 67, 70). Each of these echemes lasts for about 1—2 s and consists of a soft, continuous and rather amorphous sound. Between the echemes there are intervals of varying duration (usually between 3 and 15 s), during which quieter sounds are often produced. DisTRIBUTION. Known only from upland areas in southern France and the Iberian Peninsula. Stenobothrus bolivarii (Brunner) (Figs 73-76) Gomphocerus (Stenobothrus) bolivarii Brunner, in Bolivar, 1876: 327. LECTOTYPE: ©’, Spain: Sierra de Guadarrama, La Granja (NM, Vienna), here designated (see p. 399) [examined]. REFERENCES TO SONG. No published work known to me. RECOGNITION. Males of this species may be recognized by the maxillary palps, in which the last segment is bulbous and usually coloured orange-pink or reddish brown. Females are more difficult to recognize, as the last segment of the maxillary palps is more normal in shape and is not always distinctively coloured like that of the male; however, in the upland areas of the Iberian Peninsula where bolivarii occurs the only other species of Stenobothrus with which females are likely to be confused is grammicus, in which both pairs of palps have black or dark brown tips in both sexes. In the field the short, rapidly repeated echemes of the calling song enable males to be easily distinguished from most other species of Stenobothrus; the calling song of grammicus is, however, rather similar and oscillographic analysis is necessary to show clearly that bolivarii has longer echemes with a different syllable structure. SonG (Figs 73-76). The calling song is quite similar to that of grammicus, consisting of a sequence of about 10—20 echemes lasting about 4—8 s; the echemes are repeated at the rate of about 2—3/s. Each echeme lasts about 80-150 ms and is composed of a series of about 5-10 syllables; the echemes are thus rather longer than those of grammicus and show a different syllable structure, with the loudest sound at the beginning of each echeme (cf. Figs 71, 76). DisTRIBUTION. Known only from upland areas in the Iberian Peninsula. Stenobothrus stigmaticus (Rambur) (Figs 7, 77-93) Gryllus stigmaticus Rambur, [1838]: 93. Syntypes of both sexes, Spain: Sierra Nevada (lost); the two ‘neotypes’ designated by Harz (1975: 768) are clearly invalid. REFERENCES TO SONG. Oscillogram: Grein, 1984. Diagram: Bellmann, 1985a; Duijm & Kruseman, 1983; D. R. RAGGE 414 ‘(s6¢ “d) T 21qe,L WoL eIep [[NJ oY} UIeJGO 0} pasn oq ULd pue OPEU DIOM SUILIBOT[IOSO OY} YOIYM WOIJ SSUIPIOIII OY} 0} IOJOI SIOQUINU [PWS dT, *g9 ‘SO ‘ZO SBLY Se OPW OURS OY} JO BUOS diys}INOD ‘C/ ‘19 “pg ‘s8uos SulT[eD *69 ‘89 “99 ‘SO “EQ ‘ZO “SNIIMMUDAS SNAYJOGOUAJS JO SI|PUI OA} JO SBUOS dy} JO Spoods JUDIOIJIP 9OIY} Je SWILISOT[IISO. OL—-7Z9 SBA I SLO Nl a7 on 7‘ Nh OZ | H l Hy { oz “614 | 99 69 ‘614 S9 ¥/EET v9 \/vETZ 9 99 Bly weet TO $A A + 415 SONGS OF THE WESTERN EUROPEAN Stenobothrus “(c6¢ “d) T 91q4eL Woy ejep [[NJ 94} UTe}qO 0} pasn 9q ued puke ope oIOM SWIPIZO][INSO ay} YSIYM Wo Zurpi0se1 OY} 0} SIoJol 8/TOS Joquinu oy], 1740A1/0qg ‘g JO SuOs Sur|]ed a[eu ev Jo spsads JUSIIJFIP INOJ Je SWIBIBOTIIOSC *9/—€/ *69 ‘QQ SBLJ UI UMOYs sno "§ JO Ssuos oy} jo sired pojeoipur oy} Jo sureSO]]INsO Josey “ZL “TL “apaljog pure snomuupss sniyjoqouajs Jo s8uos Suly[ed Q[eW BY} JOSWIRISOT[INSO 9L-TL S814 “sion? 510-0 92 SZ ‘B14 8/t0S €Z Ts10-0 | CZ LZ D. R. RAGGE 416 (96 ‘c6¢ dd) Z pur | sajqe yy, Woy eyep [[Ny Oy} UTe}GO 0} pasn oq UD pu apeUl IIOM SWILISOTIIOSO 9Y} YOTYM WIJ SBUIPIOIII 9Y} O} JOJoI SIoquinu [Jews OU,L, “T8-LL S8lq Ul UMOYs s8UOs a4} Jo sj1ed pojyeoIpuT OY} Jo SWIRIDOTIIOSO 19}se,J “Qg—Z “SUOS diysjinod v Jo Ae “1g “B9] puTY oUO (98) pue sso] pury OM (6/—L) YIM soyeU Jo ssuos Sued “Y8—LL “SNINDWIYS sn4yOgouaG JO S2|BW SAY JO SSUOS 94} JO SWILISOTIOSO ~=98-LL SBA 5¢-0 98 c8 68 ‘614 £8 c8 1ZL 8 98 ‘614 “ey OR eos 6/ “Br ‘Bi se ‘614 waz ee él ff See) ————_——— 8 “614 zg 64 417 “S9] pury ouo ATUO YyIM seu B WO J1e €6 PUL 16 SSIJ “O8—L/ SSIJ Ul UMOYS SnoDUSYS Sn4yoqouay JO SBUOS dy} JO sj1ed poyeoIpuT ay) Jo suTeAZOT|IOSO I9}seJ €6-LS S3Iq oon €6 HN | HAN i Wht | C6 06 €6 ‘614 SONGS OF THE WESTERN EUROPEAN Stenobothrus 7 614 £8 418 D. R. RAGGE Jacobs, 1950, 1953; Ragge, 1965. Verbal description only: Broughton, 1972; Faber, 1928, 1953; Harz, 1957. Disc recording: Grein, 1984; Ragge, Burton & Wade, 1965. Cassette recording: Bellmann, 1985b. RECOGNITION. S. stigmaticus is conspicuously smaller than any other fully winged species of Stenobothrus occurring in western Europe. Were it not for the size difference it could be confused with nigromaculatus, but the veins Cu, and Cu, are much more widely separated in the fore wings than in that species. In the field the male calling song, usually consisting of a single echeme lasting 1—3 s, enables this species to be distinguished easily from all other species of Stenobothrus with overlapping distributions except fischeri and festivus; these species produce single echemes of a similar kind to stigmaticus, but in fischeri the echeme is longer, with more than 40 syllables (fewer than 40 in stigmaticus), and in festivus the syllable repetition rate is less than 12/s (usually more than 12/s in stigmaticus). Both the calling and courtship songs are extremely similar to those of apenninus, but as that species is confined to the Apennines, where stigmaticus does not occur, confusion between the songs of these two species is unlikely. (See also the remarks on pp. 397, 399.) SonG (Figs 77-93). The calling song is a rather quiet echeme lasting about 1-3 s and consisting of about 25-40 syllables repeated at the rate of about 10—20/s (usually 14-18/s). The echeme usually begins quietly, reaching maximum intensity within about one second. Oscillographic analysis shows that there are usually gaps in the syllables, especially near the beginning of the echeme, but they are not always very clear. The echemes are usually produced singly and repeated at irregular intervals, but occasionally two echemes are produced within a few seconds of each other (Fig. 79). In the presence of a female the male produces a series of echemes separated by intervals of about 2-7 s (Figs 81, 86, 91). Each echeme is similar to that of the calling song, but is quieter, longer (usually lasting 3—S s and composed of about 50-70 syllables) and with one hind leg moving more vigorously than the other; the last few syllables are rather modified and then the hind leg that has been moving more vigorously produces a short, loud syllable by a very quick downstroke. The more vigorously moving hind leg alternates from one side to the other in successive echemes. Sometimes the loud syllable at the end of each echeme is omitted. DIsTRIBUTION. Widespread in western and central Europe, but not reaching as far north as Scandinavia. In southern Europe it occurs in the mountainous parts of the Iberian Peninsula and the northern part of the Balkan Peninsula but is apparently absent from the Italian Peninsula (although occurring in the Italian Alps). It is recorded from the Rif mountains in Morocco and its range extends eastwards into European Russia and Asia Minor. Stenobothrus apenninus Ebner (Figs 94-102) Stenobothrus apenninus Ebner, 1915: 551. LECTOTYPE ©, Ivaty: Lazio, Monti Reatini, Terminillo, 6.viii.1912 (R. Ebner) (NM, Vienna), here designated (see p. 399) [examined]. REFERENCES TO SONG. The only published reference to the song known to me is that of Harz (1975: 770), in which he quotes Faber as having told him in 1971 that the song was indistinguishable from that of stigmaticus. RECOGNITION. This montane species is endemic to the Apennine range in the Italian Peninsula, where it may be easily distinguished from other species of Stenobothrus by the vestigial hind wings of both sexes and the much reduced fore wings of the female. In the field the calling song of the males, consisting of a single echeme lasting 1-2 s, also enables this species to be distinguished easily from Italian species of the genus occurring in the Apennines. SonG (Figs 94-102). The calling song is a single echeme lasting about 1—2 s and consisting of about 20-30 syllables repeated at the rate of about 15—20/s. The echeme usually begins quietly but reaches maximum intensity within 0-5 s. Oscillographic analysis shows that there are clear gaps in the syllables throughout the echeme. In the presence of a female the male produces a quieter courtship song (Figs 96, 99, 102) very similar to that of stigmaticus (see above) but with shorter echemes (often lasting about 2 s and composed of about 30 syllables). DISTRIBUTION. Known only from the Apennine range in Italy. 419 SONGS OF THE WESTERN EUROPEAN Stenobothrus (96€ “d) T FIquJ, Wo4y eJep [IJ ay} UIE}QO 0} PAsN oq URS PUL aPeU 919M SWILISOT[IOSO BY} YOIYM WO sdUIPIOIAI ay} 0} 19J91 sIaquINU [JeUIS oY |, “3UOS diys1inod *Z0T ‘66 ‘96 ‘ssuos SUTTIED “TOT ‘OOT ‘86 ‘26 ‘S6 ‘P6 ‘Snuiuuadn snayjoqoualg Jo sayew 9014) Jo sduos ay) Jo spaads jusrayIp sary) ye sweISOT[IOSO ZOT—b6 SBIq SLO ZOL — 5 al zol ‘614 ' 86 | LS 5 SH 101614 | | L6 _ elt | ool 614 s¢ £609 O96 , EEE | 6 66°Pld z/€09 1/809 G6 V6 86 ‘614 26 814 ‘(96¢ ‘d) Z puke | solqe, Wory eJep [[NJ Oy) UTe]GO 0} pasn aq UkD PU Ope 919M SWILIDO][INSO BY} YOIYM WO] SSUIPIOIII DY} 0} JOJOI SIOQUINU [[eWs SUL “POT PUe EOT SITY Ul UMOYs UONR[NpPLYs Are[e-O1OW9y ay} Jo sjied poiedIpU dy} JO SWIRIZO][IOSO 19\Se4 “SOT—COT ‘punoss oy} UO poUleWal se dy} o[IYM ZzNq-3uIM e Aq AjoeIPOWIT POMOT[O} oWOYydE AIe[e-O1OWI] Y “POT “UONR[NpIys Are[e-o1OWsy A[aInd Jo aurayo9 UY “EOL ‘SN/NpUNsIgna snayjoqouaTs JO S2[eUl OM) JO ZZNQ-SUIM pu ssUOs SUT][ed ay} Jo spsads yUSJJIP 9914} Je SWIRISOT[IOSE OT-EOT SBT I SLO l ty | Ky 90l D. R. RAGGE S/l09 vol Mra pig ATE agg UNE SS veer pe ' ay 4 1 6 €0Ol 420 421 (96¢ “d) Z pur | saiqeL Wo4y ere [Ny oy} UIe}GO O} Pasn oq ULD PUR IPR 919M SWILIZOT[IOSO OY} YOIYM WOIJ SSUIPIOIII JY} O} JOJOI SIDQUINU [[eWIS OY, “}YSIY SuLAOY SuLINp poonpoid SBM (TT ‘8LJ Ul UMOYS ZZNQ-3UIM PUOdas oY} JO ed nq ‘puNoIS oy} UO seM JPW 9Y} O[IYM Poonpold o10M YT “BLJ Ul UMOYS ZzNQ-sUIM }SIY JY} pUL GOT “SIA Ul UMOYS ZZNG-3UIM OU], ‘SMNpUuNdIgn4 Sn4yJOgoua]gS JO S2{BUI 9d1Y} JO SOZZNG-SUIM 9Y} JO SWILISOT[INSO = 9TI-60T SBA SLO OL 4 SIL = 4 ~ < ~ 2 S PLL wa | | ry | X 2) YL | < ww 5 l sc, 1 % S-0 =] ell 5) V4 [4 | | | | | : { | i) | i H = ; 91 ‘B14 a cll = matin w oa / 1 1} | = a fe su Big LLL ° | 1| | yyy | ty | | lyyid] | a ; ie) Zi HW : bie | je) eae ee. eee nec ee Seee j a vit Big wore OLL 422 D. R. RAGGE Stenobothrus rubicundulus Kruseman & Jeekel (Figs 8, 103-116) Gryllus miniatus Charpentier, 1825: 155. Syntypes of both sexes, Austria: Alps, and GERMANY: ‘in Saxoniae montibus’ (lost); the two ‘neotypes’ designated by Harz (1975: 775) are clearly invalid. [Primary homonym of Gryllus miniatus Pallas, 1771: 467.] [‘Stenobothrus rubicundus (Germar)’; Yakobson & Bianki, 1902: 223, and other authors. Misinterpreta- tion of Germar (1817: 256) and misidentification of Gryllus (Locusta) rubicundus Goeze, 1778: 108.] Stenobothrus rubicundulus Kruseman & Jeekel, 1967: 79. [Replacement name for Gryllus miniatus Charpentier. | REFERENCES TO SONG. Oscillogram: Elsner, 1974b. Verbal description only: Chopard, 1952; Faber, 1953; Harz, 1957. RECOGNITION. The very broad fore wings, with unbranched M, regular ladder-like cross-veins in the medial area, and Cu, and Cu, quite clearly separated (Fig. 8), enable males of this species to be recognized quite easily. In the female these characters of the fore wing are not so clearly developed, but the hind wings of both sexes have an unbranched M and greatly enlarged medial area, and the more distal part of the anterior margin is heavily sclerotized and conspicuously dark in colour (Fig. 8). In the field males may be easily recognized by their highly characteristic calling song, which includes a loud buzzing sound produced by wing-vibration, either on the ground or in hovering flight. Sonc (Figs 103-116). The calling song is composed of two quite different kinds of sound: one is produced by the usual femoro-alary stridulation and the other by wing-vibration, either on the ground or in flight. The song sometimes consists of femoro-alary stridulation immediately followed by wing-vibration, but each kind of sound production may occur without the other. The femoro-alary stridulation (Figs 103-108) usually begins with a series of syllables in which the upstroke and downstroke of the hind legs produce clearly separated, simple sounds (Figs 105, 107); the syllable repetition rate is variable, but is usually 8—13/s. After a few seconds the pattern is often complicated by some of the downstrokes being prolonged and broken up by gaps (Figs 106, 108). This femoro-alary echeme usually lasts for about 6-10 s, after which it may be followed immediately by wing-vibration (Fig. 104). The wing-vibration produces a loud buzzing sound (Figs 109-116) usually lasting about 4-10 s. According to Elsner (19745) the sound is produced by the strongly sclerotized anterior margins of the two hind wings being beaten against each other. Oscillographic analysis shows the buzzing to consists of a series of sharply distinct short-lived sounds repeated at the rate of about 55-85/s (Figs 111-116). The whole of the wing-buzz may be produced while the male is sitting on the ground, but often the male takes off and continues to produce the buzzing sound in flight, often hovering above the ground (Fig. 110, second wing-buzz). In the presence of a female the male begins with femoro-alary syllables similar in kind to those of the calling song, but the echeme is often greatly prolonged, sometimes lasting for more than 30 minutes. This is followed by wing-vibration and there is then often an alternation of shorter echemes of femoro-alary stridulation and bursts of wing-vibration. A sequence of this kind may end in the male taking off during a burst of wing-vibration and finally an attempt to copulate with the female. DIsTRIBUTION. This montane species is known only from the Alps, higher Apennines and mountains of the Balkan Peninsula. References Andrieu, A.-J. & Dumortier, B. 1963. Chantes d’insectes. 30 cm disc, 33 rpm. Neuilly (Pacific). Baier, L. J. 1930. Contribution to the physiology of the stridulation and hearing of insects. Zoologische Jahrbiicher (Zoologie und Physiologie) 47: 151-248. Bei-Bienko, G. Ya. & Mishchenko, L. L. 1951. Locusts and grasshoppers of the U.S.S.R. and adjacent countries. Part II. Opredeliteli po Faune SSR, Izdavaemye Zoologicheskim Muzeem Akademii Nauk, Leningrad 40: 381-667. [In Russian. English translation published by Israel Program for Scientific Translations, Jerusalem, 1964, iv + 291 + xxi pp.] Bellmann, H. 1985a. Heuschrecken. Beobachten, bestimmen. 210 pp. Melsungen. SONGS OF THE WESTERN EUROPEAN Stenobothrus 423 — 1985b. Die Stimmen der Heimischen Heuschrecken. Tape cassette. Melsungen (Neumann- Neudamm). Bolivar, I. 1876. Sindpsis de los ort6pteros de Espana y Portugal. Segunda parte. Anales de la Sociedad Espanola de Historia Natural 5: 259-372. — 1887. Especies nuevas 6 criticas de ort6pteros. Anales de la Sociedad Espanola de Historia Natural 16: 90-114. Broughton, W. B. 1972. The grasshopper and the taxonomer. III. Keys to the species. Journal of Biological Education 6: 385-395. Cazurro y Ruiz, M. 1888. Enumeracién de los Ort6pteros de Espana y Portugal. Anales de la Sociedad Espanola de Historia Natural 17: 435-513. Charpentier, T. de 1825. Horae entomologicae, adjectis tabulis novem coloratis. xvi + 262 pp. Bratislava. Chopard, L. 1922. Orthoptéres et Dermaptéres. Faune de France 3, vi + 212 pp. — 1952. Orthoptéroides. Faune de France 56 (1951), 359 pp. Duijm, M. & Kruseman, G. 1983. De Krekels en Sprinkhanen in de Benelux. 186 pp. Amsterdam. Dumortier, B. 1963. The physical characteristics of sound emissions in Arthropoda (pp. 346-373). In Busnel, R.-G. [Ed.], Acoustic behaviour of Animals. xx + 933 pp. Amsterdam, London & New York. Ebner, R. 1915. Zur Kenntnis der Orthopterenfauna der Abruzzen. Deutsche Entomologische Zeitschrift 1915: 545-570. Elsner, N. 1974a. Neuroethology of sound production in gomphocerine grasshoppers (Orthoptera: Acrididae). I. Song patterns and stridulatory movements. Journal of Comparative Physiology 88: 67-102. — 1974b. Neural economy: bifunctional muscles and common central pattern elements in leg and wing stridulation of the grasshopper Stenobothrus rubicundus Germ. (Orthoptera: Acrididae). Journal of Comparative Physiology 89: 227-236. — 1975. Neuroethology of sound production in gomphocerine grasshoppers (Orthoptera: Acrididae). II. Neuromuscular activity underlying stridulation. Journal of Comparative Physiology 97: 291-322. Elsner, N. & Popov, A. V. 1978. Neuroethology of acoustic communication. Advances in Insect Physiology 13: 229-355. Eversmann, E. 1848. Additamenta quaedam levia ad Fischeri de Waldheim celeberrimi Orthoptera Rossica. 15 pp. Moscow. Faber, A. 1928. Die Bestimmung der deutschen Geradfliiger (Orthopteren) nach ihren Lautausserungen. Zeitschrift fiir Wissenschaftliche Insektenbiologie 23: 209-234. — 1953. Laut- und Gebdardensprache bei Insekten. Orthoptera (Geradfliigler). Teil 1. 198 pp. Stuttgart. Fischer, L. H. 1853. Orthoptera Europaea. xx + 454 pp. Leipzig. Germar, E. F. 1817. Reise nach Dalmatien und in das Gebiet von Ragusa. xii + 323 pp. Leipzig & Altenburg. Goeze, J. A. E. 1778. Entomologische Beytrage zu des Ritter Linné zwolften Ausgabe des Natursystems 2, Ixxii + 352 pp. Leipzig. Grein, G. 1984. Gesdnge der heimischen Heuschrecken. Akustisch-optische Bestimmungshilfe. 30 cm disc, 33 rpm [with oscillograms]. Hannover (Niedersachsisches Landesverwaltungsamt). Harz, K. 1957. Die Geradfliigler Mitteleuropas. xxiv + 494 pp. Jena. —— 1975. Die Orthopteren Europas II. Series Entomologica 11, [viii +] 939 pp. Haskell, P. T. 1955. Intensité sonore des stridulations de quelques Orthoptéres britanniques [pp. 154-167]. Jn Busnel, R.-G. [Ed.], Colloque sur l’Acoustique des Orthoptéres. Annales des Epiphyties (fascicule spécial de 1954), 448 pp. — 1957. Stridulation and associated behaviour in certain Orthoptera. 1. Analysis of the stridulation of, and behaviour between, males. British Journal of Animal Behaviour 5: 139-148. — 1958. Stridulation and associated behaviour in certain Orthoptera. 2. Stridulation of females and their behaviour with males. Animal Behaviour 6: 27-42. Herrich-Schaffer, [G. A. W.] 1840. Nomenclator entomologicus. Verzeichniss der europdischen Insecten zur Erleichterung des Tauschverkehrs mit Preisen versehen 2, viii + 244 pp. Regensburg. Jacobs, W. 1950. Vergleichende Verhaltensstudien an Feldheuschrecken. Zeitschrift fiir Tierpsychologie 7: 169-216. —— 1953. Verhaltensbiologische Studien an Feldheuschrecken. Zeitschrift fiir Tierpsychologie, Beiheft 1, vii + 228 pp. Kirby, W. F. 1910. A synonymic catalogue of Orthoptera. Vol. III. Orthoptera Saltatoria. Part II. (Locustid@ vel Acridiide). x + 674 pp. London. Kruseman, G. & Jeekel, C. A. W. 1967. Stenobothrus rubicundus (Germar, 1817): an invalid name (Orthoptera). Entomologische Berichten, Amsterdam 27: 78-80. 424 D. R. RAGGE La Greca, M. 1965. Su alcuni Blattodei, Mantodei ed Ortotteri della fauna italiana, nuovo o poco noti. Fragmenta Entomologica 4: 64-71. Luquet, G. C. 1978. La systématique des Acridiens Gomphocerinae du Mont Ventoux (Vaucluse) abordée par le biais du comportement acoustique [Orthoptera, Acrididae]. Annales de la Société Entomologique de France (N.S.) 14: 415-450. Pallas, P. S. 1771. Reise durch verschiedene Provinzen des russischen Reichs 1, [viii +] 504 pp. St. Petersburg. Panzer, G. W. F. 1796. Faunae insectorum Germanicae initia oder Deutschlands Insecten. Heft 33. 24 pp. Nurnberg. Ragge, D. R. 1965. Grasshoppers, crickets and cockroaches of the British Isles. xii + 299 pp. London & New York. —— 1986. The songs of the western European grasshoppers of the genus Omocestus in relation to their taxonomy (Orthoptera: Acrididae). Bulletin of the British Museum (Natural History) (Entomology) 53: 213-249. —— In press. Speciation and biogeography of some southern European Orthoptera, as revealed by their songs. Ragge, D. R., Burton, J. F. & Wade, G. F. 1965. Songs of the British grasshoppers and crickets. 17 cm disc, 33 rpm. London (Warne). Rambur, P. 1838-1842. Faune entomologique de l’Andalousie 2, 336 pp. Paris. Schmidt, G. H. & Schach, G. 1978. Biotopmassige Verteilung, Vergesellschaftung und Stridulation der Saltatorien in der Umgebung des Neusiedlersees. Zoologische Beitrage (N.F.) 24: 201-308. Tarbinskii, S. P. 1948. Order Saltatoria (Orthoptera) — jumping orthopterans [pp. 76-127]. In Tarbinskii, S. P. & Pavilshchikov, N. N. [Eds], Keys to the insects of the European USSR. 1128 pp. Moscow & Leningrad. [In Russian. ] Tsyplenkov, E. P. 1970. Harmful Acridoidea of the U.S.S.R. 272 pp. Leningrad. [In Russian. English translation published by Amerind Publishing Co., New Delhi, 1978, viii + 208 pp.] Yakobson, G. G. & Bianki, V. L. 1902-1905. Orthoptera and Pseudoneuroptera of the Russian Empire and neighbouring countries. x + 952 pp. Leningrad. [In Russian. ] Yersin, [A.] 1852. Mémoire sur quelques faits relatifs a la stridulation des Orthopteéres et a leur distribution géographique en Europe. Bulletins des Séances de la Société Vaudoise des Sciences Naturelles 4: 108-128. Index Invalid names are in italics; principal page references in bold. amoenus 400 apenninus 394, 397, 399, 400, 404, 418 berberus 400 bolivarii 394, 397, 399, 400, 402, 405, 410, 413 broelemanni 399 Bruneria 400 bulgaricus 400 burri 400 carbonarius 400 caucasicus 400 Chorthippus 399, 400 clavatus 399, 400 cobresianus 400 cotticus 397, 400 crassipes 400 croaticus 400 Crotalacris 399, 400 eurasius 399, 400 festivus 394, 397, 400, 402, 404, 405, 410, 418 fischeri 393, 394, 397, 399, 400, 402, 404, 405, 410, 418 graecus 400 grammicus 394, 397, 399, 400, 402, 405, 410, 413 kirgizorum 400 lineatus 393, 397, 400, 402, 404, 405 maroccanus 400 miniatus 422 miramae 400 Myrmeleotettix 400 newskii 400 nigromaculatus 393, 399, 400, 402, 404, 405, 418 Omocestus 397, 399, 400 palpalis 400 posthumus 400 raymondi 399 rubicundulus 394, 399, 400, 402, 404, 422 rubicundus 399, 422 Stauroderus 400 Stenobothrodes 399, 400 stigmaticus 394, 397, 399, 400, 402, 404, 405, 410, 413, 418 sviridenkoi 400 tadzhicus 400 ursulae 400 werneri 400 zubowskyi 400 British Museum (Natural History) Milkweed butterflies: their cladistics and biology P.R. Ackery & R. I. Vane-Wright The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of their biology have stimulated far more attention than can be justified by species numbers alone. In recent years, an expansive literature has grown, considering aspects of their courtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry and genetics. The popularity of danaines among biologists can certainly be attributed to this combination, within one small group, of so many of the factors that make butterflies such an interesting group to study. The obvious need to place this wealth of biological data within an acceptable systematic framework provided theimpetus for this volume. Started in 1976 within the conventions of evolution by natural selection and Hennig’s phylogenetic systematics, the book is now largely about natural history (what the animals have and do, where they live and how they develop) and natural groups — as revealed by a form of analysis approaching that practised by the new school of ‘transformed cladistics’. The authors have prepared a handbook that will appeal to a wide range of biologists, from museum taxonomists to field ecologists. 425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. ISBN 0 565 00893 5. 1984. Price £50. Titles published in Volume 55 A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of certain genera By M. W. R. de V. Graham The songs of the western European grasshoppers of the genus Stenobothrus in relation to their taxonomy (Orthoptera: Acrididae) By D. R. Ragge Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester