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F:\REJ\16-3\265-272 (Tishechkin) Russian Entomol. J. 16(3): 265–272 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2007 Similar calling signals in different species of leafhoppers (Homoptera: Cicadellidae): an example of Paralimnini Ñõîäíûå ïðèçûâíûå ñèãíàëû ó ðàçíûõ âèäîâ öèêàäîê (Homoptera: Cicadellidae): èññëåäîâàíèå íà ïðèìåðå Paralimnini D.Yu. Tishechkin Ä.Þ. Òèøå÷êèí Department of Entomology, Faculty of Biology, M.V. Lomonosov Moscow State University, Vorobyevy Gory, Moscow 119992, Russia. E- mail: [email protected]. Кафедра энтомологии, Биологический факультет, Московский государственный университет им. М.В. Ломоносова, Воробьевы Горы, Москва, 119992 Россия. KEY WORDS: leafhoppers, taxonomy, diagnostic characters, vibratory signals, Paralimnini, Cicadellidae. КЛЮЧЕВЫЕ СЛОВА: цикадовые, систематика, диагностические признаки, вибрационные сигналы, Paralimnini, Cicadellidae. ABSTRACT. Examples of similarity of calling sig- patterns in different forms does not always indicates nals in different species of Paralimnini (Homoptera: that they belong to the same species, however. Cicadellidae) are discussed. Species producing signals For example, in certain grasshoppers, e.g., in Eu- with identical structure never come into acoustic inter- ropean Euchorthippus and in a number of Omocestus actions with each other due to differences in host spe- species (Orthoptera: Acrididae) temporal pattern of cialisation, habitat preferences or geographical distri- calling signals is quite similar and sometimes is al- bution. Therefore, using of bioacoustic characters in most identical [Ragge & Reynolds, 1984; Ragge, systematics without considering the data on biology 1986]. Recent investigation of calling songs in grass- and distribution can lead to wrong taxonomic conclu- hopper communities demonstrated that the complex sions, because the similarity or even absolute identity of temporal parameters of signal determines for each of structure of signals in different forms is not always species its own place in the acoustic environment of unequivocal evidence of their synonymy. the community, so-called acoustic niche, which is a part of the ecological niche as a whole [Bukhvalova, РЕЗЮМЕ. Рассмотрены примеры сходства при- 2006]. Signals of species belonging to the same com- зывных сигналов у разных видов Paralimnini (Homo- munity differ from each other, i.e. occupy different ptera: Cicadellidae). Во всех случаях виды, издаю- acoustic niches, whereas in members of different com- щие идентичные по структуре сигналы, не вступа- munities they sometimes are almost indistinguishable ют в акустический контакт по причине различий в in temporal pattern. This is no barrier to successful кормовой специализации, биотопической приуро- communication because such species never come into ченности или географическом распространении. Та- acoustic interactions with each other. ким образом, использование биоакустических при- Similar situation takes place in Psyllinea (Ho- знаков в систематике без учёта данных по распрос- moptera). Temporal pattern of calling songs in this транению и биологии может привести к ошибоч- group for the most part is species-specific. Nonethe- ным таксономическим заключениям, поскольку less, in a number of species signals are quite similar сходство или даже полная идентичность сигналов and cannot be told apart with certainty [Tishechkin, разных форм отнюдь не всегда является доказа- 2006]. Occasionally, such species are formally sym- тельством их синонимии. patric, i.e. live in the same biotope. Still, it should be remembered, that psyllids use not airborne sounds, It is well known that these are differences in tem- but substrate-borne vibrations for their communica- poral pattern of calling signals, which provide repro- tion. Such signals cannot be transmitted from one ductive isolation in many species of insects using plant to another without physical contact. Consequent- acoustic communication. For this reason, analysis of ly, forms with narrow host specialization dwelling on signals is successfully employed in taxonomy for different plant species as a rule cannot hear the signals discrimination between cryptic species and for estab- of each other. lishing of status of closely related forms showing Small Auchenorrhyncha (Homoptera: Cicadinea small morphological differences. Similarity of signal with the exception of singing cicadas, Cicadidae) also 266 D.Yu. Tishechkin produce substrate-borne vibratory signals. Although a used. Analysis of recordings was performed on PC great number of representatives of many orders of provided with analog/digital converter and appropri- insects possess well-developed acoustic communica- ate software. tion, signals of Cicadinea stands out because of most Data for recordings of signals of leafhoppers used elaborate temporal structure. Even in closely related in the paper are given in the Table. species the pattern of calling songs sometimes is com- pletely different. Thus, in many taxa of Cicadinea It is apparent, that in allopatric species differences acoustic analysis provides useful characters for dis- in structure of communication signals plays no role in crimination between cryptic species [for example, reproductive isolation. Consequently, such forms can Tishechkin, 1999a, b, 2002]. By contrast, in certain produce similar signals and still exist as separate tribes of leafhoppers (Cicadellidae) and in some Fulgor- species. In certain Paralimnini that is the case. oidea the structure of signals is rather simple and uniform. Differences between signals of closely relat- Sorhoanus medius (Mulsant et Rey, 1855) and S. ed species in such groups are indistinct or absent. hilaris (Melichar, 1900) are common and abundant Therefore, signal analysis for taxonomic purposes steppe-dwelling species feeding on grasses (Poaceae). must be used with caution in this situation. They inhabit similar biotopes, but the range of the Certain examples of similarity of calling signals in former one includes Europe, Kazakhstan and southern the representatives of the tribe Paralimnini (Homoptera: part of Western Siberia, whereas the latter occurs in Cicadellidae: Deltocephalinae) will be discussed here. steppes of Eastern Siberia (Irkutsk Area, Transbaikalia, Yakutia) and in Central and Eastern Mongolia. In both Recordings of vibratory signals were made by species, calling signals consist of single or regularly means of piezo-electric crystal gramophone cartridge repeating short fragments with variable shape (Figs 1– connected to the microphone input of cassette record- 10 and 11–16). Occasionally, fragments of quite differ- er “Elektronika–302–1” or minidisk recorder Sony ent shape present in the signals of specimens from the Walkman MZ–NH900 via the matching amplifier. In same population (Fig. 1). Inner structure of signals also all cases manual mode of recording level control was varies greatly. Signal consists either of sine waves (Figs Data for recordings of calling signals of the studied species of Paralimnini Äàííûå î çàïèñÿõ ïðèçûâíûõ ñèãíàëîâ èçó÷åííûõ âèäîâ Paralimnini Similar signals in different species of leafhoppers 267 1 4 2 3 2 s 2 200 3 ms 200 4 ms 5 200 ms 5 20 ms 6 200 7 ms 200 8 ms 200 9 ms 9 20 ms 10 200 ms 11 2 s 12 12 200 13 ms 200 14 ms 200 15 ms 200 ms 16 200 ms Figs 1–16. Oscillograms of calling signals of males: 1–10 — Sorhoanus medius; 1–5 — from Saratov Area; 6–9 — from Moscow Area; 10 — from Altai; 11–16 — S. hilaris; 11–15 — from Buryatia; 16 — from Chita Area. Parts of signals indicated as 2–5, 9 and 12 are given on oscillograms under the same numbers. Ðèñ. 1–16. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ ñàìöîâ: 1–10 — Sorhoanus medius; 1–5 — èç Ñàðàòîâñêîé îáë.; 6–9 — èç Ìîñêîâñêîé îáë.; 10 — ñ Àëòàÿ; 11–16 — S. hilaris; 11–15 — èç Áóðÿòèè; 16 — èç ×èòèíñêîé îáë. Ôðàãìåíòû ñèãíàëîâ, îáîçíà÷åííûå öèôðàìè 2–5, 9 è 12, ïðåäñòàâëåíû íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 268 D.Yu. Tishechkin 17 2 s 18 20 18 200 19 ms 200 20 ms 200 21 ms 200 ms 22 200 ms 23 200 24 ms 200 ms 25 2 s 26 26 200 27 ms 200 ms 28 29 2 s 29 200 30 ms 31 2 s 32 33 31 50 ms 32 50 ms 33 50 ms Figs 17–33. Oscillograms of calling signals: 17–21 — Mocuellus collinus; 22–24 — M. angustiarum; 25–27 — Psammotettix atropidicola; 28–33 — P. kaszabi. Parts of signals indicated as 18, 20, 26, 29 and 31–33 are given on oscillograms under the same numbers. Ðèñ. 17–33. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ: 17–21 —Mocuellus collinus; 22–24 — M. angustiarum; 25–27 — Psammotettix atropidicola; 28–33 — P. kaszabi. Ôðàãìåíòû ñèãíàëîâ, îáîçíà÷åííûå öèôðàìè 18, 20, 26, 29 è 31–33, ïðåäñòàâëåíû íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Similar signals in different species of leafhoppers 269 4–5), or of vibrations with more complex and less the same male are presented on oscillograms) as well regular shape (Figs 8–9). Evidently, variability of waves as the outline of syllables and their duration (Figs 34– shape is a result of different transmission properties of 35, 37 and 39–46) provide no reliable diagnostic the parts of twigs or stems and of differences in relative characters. All species are partly sympatric. The rang- position of singing insect and vibrotransducer detect- es of D.bohemani and D. frauenfeldi include Europe, ing vibrational signal. So, as is seen from the oscillo- Kazakhstan and Southern Siberia (possibly, with the grams on Figs 1–16, no clear-cut distinction exists exception of Eastern Transbaikalia). D. suttholli in- between calling signals of these two species. habits steppes of Mongolia and Southern Siberia (Tyva, Buryatia). These species were never found in the same Mocuellus
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