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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">2</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">ZooKeys</journal-title>
        <abbrev-journal-title xml:lang="en">ZK</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1313-2989</issn>
      <issn pub-type="epub">1313-2970</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zookeys.1200.118422</article-id>
      <article-id pub-id-type="publisher-id">118422</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Acrididae</subject>
          <subject>Orthoptera</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Ecology &amp; Environmental sciences</subject>
          <subject>Systematics</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Cenozoic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Asia</subject>
          <subject>Europe</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿New data on bioacoustics and courtship behaviour in grasshoppers (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Orthoptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Acrididae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name>) from Russia and adjacent countries</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Vedenina</surname>
            <given-names>Varvara</given-names>
          </name>
          <email xlink:type="simple">vvedenina@googlemail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-2694-4152</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Sevastianov</surname>
            <given-names>Nikita</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-1563-5194</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Kovalyova</surname>
            <given-names>Evgenia</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Institute for Information Transmission Problems, Russian Academy of Sciences, Bolshoy Karetny per. 19, Moscow 127051, Russia</addr-line>
        <institution>Institute for Information Transmission Problems, Russian Academy of Sciences</institution>
        <addr-line content-type="city">Moscow</addr-line>
        <country>Russia</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Varvara Vedenina (<email xlink:type="simple">vvedenina@googlemail.com)</email></p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Jun-Jie Gu</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2024</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>02</day>
        <month>05</month>
        <year>2024</year>
      </pub-date>
      <volume>1200</volume>
      <fpage>1</fpage>
      <lpage>26</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/06FC1BC1-1DF0-58A1-9095-9C5F8A889450">06FC1BC1-1DF0-58A1-9095-9C5F8A889450</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/4F62E5E1-95BA-4F1C-99E2-765E6340EFA9">4F62E5E1-95BA-4F1C-99E2-765E6340EFA9</uri>
      <history>
        <date date-type="received">
          <day>08</day>
          <month>01</month>
          <year>2024</year>
        </date>
        <date date-type="accepted">
          <day>31</day>
          <month>03</month>
          <year>2024</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Varvara Vedenina, Nikita Sevastianov, Evgenia Kovalyova</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <self-uri content-type="zoobank" xlink:type="simple">http://zoobank.org/4F62E5E1-95BA-4F1C-99E2-765E6340EFA9</self-uri>
      <abstract>
        <label>﻿Abstract</label>
        <p>The songs of seven grasshopper species of subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name> from Russia, Ukraine, Georgia, and Kazakhstan were studied. We analysed not only the sound, but also the stridulatory movements of the hind legs to more entirely describe the songs. In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">Mesasippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic>, the legs are moved in a relatively simple pattern; four other species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> demonstrate more complex leg movements. In six of the seven species studied, the courtship songs contain more sound elements than the calling songs. The highest number of courtship sound elements was found in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus.</tp:taxon-name-part></tp:taxon-name></italic> The different parts of a remarkably long stridulatory file in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> are thought to participate in the production of different sound elements. The songs in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic> are shown for the first time. This species produces sound not only by common stridulation but also by wing beats. A relationship of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic> to some other species of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part></tp:taxon-name></italic>, which are able to crepitate, is discussed. We also analyse the frequency spectra of the songs. A maximum energy of the song power spectra in six species studied lies in ultrasound range (higher than 20 kHz). In only <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>, the main peaks in the song power spectra lie lower than 20 kHz. The courtship behaviour in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic> was shown to include conspicuous visual display (movements of antennae, palps and the whole body).</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>Calling song</kwd>
        <kwd>courtship song</kwd>
        <kwd>frequency spectrum</kwd>
        <kwd>stridulatory leg movements</kwd>
        <kwd>visual display</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>Russian Scientific Foundation (grant 23-24-00533)</funding-statement>
      </funding-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0EBCAC">
        <title>Citation</title>
        <p>Vedenina V, Sevastianov N, Kovalyova E (2024) New data on bioacoustics and courtship behaviour in grasshoppers (Orthoptera, Acrididae, Gomphocerinae) from Russia and adjacent countries. ZooKeys 1200: 1–26. <ext-link xlink:href="10.3897/zookeys.1200.118422" ext-link-type="doi" xlink:type="simple">https://doi.org/10.3897/zookeys.1200.118422</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0EADAC">
      <title>﻿Introduction</title>
      <p>In many species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Orthoptera</tp:taxon-name-part></tp:taxon-name>, the song is an important component of reproductive isolation. This is the reason why acoustic signals are often used in taxonomy, when sibling species are similar in morphology, but have quite different songs. Among the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Acrididae</tp:taxon-name-part></tp:taxon-name> subfamilies, acoustic communication in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name> is most developed in terms of structure of acoustic apparatus, temporal pattern of the song, and mating strategies (e.g., <xref ref-type="bibr" rid="B18">Otte 1970</xref>; <xref ref-type="bibr" rid="B33">von Helversen and von Helversen 1994</xref>; <xref ref-type="bibr" rid="B19">Ragge and Reynolds 1998</xref>). The song is produced by stroking a stridulatory file on each hind femur across a raised vein on the fore wing. Using both hind legs, the grasshoppers have two separate sound-producing devices that must be coordinated with one another. The stridulatory movements of the two legs often differ in amplitude and form, and the legs can exchange roles from time to time, leading to increased song complexity (<xref ref-type="bibr" rid="B9">Elsner 1974</xref>; <xref ref-type="bibr" rid="B32">von Helversen and Elsner 1977</xref>; <xref ref-type="bibr" rid="B10">Elsner 1994</xref>). Various species demonstrate different degrees of song complexity. The song in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name> also varies according to the behavioural situation. A solitary male produces a calling song, listening for the response song of a female that is ready to mate. Several males sitting in a close vicinity can produce rival songs. When a male finds a female, in many species the male begins a special courtship song, which may reach a high complexity and may be accompanied by conspicuous movements of different parts of the body such as the abdomen, head, antennae, or palps (<xref ref-type="bibr" rid="B12">Faber 1953</xref>; <xref ref-type="bibr" rid="B18">Otte 1970</xref>; <xref ref-type="bibr" rid="B33">von Helversen and von Helversen 1994</xref>).</p>
      <p>To make a comprehensive analysis of songs between the species, it is necessary to compare not only the sound but also the stridulatory leg movements. Sometimes, a similar sound pattern can be produced by completely different leg movements (<xref ref-type="bibr" rid="B27">Vedenina and Mugue 2011</xref>). The leg movement analysis may help in the sound analysis when the gaps between sound elements are not distinct because of the phase shift between the two legs. A comparison of the leg movements in different species rather than the sound analysis may indicate a relationship between the species (<xref ref-type="bibr" rid="B20">Sevastianov et al. 2023</xref>). During the courtship behaviour, a male may also demonstrate species-specific leg movements without producing the sound.</p>
      <p>It was previously argued that the specificity of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name> songs lies not in their frequency band but almost without exception in the pattern of amplitude over time. However, several studies showed that despite a relatively broad spectra of the grasshopper songs, there are pronounced interspecific differences in maxima or peaks (Meyer and Elsner 1996, <xref ref-type="bibr" rid="B16">1997</xref>). It was also shown that male calling and female response songs may differ in the frequency spectra, and these differences can be used during species recognition (<xref ref-type="bibr" rid="B34">von Helversen and von Helversen 1997</xref>). It was also shown that various parts of elaborate courtship songs may significantly differ in the carrier frequency (<xref ref-type="bibr" rid="B30">Vedenina et al. 2007</xref>; <xref ref-type="bibr" rid="B17">Ostrowski et al. 2009</xref>; <xref ref-type="bibr" rid="B31">Vedenina et al. 2020</xref>). The differences in the frequency spectra between the various song elements may influence the amplitude ratio on the oscillogram. If the song is recorded by portable recorders with a frequency range not exceeding 12.5–15 kHz, the amplitude ratio of different elements may be distorted (<xref ref-type="bibr" rid="B28">Vedenina and Shestakov 2014</xref>).</p>
      <p>In the current paper, we describe the calling and courtship songs in seven species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name> from Russia, Ukraine, Kazakhstan, and Georgia. To gain a better description of the songs, we analyse not only the sound, but also the underlying stridulatory movements of the hind legs. We also consider the whole visual display accompanying the courtship song in some species. And finally, we analyse the frequency spectra of the songs and different song elements.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EOGAC">
      <title>﻿Materials and methods</title>
      <p>The calling song was recorded from a solitary male; the courtship song was recorded when a male was sitting near a female. All song recordings were made in the laboratory. Both the sound and the movements of the hind legs were recorded with a custom-built opto-electronic device (<xref ref-type="bibr" rid="B32">von Helversen and Elsner 1977</xref>; <xref ref-type="bibr" rid="B13">Hedwig 2000</xref>). A piece of reflecting foil was glued to the distal part of each hind leg femur of the male and two opto-electronic cameras were focused on the illuminated reflecting dots. Each camera was equipped with a position-sensitive photodiode that converted the upward and downward movements of the hind legs into voltage signals. These signals, together with the recordings of the sounds (a microphone type 4191, ½ inch; a conditioning amplifier type 2690; Brüel &amp; Kjaer, Nærum, Denmark), were A/D-converted with a custom-built PC card. The sampling rate was 1325 Hz for recording the stridulatory movements and 100 kHz for sound recordings. The ambient temperature near the singing male was 30–32 °C. The temporal parameters and power spectra of the songs were analysed with COOLEDIT (Syntrillium, Seattle, WA) and TURBOLAB 4.0 (Bressner Technology, Gröbenzell, Germany). Courtship behaviour was also recorded with a Sony HDR-CX 260E digital video camera; the video signals were analysed with the VIRTUAL DUB program.</p>
      <p>Localities where the song recordings were made are shown in Fig. <xref ref-type="fig" rid="F1">1</xref>. The numbers of localities in the text (paragraphs Material in description of each species) correspond to the numbers on the map. Data on species distribution were obtained from <xref ref-type="bibr" rid="B6">Bey-Bienko and Mistshenko (1951)</xref> and <xref ref-type="bibr" rid="B19">Ragge and Reynolds (1998)</xref>.</p>
      <fig id="F1" position="float" orientation="portrait">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.figure1</object-id>
        <object-id content-type="arpha">10569CA6-2F90-531C-8975-331147997AC3</object-id>
        <label>Figure 1.</label>
        <caption>
          <p>Map of localities where the specimens were collected for the song recordings. 1–6: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>; 7, 8: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic>; 9: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">Mesasippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part></tp:taxon-name></italic>; 10–13: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>; 14, 15 <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic>; 16: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>; 17, 18: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii.</tp:taxon-name-part></tp:taxon-name></italic> The localities of the same species are indicated by the same colour.</p>
        </caption>
        <graphic xlink:href="zookeys-1200-001_article-118422__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040930.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1040930</uri>
        </graphic>
      </fig>
      <p>For the song description we used the following terms: <bold>pulse</bold> – the sound produced by one stroke of a hind leg and representing the shortest measurable unit; <bold>syllable</bold> – the sound produced by one complete up and down movement of the hind legs, starting when the legs leave their initial position and ending when the legs return to their original position; <bold>element</bold> – the sound produced by the same leg movements and usually including a series of equal syllables; <bold>echeme</bold> – series of consistent syllables separated by pauses (Fig. <xref ref-type="fig" rid="F2">2</xref>).</p>
      <fig id="F2" position="float" orientation="portrait">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.figure2</object-id>
        <object-id content-type="arpha">9C0F4DBA-9504-5969-A74B-3499ECEDDA25</object-id>
        <label>Figure 2.</label>
        <caption>
          <p>Oscillograms of the calling song <bold>A, B</bold>, courtship songs <bold>C–G</bold> and frequency spectra <bold>H–J</bold> in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>. Courtship songs of two males are shown in <bold>C</bold> and <bold>D</bold>. Song recordings are presented at three different speeds. In oscillograms <bold>C–G</bold> the two upper lines are recordings of hind leg movements and the lower line is the sound recording. Different elements of the courtship song are shown in <bold>E, F</bold>. Frequency spectra are shown in kHz for the courtship elements 1 <bold>H</bold>, 2 <bold>I</bold> and 3 <bold>J</bold>.</p>
        </caption>
        <graphic xlink:href="zookeys-1200-001_article-118422__-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040931.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1040931</uri>
        </graphic>
      </fig>
    </sec>
    <sec sec-type="﻿Results and discussion" id="SECID0E5LAC">
      <title>﻿Results and discussion</title>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Orthoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Acrididae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">FE0D4EB4-6C99-5797-B76C-F3C391F3ED21</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>(Zubowsky, 1900)</tp:taxon-authority>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="distribution" id="SECID0EGNAC">
          <title>Distribution.</title>
          <p>Southern Siberia from Altai to Transbaikalia, south-west of Amur region to Mongolia. Abundant in dry steppes and semi-deserts.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0ELNAC">
          <title>Material</title>
          <p><bold>(Fig. <xref ref-type="fig" rid="F1">1</xref>).</bold> 16. Russia: Altai Republic, ~ 26 km SE of Ongudai, environs of Kupchegen’, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[86.436667,50.621667]}" id="NCID0E2NAC">50°37.3'N, 86°26.2'E</named-content></named-content>, 922 m a.s.l., 05.08.2023, song recordings in 3 ♂.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="References to song" id="SECID0EAOAC">
          <title>References to song.</title>
          <p><xref ref-type="bibr" rid="B24">Tishechkin and Bukhvalova 2009</xref>: recordings of calling song from Buryatia and Chita region.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Song" id="SECID0EKOAC">
          <title>Song.</title>
          <p>The calling song is an echeme-sequence lasting for ~ 7 s and consisting of ~ 14 echemes (Fig. <xref ref-type="fig" rid="F2">2A–B</xref>). The echemes usually lasts ~ 0.2–0.25 s and the intervals between them are ~ 0.3–0.4 s. Each echeme consists of ~ 25 pulses repeated at the rate of 120–125 /s. Oscillograph analysis shows that the low-amplitude pulses are sometimes produced in gaps between the main pulses.</p>
          <p>In the courtship song, one can distinguish three sound elements (Fig. <xref ref-type="fig" rid="F2">2C–J</xref>; see Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>). The element 1 is similar to the calling echeme, although with clear gaps between pulses. The pulses are produced by synchronous leg movements repeated at the rate of ~ 115–120 /s. Each pulse is generated by only downstroke. The element 1 gradually transforms to element 2: the rate of pulses decreases to ~ 40–50 /s and they become ~ 2 × as long. It is remarkable that they are produced by very weak up-movement of one leg. After three alternations of elements 1 and 2, a male produces a high-amplitude stroke with both legs, despite two legs produce different patterns. One leg is moved up in a stepwise manner, which results to generation of 3–7 low-amplitude pulses (element 3), whereas another leg is moved up straighter without low-amplitude vibrations. Then elements 1 and 2 alternate again 3×, followed by the high-amplitude stroke of the legs, which change the roles. After repeating ~ 6–7 cycles with high-amplitude strokes, the two legs are moved synchronously producing element 3, followed by precopulatory leg movements. The frequency spectra of elements 1 and 2 are similar, occupying a broad range from 15 to 40 kHz with maximum energy between 25 and 35 kHz (Fig. <xref ref-type="fig" rid="F2">2H, I</xref>). The spectrum of element 3 has three maxima at ~ 15, 30, and 40 kHz (Fig. <xref ref-type="fig" rid="F2">2J</xref>).</p>
          <p>When producing alternation of elements 1 and 2, a male slightly moves his body from side to side, generates low-amplitude movements with antennae keeping them turned to the sides, and conspicuously moves with black and white palps up and down (Fig. <xref ref-type="fig" rid="F3">3</xref>).</p>
          <fig id="F3" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure3</object-id>
            <object-id content-type="arpha">0B23D50A-4B01-58C3-9D57-BA7E610FFEA1</object-id>
            <label>Figure 3.</label>
            <caption>
              <p>Movements of antennae, black and white palps and the whole body in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> during courtship. The palps are raised up <bold>A</bold> lowered down <bold>B</bold> and raised up again <bold>C</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040932.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040932</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Comparative remarks" id="SECID0ELAAE">
          <title>Comparative remarks.</title>
          <p>The recordings of calling song in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> from Altai are similar to the recordings from Buryatia and Chita region (<xref ref-type="bibr" rid="B24">Tishechkin and Bukhvalova 2009</xref>). The courtship song of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> is shown for the first time. The calling song and the courtship element 1 are produced during the leg movements repeated at the rate of ~ 115–120 /s. According to <xref ref-type="bibr" rid="B27">Vedenina and Mugue (2011)</xref> and to <xref ref-type="bibr" rid="B20">Sevastianov et al. (2023)</xref>, this pattern can be considered as relatively complex and advanced pattern: the leg-movement rate could originate from the double rates of the wing beat. The complex structure of the courtship song with several sound elements found in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> are concordant with the overall complexity of the courtship songs in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part></tp:taxon-name></italic>. (<xref ref-type="bibr" rid="B19">Ragge and Reynolds 1998</xref>; <xref ref-type="bibr" rid="B4">Berger and Gottsberger 2010</xref>; <xref ref-type="bibr" rid="B27">Vedenina and Mugue 2011</xref>; <xref ref-type="bibr" rid="B31">Vedenina et al. 2020</xref>).</p>
          <p>We suggest the movements with palps to be a remarkable visual display that distinguishes <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> from most gomphocerine species. In one more species of this genus, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="antennatus">antennatus</tp:taxon-name-part></tp:taxon-name></italic>, the palp movements were also described during courtship (<xref ref-type="bibr" rid="B4">Berger and Gottsberger 2010</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="antennatus">antennatus</tp:taxon-name-part></tp:taxon-name></italic>, however, moves palps much more rapidly than <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>, and the most conspicuous visual display in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="antennatus">antennatus</tp:taxon-name-part></tp:taxon-name></italic> comprises a large swing of antennae with well-developed clubs. In contrast to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>, pulps in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="antennatus">antennatus</tp:taxon-name-part></tp:taxon-name></italic> are not coloured in black and white. Within the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part></tp:taxon-name></italic>, antennae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> are least thickened at the ends, which is probably correlated with very weak antennal movements during courtship. Other two species of this genus, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maculatus">maculatus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B19">Ragge and Reynolds 1998</xref>; <xref ref-type="bibr" rid="B28">Vedenina and Shestakov 2014</xref>) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pallidus">pallidus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B31">Vedenina et al. 2020</xref>) move conspicuously with antennae but not with palps. Thus, the visual display in various <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part></tp:taxon-name></italic> species seems to evolve independently. The palp movements are also known in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Aeropus">Aeropus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sibiricus">sibiricus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gomphocerippus">Gomphocerippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="rufus">rufus</tp:taxon-name-part></tp:taxon-name></italic> (e.g., <xref ref-type="bibr" rid="B9">Elsner 1974</xref>; <xref ref-type="bibr" rid="B19">Ragge and Reynolds 1998</xref>). However, these species that are distantly related to the species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part></tp:taxon-name></italic>, demonstrate very different patterns of the palp movements. It is evident that the pulp movements in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Aeropus">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sibiricus">sibiricus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gomphocerippus">G.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="rufus">rufus</tp:taxon-name-part></tp:taxon-name></italic> evolved convergently.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Orthoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Acrididae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">3FFF7DCA-F3EF-58E3-AE01-1222D92A2B0D</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>Zubowsky, 1900</tp:taxon-authority>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="distribution" id="SECID0EJKAE">
          <title>Distribution.</title>
          <p>Altai Mountains, Tuva, NW Mongolia. Usually associated with alpine meadows.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0EOKAE">
          <title>Material</title>
          <p><bold>(Fig. <xref ref-type="fig" rid="F1">1</xref>).</bold> Russia: 17. Altai Republic, Ulagan district, 3.5 km N of Lake Cheybek-Kohl, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[87.576017,50.430900]}" id="NCID0E5KAE">50°25.854'N, 87°34.561'E</named-content></named-content>, 1907 m a.s.l., 06.08.2023, song recordings in 3 ♂; 18. Altai Republic, Ulagan district, ab. 10 km N of Aktash, near Lake Cheybek-Kohl, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[87.596667,50.408333]}" id="NCID0EGLAE">50°24.5'N, 87°35.8'E</named-content></named-content>, 1821 m a.s.l., 14.08.2021, song recordings in 5 ♂, 06.08.2023, song recordings in 3 ♂.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="References to song" id="SECID0ELLAE">
          <title>References to song.</title>
          <p>Unknown.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Song" id="SECID0EQLAE">
          <title>Song.</title>
          <p>The calling song is an echeme-sequence that may last for tens of seconds, up to a minute (Fig. <xref ref-type="fig" rid="F4">4A–C</xref>). The echemes usually lasts ~ 0.25 s and the intervals between them are ~ 0.6 s. Each echeme begins quietly, reaching maximum intensity at the second half of its duration. Each echeme is generated by the low-amplitude, antidromic up and down leg movements at the rate of ~ 120 /s. During both up and down movements, the legs produce distinct pulses so, that the pulse rate is 2× as high as the leg-movement rate.</p>
          <fig id="F4" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure4</object-id>
            <object-id content-type="arpha">4C9913AF-1BC7-58B4-A77A-70EA4527BCAD</object-id>
            <label>Figure 4.</label>
            <caption>
              <p>Oscillograms of the calling song <bold>A–C</bold> courtship song <bold>D–G</bold> and frequency spectra <bold>H, I</bold> in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic>. Courtship songs of two males are shown in <bold>E</bold> and <bold>F</bold>. Song recordings are presented at three different speeds. In oscillograms <bold>A–C, F–G</bold> the two upper lines are recordings of hind leg movements and the lower line is the sound recording. The drawing shows the wing clapping, which generates the high-amplitude pulses shown in <bold>D–G</bold>. Frequency spectra are shown in kHz for the main echeme <bold>H</bold> and wing beats <bold>I</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040933.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040933</uri>
            </graphic>
          </fig>
          <p>During courtship, the males generate a sequence of echemes almost identical to the calling echeme-sequence. However, sometimes a courting male shortly crepitates by wings (Fig. <xref ref-type="fig" rid="F4">4D–G</xref>; see Suppl. material <xref ref-type="supplementary-material" rid="S2">2</xref>) or / and starts a noisy flight. After such a flight, the male is trying to copulate. The frequency spectrum of crepitation has a rather narrow maximum around 20 kHz; by contrast, the spectrum of echeme is more usual for <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name>, ranging from 10 to 40 kHz with numerous maxima between 18 and 30 kHz.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Comparative remarks" id="SECID0ETNAE">
          <title>Comparative remarks.</title>
          <p>The acoustic behaviour in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic> is described for the first time. It is remarkable that this species crepitates in flight and is also able to generate short sequences of wing beats sitting on the ground. Such crepitation on the ground is also known in some other species of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part></tp:taxon-name></italic>, namely, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="rubicundulus">rubicundulus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B11">Elsner and Wasser 1995</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cotticus">cotticus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B5">Berger et al. 2010</xref>), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hyalosuperficies">hyalosuperficies</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B22">Tarasova et al. 2021</xref>).</p>
          <p>The song and mating behaviour of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic> is almost identical to those in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cotticus">cotticus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B19">Ragge and Reynolds 1998</xref>; <xref ref-type="bibr" rid="B3">Berger 2008</xref>; <xref ref-type="bibr" rid="B5">Berger et al. 2010</xref>). It is surprising considering the large distance between the localities of the two species. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cotticus">cotticus</tp:taxon-name-part></tp:taxon-name></italic> was originally assumed to be endemic to the southwestern Alps, and it was later found in the Rila mountains in Bulgaria (<xref ref-type="bibr" rid="B5">Berger et al. 2010</xref>). The authors suggested that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cotticus">cotticus</tp:taxon-name-part></tp:taxon-name></italic> had a wider distribution during colder periods, when its range was expanded to lower altitudes. It is remarkable that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic> is similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cotticus">cotticus</tp:taxon-name-part></tp:taxon-name></italic> not only in song but also in morphology (<xref ref-type="bibr" rid="B36">Zubowsky 1900</xref>; <xref ref-type="bibr" rid="B14">Kruseman and Jeekel 1967</xref>) and ecological preferences: both species occur in alpine meadows at altitudes higher than 1700 m a.s.l. Taking into account that the leg-movement patterns of both species are relatively complex and may be considered as evolutionary advanced (<xref ref-type="bibr" rid="B27">Vedenina and Mugue 2011</xref>; <xref ref-type="bibr" rid="B20">Sevastianov et al. 2023</xref>), we suppose that the two species may represent one taxon. However, this conclusion requires more confirmation.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Orthoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Acrididae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">665CEF67-C888-5577-BAD7-AB9A962A66B3</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">Mesasippus</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>Tarbinski, 1925</tp:taxon-authority>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="distribution" id="SECID0EAUAE">
          <title>Distribution.</title>
          <p>Eastern and southern Kazakhstan, Uzbekistan.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0EFUAE">
          <title>Material</title>
          <p><bold>(Fig. <xref ref-type="fig" rid="F1">1</xref>).</bold> 9. Kazakhstan: Almaty region, national park Altyn-Emel’, environs of Basshi, along stream, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[78.751667,44.168333]}" id="NCID0EVUAE">44°10.1'N, 78°45.1'E</named-content></named-content>, 05.07.2016, song recordings in 2 ♂, 22.06.2023, song recordings in 3 ♂.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="References to song" id="SECID0E1UAE">
          <title>References to song.</title>
          <p><xref ref-type="bibr" rid="B7">Bukhvalova and Vedenina 1998</xref>: recordings of calling song from Kazakhstan.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Song" id="SECID0EEVAE">
          <title>Song.</title>
          <p>The calling song is an echeme lasting for 3–4 s (Fig. <xref ref-type="fig" rid="F5">5</xref>). It begins quietly and reaches maximum intensity ~ 2/3 of its duration (Fig. <xref ref-type="fig" rid="F5">5A–B</xref>). The two legs are moved synchronously at the rate of 21–25 /s, generating homogenic syllables. During the upstroke, the legs generate a soft sound, whereas during the downstroke, the sound intensity gradually increases (Fig. <xref ref-type="fig" rid="F5">5C</xref>).</p>
          <fig id="F5" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure5</object-id>
            <object-id content-type="arpha">E3B35129-353C-5C4E-BF3D-49E547150E58</object-id>
            <label>Figure 5.</label>
            <caption>
              <p>Oscillograms of the calling song <bold>A, B</bold> courtship song <bold>C–H</bold> and frequency spectra <bold>I, J</bold> in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">Mesasippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part></tp:taxon-name></italic>. Courtship songs of two males are shown in <bold>D</bold> and <bold>G</bold>. Song recordings are presented at three different speeds. In all oscillograms the two upper lines are recordings of hind leg movements and the lower line is the sound recording. Frequency spectra are shown in kHz for the short syllables <bold>I</bold> and the main echeme <bold>J</bold> of courtship.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g005.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040934.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040934</uri>
            </graphic>
          </fig>
          <p>The courtship song starts with producing soft and short syllables, repeated at the rate of ~ 2–3 /s (Fig. <xref ref-type="fig" rid="F5">5G</xref>); however, sometimes these syllables are repeated very irregularly (Fig. <xref ref-type="fig" rid="F5">5D</xref>). When producing element 1, the legs are moved with a very small amplitude. In ~ 30 s–1 min a male starts an element 2 that is similar to the calling song. However, the echeme duration is longer, up to ~ 7–8 s. The echeme also begins quietly, reaching maximum intensity in ~ 2–4 s. The amplitude of the leg movements gradually increases but abruptly decreases close to the end; as a result, the syllables of smaller amplitude are generated at the end of each echeme. Then, the long element 1 is again produced, followed by element 2. The frequency spectrum of element 1 shows three maxima in a wide range (Fig. <xref ref-type="fig" rid="F5">5I</xref>), whereas the spectrum of element 2 is characterised by many peaks between 18 and 35 kHz (Fig. <xref ref-type="fig" rid="F5">5J</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Comparative remarks" id="SECID0EOXAE">
          <title>Comparative remarks.</title>
          <p>The current recordings of calling song are similar to the recordings published by <xref ref-type="bibr" rid="B7">Bukhvalova and Vedenina (1998)</xref>, despite usage of the different recording equipment. The courtship song of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part></tp:taxon-name></italic> is shown for the first time. According to <xref ref-type="bibr" rid="B6">Bey-Bienko and Mistshenko (1951)</xref>, the localities where our material and that collected by <xref ref-type="bibr" rid="B7">Bukhvalova and Vedenina (1998)</xref> are situated within the range of subspecies <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="iliensis">iliensis</tp:taxon-name-part></tp:taxon-name></italic> Mistsh. In the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">Mesasippus</tp:taxon-name-part></tp:taxon-name></italic>, there are nine species occurring in Kazakhstan, Uzbekistan, western China, and north-western Mongolia. Currently, we have information on bioacoustics in only one species of the genus.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Orthoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Acrididae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">D2453D6C-0820-561B-B458-9739FA6B0DAD</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>(Philippi, 1830)</tp:taxon-authority>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="distribution" id="SECID0EF1AE">
          <title>Distribution.</title>
          <p>Europe from France to the east of European Russia, reaching as far north as Leningrad region and as far south as the northern Caucasus. This species occurs very locally, either in mountains or in dry pine forests (sandy heathlands and forest clearings).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0EK1AE">
          <title>Material</title>
          <p><bold>(Fig. <xref ref-type="fig" rid="F1">1</xref>).</bold> 7. Ukraine: Cherkassy region, ~ 17 km S of Kanev, glades in pine forest, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[31.491833,49.593000]}" id="NCID0E11AE">49°35.58'N, 31°29.51'E</named-content></named-content>, 22.06.2010, song recordings in 3 ♂; 8. Russia: Ul’yanovsk region, Novospassky district, Monastyrsky Sungur, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[47.663983,53.241383]}" id="NCID0EC2AE">53°14.483'N, 47°39.839'E</named-content></named-content>, 05.07.2022, song recordings in 2 ♂.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="References to song" id="SECID0EH2AE">
          <title>References to song.</title>
          <p><xref ref-type="bibr" rid="B19">Ragge and Reynolds 1998</xref>: recordings of calling song from Germany. <xref ref-type="bibr" rid="B7">Bukhvalova and Vedenina 1998</xref>: recordings of calling song from Ukraine, Zakarpat’je.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Song" id="SECID0EV2AE">
          <title>Song.</title>
          <p>During courtship, a male generates several echemes each lasting ~ 2–4 s and repeated at ~ 3–4 s intervals (Fig. <xref ref-type="fig" rid="F6">6A</xref>). Each echeme has either one or two elements. The first element is a whizzing sound produced by the low-amplitude leg movements at the rate of ~ 34 /s. It begins quietly showing a gradual crescendo for approximately the first half of its duration. The two legs are moved with the notable phase shift (Fig. <xref ref-type="fig" rid="F6">6F</xref>). During the short up movement, each leg generates one soft pulse, whereas during stepwise down movement, each leg produces four pulses of increasing sound intensity. Thus, each syllable contains five pulses repeated at the rate of ~ 170 /s. Sometimes, immediately after the first echeme element, the legs are moved asynchronously with the high amplitude, which produces almost no sound (Fig. <xref ref-type="fig" rid="F6">6A, D</xref>; see Suppl. material <xref ref-type="supplementary-material" rid="S3">3</xref>). Before the copulation attempts, a male moves the legs synchronously with high amplitude generating several noisy syllables repeated at the rate of ~ 3.5 /s (Fig. <xref ref-type="fig" rid="F6">6A, C, E</xref>). The frequency spectrum of the main echeme is broad with maximum energy between 13 and 30 kHz (Fig. <xref ref-type="fig" rid="F6">6H</xref>); the spectrum of precopulatory sound has several maxima between 5 and 20 kHz (Fig. <xref ref-type="fig" rid="F6">6I</xref>).</p>
          <fig id="F6" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure6</object-id>
            <object-id content-type="arpha">A3163C76-3493-5725-831B-DBC3B5039D9B</object-id>
            <label>Figure 6.</label>
            <caption>
              <p>Oscillograms of the courtship song <bold>A–G</bold> and frequency spectra <bold>H, I</bold> in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic>. Courtship songs are shown in males from Ukraine <bold>A, D, F</bold> and Russia <bold>B, C, E, G</bold>. Song recordings are presented at three different speeds. In all oscillograms except <bold>B, G</bold> the two upper lines are recordings of hind leg movements and the lower line is the sound recording. Courtship song of one-legged male is shown in <bold>B, G</bold>. Frequency spectra are shown in kHz for the main echeme <bold>H</bold> and precopulatory sound <bold>I</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g006.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040935.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040935</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Comparative remarks" id="SECID0E64AE">
          <title>Comparative remarks.</title>
          <p><xref ref-type="bibr" rid="B19">Ragge and Reynolds (1998)</xref> suggested sometimes two elements in the calling song of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic>, the first one being produced by the fast, low-amplitude leg vibrations and the second one by the slower, high-amplitude leg movements. We suggest that the second element mainly serves as a visual display during courtship, especially given the absence of the sound during the high-amplitude leg movements. <xref ref-type="bibr" rid="B19">Ragge and Reynolds (1998)</xref> also supposed that the leg movements producing the whizzing noise could be complex and asynchronous. We support their assumption by analysing the leg-movement pattern. The main rhythm of the leg movements (34 /s) could originate from the half rates of the wing beat (<xref ref-type="bibr" rid="B27">Vedenina and Mugue 2011</xref>). However, rapid vibratory movements during each downstroke (170 /s) are higher than the double rates of the wing beat. We consider the leg-movement pattern of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic> to be rare, if not unique, within <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name>.</p>
          <p>The uniqueness of the leg-movement pattern of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic> within <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name> is in a concordance with its controversial taxonomic status. Despite this species is attributed to the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part></tp:taxon-name></italic>, different phylogenetic reconstructions based on various molecular markers (<xref ref-type="bibr" rid="B20">Sevastianov et al. 2023</xref>; <xref ref-type="bibr" rid="B21">Schmidt et al. 2024</xref>) indicate that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic> forms an outgroup not only to the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part></tp:taxon-name></italic>, but even to the tribes <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Stenobothrini</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Gomphocerini</tp:taxon-name-part></tp:taxon-name>. At the same time, morphologically this species could be easily attributed to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part></tp:taxon-name></italic>, although being a rather brachypterous species.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Orthoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Acrididae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">B5A1FBD7-2EB9-563D-8778-D5B6ED368637</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>(Fischer-Waldheim, 1846)</tp:taxon-authority>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="distribution" id="SECID0EVCAG">
          <title>Distribution.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>: Transcaucasia, Asia Minor, Iraq, northern Iran. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> (Vorontsovski, 1928): from Ukraine to northern and western Kazakhstan, reaching as far south the northern Caucasus.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0EZDAG">
          <title>Material</title>
          <p><bold>(Fig. <xref ref-type="fig" rid="F1">1</xref>).</bold> 1. Ukraine: Nikolaev region, Pervomaisk district, Ostapovka, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[31.096667,47.970000]}" id="NCID0EJEAG">47°58.2'N, 31°05.8'E</named-content></named-content>, 05.07.2005, song recordings in 1 ♂; Georgia: 2. Algeti national park, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[44.359167,41.675833]}" id="NCID0EREAG">41°40.55'N, 44°21.55'E</named-content></named-content>, 1252 m a.s.l., 27.08.2023, song recordings in 5 ♂; 3. environs of Sighnaghi, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[45.853000,41.598500]}" id="NCID0EZEAG">41°35.91'N, 45°51.18'E</named-content></named-content>, 770 m a.s.l., song recordings in 1 ♂; Russia: 4. Samara region, Alekseevka district, Gerasimovka, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[51.509733,52.710600]}" id="NCID0EBFAG">52°42.636'N, 51°30.584'E</named-content></named-content>, 12.07.2012, song recordings in 2 ♂; 5. Orenburg region, environs of Buzuluk, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[52.296667,52.678333]}" id="NCID0EJFAG">52°40.7'N, 52°17.8'E</named-content></named-content>, 30.06.2020, song recording in 1 ♂; 6. Orenburg region, Saraktash district, Studentzy, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[55.855200,51.860650]}" id="NCID0ERFAG">51°51.639'N, 55°51.312'E</named-content></named-content>, 14.07.2012, song recordings in 1 ♂.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="References to song" id="SECID0EWFAG">
          <title>References to song.</title>
          <p><xref ref-type="bibr" rid="B29">Vedenina and Zhantiev 1990</xref>; <xref ref-type="bibr" rid="B8">Bukhvalova and Zhantiev 1994</xref>; <xref ref-type="bibr" rid="B26">Vedenina and Bukhvalova 2001</xref>: recordings of calling song from Moldova, Ukraine, south-eastern part of European Russia and northern Caucasus.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Song" id="SECID0EIGAG">
          <title>Song.</title>
          <p>The calling song is an echeme lasting 14–18 s in nominative subspecies, and shorter, ~ 4–10 s, in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> (Fig. <xref ref-type="fig" rid="F7">7</xref>). The legs are moved with a small phase shift at the rate of ~ 3.5–4.5 /s in nominative subspecies, and faster, at the rate of ~ 5–7 /s, in another subspecies. At the beginning of the echeme, each upstroke of the legs produces almost no sound, whereas each downstroke generates syllables of distinct pulses (Fig. <xref ref-type="fig" rid="F7">7C</xref>). Close to the echeme end, the legs produce a louder, but still relatively soft sound during each upstroke. Very often, the legs are moved faster close to the echeme end than at the beginning; as a result, distinct pulses disappear in the last syllables (Fig. <xref ref-type="fig" rid="F7">7D</xref>). This, however, occur only in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic>. The nominative subspecies usually does not increase the rate of the leg movements close to the echeme end.</p>
          <fig id="F7" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure7</object-id>
            <object-id content-type="arpha">BDE56785-91D5-51E1-B1B5-4C2E19912D2F</object-id>
            <label>Figure 7.</label>
            <caption>
              <p>Oscillograms of the calling song <bold>A–D</bold>, courtship songs <bold>E–H</bold> and frequency spectra <bold>I, J</bold> in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>. Courtship songs are shown in males from Orenburg region <bold>E–F</bold> and from Georgia <bold>G–H</bold>. Song recordings are presented at three different speeds. In all oscillograms the two upper lines are recordings of hind leg movements and the lower line is the sound recording. Frequency spectra are shown in kHz for the courtship songs from Orenburg region <bold>I</bold> and from Georgia <bold>J</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g007.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040936.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040936</uri>
            </graphic>
          </fig>
          <p>The courtship song is similar to the calling song but lasts longer. For example, in nominative subspecies the echeme duration varies in the range of 18–30 s (Fig. <xref ref-type="fig" rid="F7">7G</xref>). The frequency spectra are also similar in the calling and courtship songs. We, however, found some differences between the spectra of two subspecies. The spectrum of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> song has the main maximum around 35 kHz but also a small peak around 10 kHz (Fig. <xref ref-type="fig" rid="F7">7I</xref>). The spectrum of the song in nominative subspecies has many periodical peaks in the broad range of 5–40 kHz (Fig. <xref ref-type="fig" rid="F7">7J</xref>).</p>
          <p>After finishing the long courtship echeme, a male produces a very conspicuous display with his long antennae (the ratio of antennae length to head and pronotum length averages 1.89±0.07 in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic> in contrast to 1.64±0.1 in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="apricarius">apricarius</tp:taxon-name-part></tp:taxon-name></italic> or 1.70±0.17 in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fallax">fallax</tp:taxon-name-part></tp:taxon-name></italic>). First, antennae are moved in longitudinal plane backwards, and the two antennae are moved asymmetrically (Fig. <xref ref-type="fig" rid="F8">8A</xref>). Then, antennae are moved in horizontal plane, from side to side (Fig. <xref ref-type="fig" rid="F8">8B</xref>) and finally, antennae are moved in a circular manner (Fig. <xref ref-type="fig" rid="F8">8C</xref>). Immediately after the antennal movements (see Suppl. material <xref ref-type="supplementary-material" rid="S4">4</xref>), a male is trying to mate. We documented the antennal movements in the only males from Georgia (nominative subspecies). Concerning the males from other localities (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic>), we cannot say whether they show such visual display or not, since we did not pay attention to the antennal movements.</p>
          <fig id="F8" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure8</object-id>
            <object-id content-type="arpha">CDEB3E6A-1E02-5F23-9073-6FE8204CB65B</object-id>
            <label>Figure 8.</label>
            <caption>
              <p>Movements of antennae during courtship in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>. Successive stages of the antennae position are shown in <bold>A–C</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g008.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040937.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040937</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Comparative remarks" id="SECID0EXMAG">
          <title>Comparative remarks.</title>
          <p>The current recordings of calling song in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> are similar to the previous recordings. <xref ref-type="bibr" rid="B26">Vedenina and Bukhvalova (2001)</xref> showed that in two subspecies, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="ponticus">ponticus</tp:taxon-name-part></tp:taxon-name></italic>, the echeme duration varies greatly (in the range of ~ 3.5–15 s); however, it varies in different specimens referred to the same subspecies. On the basis of this similarity and similarity in morphology between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="ponticus">ponticus</tp:taxon-name-part></tp:taxon-name></italic>, the authors doubt whether these subspecies should be distinguished. By contrast, nominative subspecies differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> by some morphological characters (the body and wing lengths are larger in nominative subspecies) and some song parameters (the echeme duration is higher and the syllable rate is less in nominative subspecies).</p>
          <p>The leg-movement and song patterns in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic> are relatively simple and may be considered as plesiomorphic (<xref ref-type="bibr" rid="B27">Vedenina and Mugue 2011</xref>; <xref ref-type="bibr" rid="B20">Sevastianov et al. 2023</xref>). Moreover, it was suggested that the calling and courtship songs are similar in this species. Therefore, our discover of specific movements with antennae during courtship in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic> is remarkable. In future, it would be important to study if the males in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="purpuratus">purpuratus</tp:taxon-name-part></tp:taxon-name></italic> demonstrate similar movements with antennae as it was found in nominative subspecies. Considering the very long antennae typical for this species, we expect similar visual display in different subspecies.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Orthoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Acrididae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">87437CCA-B3CB-5CA6-9C95-F50B3D387206</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>(Miram, 1907)</tp:taxon-authority>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="distribution" id="SECID0EOSAG">
          <title>Distribution.</title>
          <p>Southern Siberia from Altai to Transbaikalia, southern part of the Russian Far East, Mongolia, China.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0ETSAG">
          <title>Material</title>
          <p><bold>(Fig. <xref ref-type="fig" rid="F1">1</xref>).</bold> Russia: 14. Altai Republic, Chemal district, Elekmonar, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[86.042067,51.456200]}" id="NCID0EDTAG">51°27.372'N, 86°02.524'E</named-content></named-content>, 459 m a.s.l., 03.08.2023, song recordings in 2 ♂; 15. Altai Republic, Ongudai district, ab. 3.5 km S of Inya, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[86.635333,50.414000]}" id="NCID0ELTAG">50°24.840'N, 86°38.120'E</named-content></named-content>, 769 m a.s.l., 05.08.2023, song recordings in 6 ♂.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="References to song" id="SECID0EQTAG">
          <title>References to song.</title>
          <p><xref ref-type="bibr" rid="B2">Benediktov 2005</xref>: recordings of calling song from Tuva; <xref ref-type="bibr" rid="B24">Tishechkin and Bukhvalova 2009</xref>: recordings of calling and courtship songs from Buryatia, Chita region and Maritime Province.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Song" id="SECID0E5TAG">
          <title>Song.</title>
          <p>The calling song is an echeme of variable duration ranging from ~ 6 to 20 s. Sometimes a male can produce several echemes with intervals of ~ 4–6 s (Fig. <xref ref-type="fig" rid="F9">9A</xref>). The legs being moved with a small phase shift at the rate of ~ 4–4.5 /s generate syllables where one can distinguish soft and loud parts (Fig. <xref ref-type="fig" rid="F9">9B, C</xref>). During each upstroke, almost no (in the beginning of echeme) or relatively soft (in the end of echeme) sound is produced; during downstroke, a loud sound is generated. The oscillograph analysis shows that the legs slightly vibrate during downstroke, which result to producing distinct pulses, especially at the beginning of echeme.</p>
          <fig id="F9" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure9</object-id>
            <object-id content-type="arpha">117DE3AD-DDBD-5EC5-A60A-0BF2D8B89F25</object-id>
            <label>Figure 9.</label>
            <caption>
              <p>Oscillograms of the calling song <bold>A–C</bold> rivalry song <bold>D–E</bold> courtship song <bold>F–G</bold> and frequency spectra <bold>H–J</bold> in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic>. Song recordings are presented at three different speeds. In all oscillograms except <bold>F, G</bold> the two upper lines are recordings of hind leg movements and the lower line is the sound recording. In <bold>F, G</bold> the movements of only one leg are shown. Frequency spectra are shown in kHz for the calling song <bold>H</bold> rivalry song <bold>I</bold> and courtship song <bold>J</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g009.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040938.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040938</uri>
            </graphic>
          </fig>
          <p>The rivalry song is a sequence of short echemes (Fig. <xref ref-type="fig" rid="F9">9D</xref>), which number can greatly vary. Each echeme consists of two or three syllables. The legs are moved almost synchronously at the rate of ~ 7.5 /s. The first syllable usually contains distinct pulses that are similar to those in the calling song. In the noisy second and third syllables the pulses are not distinguishable (Fig. <xref ref-type="fig" rid="F9">9E</xref>).</p>
          <p>A courted male generates an echeme or several echemes that are similar to the calling song. The legs are moved at the slightly higher rate than during calling (of ~ 5–6 /s). After this, a male can produce noisy syllables by the high-amplitude synchronous leg movements at the rate of ~ 2–3 /s (Fig. <xref ref-type="fig" rid="F9">9F, G</xref>). Before copulation attempt, males move with the long antennae (the ratio of antennae length to head and pronotum length averages 2.07±0.14). The movement pattern is simpler than in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>: antennae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic> are only moved in a circular manner as in Fig. <xref ref-type="fig" rid="F8">8C</xref>. Sometimes, the courting male produces the song similar to rivalry song, and the short echemes containing two types of syllables (as in Fig. <xref ref-type="fig" rid="F9">9E</xref>) can alternate with the noisy courtship element (as in Fig. <xref ref-type="fig" rid="F9">9G</xref>).</p>
          <p>The power spectra of the calling and rivalry songs are similar and have maximum energy between 20 and 35 kHz (Fig. <xref ref-type="fig" rid="F9">9H, I</xref>). The power spectrum of specific courtship element has two broad peaks, between 5 and 15 kHz and between 20 and 30 kHz (Fig. <xref ref-type="fig" rid="F9">9J</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Comparative remarks" id="SECID0ESXAG">
          <title>Comparative remarks.</title>
          <p>The current recordings from Altai are similar to the previous recordings from the more eastern localities of this species (<xref ref-type="bibr" rid="B2">Benediktov 2005</xref>; <xref ref-type="bibr" rid="B24">Tishechkin and Bukhvalova 2009</xref>). We, however, found slight differences in courtship songs between specimens from Altai and those from Buryatia, Chita region, and Maritime Province (<xref ref-type="bibr" rid="B24">Tishechkin and Bukhvalova 2009</xref>). At the same time, considering variable courtship behaviour and different technique of sound recordings, we suggest no principal differences between the recordings.</p>
          <p>By contrast, the antennal movements in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic> during courtship are documented for the first time. This species is similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic> by the remarkably long antennae. The two species are also similar in plesiomorphic pattern of the leg movements during calling behaviour. <xref ref-type="bibr" rid="B25">Tishechkin and Vedenina (2016)</xref> suggested that these two allopatric species are characterised by similar song type and acoustic strategy. The usage of long antennae in courtship behaviour in both species even confirms such similarity. On the other hand, both morphological (<xref ref-type="bibr" rid="B6">Bey-Bienko and Mistshenko 1951</xref>) and molecular studies (<xref ref-type="bibr" rid="B20">Sevastianov et al. 2023</xref>) suggest that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic> are not closely related species. We suggest one of the reasons of their allopatric distribution to be a similarity of acoustic niches, which prevents them from effectively finding members of their own species at the same biotope (<xref ref-type="bibr" rid="B8">Bukhvalova and Zhantiev 1994</xref>; <xref ref-type="bibr" rid="B25">Tishechkin and Vedenina 2016</xref>).</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Orthoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Acrididae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">01564E8C-AA9F-5CAD-8D32-2377450ED86F</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>(Zubowsky, 1899)</tp:taxon-authority>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="distribution" id="SECID0EK2AG">
          <title>Distribution.</title>
          <p>Southern Siberia, the southern part of Russian Far East, the north-eastern China, Korea.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0EP2AG">
          <title>Material</title>
          <p><bold>(Fig. <xref ref-type="fig" rid="F1">1</xref>).</bold> Russia: 10. Altai Republic, ab. 6 km of Chemal, environs of Elekmonar, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[85.998333,51.483333]}" id="NCID0E62AG">51°29.0'N, 85°59.9'E</named-content></named-content>, 417 m a.s.l., 06.08.2017, song recordings in 2 ♂, 12.08.2021, song recordings in 4 ♂; 11. Altai Republic, Ongudai district, ab. 7 km N of Malyi Yaloman, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[86.563050,50.560033]}" id="NCID0EH3AG">50°33.602'N, 86°33.783'E</named-content></named-content>, 740 m a.s.l., 09.08.2023, song recordings in 1 ♂; 12. Altai Republic, Ongudai district, near Shirlak waterfall, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[87.221667,50.343333]}" id="NCID0EP3AG">50°20.6'N, 87°13.3'E</named-content></named-content>, 1064 m a.s.l., 14.08.2021, song recordings in 3 ♂; 13. Altai Republic, Ongudai district, ab. 15 km NWW of Chibit, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[87.324667,50.360617]}" id="NCID0EX3AG">50°21.637'N, 87°19.480'E</named-content></named-content>, 1056 m a.s.l., 09.08.2023, song recordings in 1 ♂.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="References to song" id="SECID0E33AG">
          <title>References to song.</title>
          <p><xref ref-type="bibr" rid="B7">Bukhvalova and Vedenina 1998</xref>; <xref ref-type="bibr" rid="B26">Vedenina and Bukhvalova 2001</xref>; <xref ref-type="bibr" rid="B2">Benediktov 2005</xref>: recordings of calling song from Altai and Maritime Province.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Song" id="SECID0EO4AG">
          <title>Song.</title>
          <p>The calling song is a sequence of several echemes lasting ~ 1.5–3 s and separated by intervals of ~ 2–6 s (Fig. <xref ref-type="fig" rid="F10">10A</xref>). In each echeme, one can distinguish two parts (see Suppl. material <xref ref-type="supplementary-material" rid="S5">5</xref>). In the first part, the legs are moved synchronously at the rate of 7.5–10 /s, which result to generation of simple regular syllables (element 1, Fig. <xref ref-type="fig" rid="F10">10B</xref>). The sound is generated mainly during downstroke. In the second part, the legs are moved asynchronously, which results to generation of the louder element 2 containing pulses of varying amplitude. The duration of element 2 is usually 2× as short as duration of element 1.</p>
          <fig id="F10" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure10</object-id>
            <object-id content-type="arpha">4FBBFD4E-3F17-5595-9398-9336BF325BF1</object-id>
            <label>Figure 10.</label>
            <caption>
              <p>Oscillograms of the calling song <bold>A, B</bold> courtship songs <bold>C–H</bold> and frequency spectra <bold>I–K</bold> in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>. Courtship songs of two males are shown in <bold>C</bold> and <bold>G</bold>. Song recordings are presented at two different speeds. In all oscillograms the two upper lines are recordings of hind leg movements and the lower line is the sound recording. Different elements of the calling song are shown in <bold>B</bold> of the courtship song – in <bold>C</bold>. Frequency spectra are shown in kHz for the calling song <bold>I</bold> for the courtship element 1 <bold>J</bold> and courtship element 2 <bold>K</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g010.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040939.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040939</uri>
            </graphic>
          </fig>
          <p>The courtship song starts similarly to the first part of the calling song: the legs are moved synchronously and generate the soft sound during upstroke and the louder sound during downstroke (Fig. <xref ref-type="fig" rid="F10">10C, D</xref>). In ~ 2–10 s, however, the temporal structure of syllables is gradually changing. The intervals between syllables become fuzzy and the very syllables become louder. Sometimes, the intervals between syllables become completely indistinguishable (Fig. <xref ref-type="fig" rid="F10">10H</xref>). This main echeme (element 1 of courtship) usually lasts for ~ 15–30 s, but can continue for more than 1 min. Oscillographic analysis of the leg movements shows that amplitude of the two neighbouring strokes slightly differs, however, this is more expressed in one leg (Fig. <xref ref-type="fig" rid="F10">10D, E</xref>). After the prolonged element 1, a much shorter (lasting ~ 6–9 s) element 2 follows. The legs are moved synchronously but at the lower position, and the leg movements have the more complex pattern than during element 1, which implies alternation of several low-amplitude strokes with one higher-amplitude stroke. The legs generate syllables repeated at the rate of 8–10 /s (Fig. <xref ref-type="fig" rid="F10">10E</xref>); each syllable contains one high-amplitude pulse and several soft pulses. In ~ 1 s after the end of element 2, the shortest (lasting ~ 1–2 s) element 3 follows (Fig. <xref ref-type="fig" rid="F10">10C, F, G</xref>; see Suppl. material <xref ref-type="supplementary-material" rid="S6">6</xref>). The leg movements have the more irregular pattern than during element 2, moreover, the two legs are moved alternately. As a result, one can distinguish sound pulses repeated at the rate of ~ 15–20 /s.</p>
          <p>The frequency spectra of both types of the song are remarkable because they occupy the band lower than 25 kHz. The spectrum of the calling song has two peaks around 7 and 18 kHz (Fig. <xref ref-type="fig" rid="F10">10I</xref>), the spectrum of the courtship element 1 has several peaks in the range from 5 to 25 kHz (Fig. <xref ref-type="fig" rid="F10">10J</xref>), and the spectrum of the courtship element 2 has two peaks at approximately 5 and 15 kHz (Fig. <xref ref-type="fig" rid="F10">10K</xref>).</p>
          <p>The elements 1 and 2 of the calling song and the elements 1–3 of the courtship song are produced at the different leg positions. During generation of the calling element 1 and the courtship elements 2 and 3, the legs are moved at the low position. Presumably the distal stridulatory pegs of the hind femora are used in generation of these elements. We compared the lengths of the stridulatory files between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>, the species with the simpler song. The file length appeared to be almost 2× longer in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> than that in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>. In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>, the most distal stridulatory pegs are at about the level of the first tibial spine if tibia is pressed to femur (Fig. <xref ref-type="fig" rid="F11">11A</xref>). In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>, the most distal stridulatory pegs are at the level between 5<sup>th</sup> and 6<sup>th</sup> tibial spines (Fig. <xref ref-type="fig" rid="F11">11B</xref>).</p>
          <fig id="F11" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1200.118422.figure11</object-id>
            <object-id content-type="arpha">3E9ED783-A650-5D56-94B8-71EF0E0E2B5D</object-id>
            <label>Figure 11.</label>
            <caption>
              <p>Stridulatory files SF on the hind femora in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic><bold>A</bold> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic><bold>B</bold>.</p>
            </caption>
            <graphic xlink:href="zookeys-1200-001_article-118422__-g011.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040940.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1040940</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Comparative remarks" id="SECID0ELEBG">
          <title>Comparative remarks.</title>
          <p>Our recordings of both types of the song are generally similar to those previously described by different authors (<xref ref-type="bibr" rid="B26">Vedenina and Bukhvalova 2001</xref>; <xref ref-type="bibr" rid="B2">Benediktov 2005</xref>). However, previous authors argued about the functions of the two song types in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>. According to <xref ref-type="bibr" rid="B26">Vedenina and Bukhvalova (2001)</xref>, different song types are produced by a male sitting alone, therefore they can be considered as the calling song variants. <xref ref-type="bibr" rid="B2">Benediktov (2005)</xref> suggested the first song type to be the calling song and the second song type to be the courtship song. Our current data are more concordant with the data of <xref ref-type="bibr" rid="B2">Benediktov (2005)</xref>. However, our observations of behaviour in this species show that the males sitting without females in laboratory for several days start to produce both song types almost equally often. In nature, by contrast, solitary males usually sing the first song type, whereas the males sitting near by the females typically produce the second song type. Therefore, we also assign the different functions to the different song types.</p>
          <p>Analysis of the elaborate leg movements during stridulation in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> allowed us to suggest unusually long stridulatory file. Up to now, only several species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="biguttulus">biguttulus</tp:taxon-name-part></tp:taxon-name></italic> group with the long stridulatory files were known (<xref ref-type="bibr" rid="B1">Benediktov 1999</xref>; <xref ref-type="bibr" rid="B35">Willemse et al. 2009</xref>; <xref ref-type="bibr" rid="B22">Tarasova et al. 2021</xref>). One of these species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="biguttulus">biguttulus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="ehedickei">ehedickei</tp:taxon-name-part></tp:taxon-name></italic>, also uses its distal part of the stridulatory file in generation of a relatively soft ‘aftersong’. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>, however, produces a relatively loud sound with the distal part of the stridulatory file.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="﻿Conclusions" id="SECID0EOHBG">
      <title>﻿Conclusions</title>
      <p>1. In seven species of subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Gomphocerinae</tp:taxon-name-part></tp:taxon-name>, the stridulatory leg movements were recorded and analysed for the first time. In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">Mesasippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic>, the legs are moved in a relatively simple pattern that is considered to be plesiomorphic (<xref ref-type="bibr" rid="B27">Vedenina and Mugue 2011</xref>; <xref ref-type="bibr" rid="B20">Sevastianov et al. 2023</xref>). Other four species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> demonstrate more complex leg movements, which are considered to be the more evolutionary advanced patterns.</p>
      <p>2. The number of sound elements in the calling and courtship songs is the same in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>. The courtship song contains one additional sound element in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesasippus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kozhevnikovi">kozhevnikovi</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic>. The highest number of courtship sound elements is found in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>.</p>
      <p>3. The songs in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic> are shown for the first time. This species is remarkable by crepitation in flight and generation of short wing beats, which brings this species closer to other three species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part></tp:taxon-name></italic> (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="rubicundulus">rubicundulus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cotticus">cotticus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hyalosuperficies">hyalosuperficies</tp:taxon-name-part></tp:taxon-name></italic>). Moreover, we found a high similarity between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cotticus">cotticus</tp:taxon-name-part></tp:taxon-name></italic> in acoustic behaviour, morphology and ecological preferences, which may indicate that these species belong to the same taxon. However, a large distance between habitats of these species do not allow us to make final conclusions.</p>
      <p>4. The courtship songs in two species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>, contain several sound elements. The complexity of the courtship song in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> is in a concordance with the complexity of courtship behaviour in other species of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix.</tp:taxon-name-part></tp:taxon-name></italic> The song complexity in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> stands apart because it is not typical for the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Gomphocerini</tp:taxon-name-part></tp:taxon-name>. The different courtship song elements in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic> are produced by vibrating hind femora at the different positions. Analysis of the leg movements revealed the participance of different parts of the long stridulatory file in sound production.</p>
      <p>5. A maximum energy of the song power spectra in 6 species studied lies in ultrasound range (higher than 20 kHz). In only <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic>, the main peaks in the song power spectra lie lower than 20 kHz. This should be considered during analysis of the recordings made by portable recorders with a frequency range not exceeding 12.5–15 kHz.</p>
      <p>6. The courtship behaviour in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic> includes a different visual display. For the first time we documented conspicuous movements with long antennae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hammarstroemi">hammarstroemi</tp:taxon-name-part></tp:taxon-name></italic>, which are demonstrated just before a copulation attempt. We suggest a correlation between the antenna length and the antennal movements during courtship. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic> shows slight movements with antennae and the whole body, and very conspicuous movements with palps during courtship, which are very different from those in other species of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part></tp:taxon-name></italic>.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgments</title>
      <p>We are grateful to Tatiana Tarasova and Lev Schestakov (Institute for Information Transmission Problems, Moscow) for their help with the field trips and song recordings. We also thank the reviewers, Charlie Woodrow (University of Lincoln, UK) and Zhu-Qing He (East China Normal University, China), who substantially improved our manuscript.</p>
    </ack>
    <sec sec-type="﻿Additional information" id="SECID0EJUBG">
      <title>﻿Additional information</title>
      <sec sec-type="Conflict of interest" id="SECID0ENUBG">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="SECID0ESUBG">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Funding" id="SECID0EXUBG">
        <title>Funding</title>
        <p>The current study was supported by the Russian Scientific Foundation (grant 23-24-00533).</p>
      </sec>
      <sec sec-type="Author contributions" id="SECID0E3UBG">
        <title>Author contributions</title>
        <p>All authors have contributed equally.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="SECID0EBVBG">
        <title>Author ORCIDs</title>
        <p>Varvara Vedenina <ext-link xlink:href="https://orcid.org/0000-0002-2694-4152" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-2694-4152</ext-link></p>
        <p>Nikita Sevastianov <ext-link xlink:href="https://orcid.org/0000-0002-1563-5194" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-1563-5194</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="SECID0ERVBG">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text or Supplementary Information.</p>
      </sec>
    </sec>
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    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.suppl1</object-id>
        <object-id content-type="arpha">92D97A44-ECA3-5F21-BC5D-651F8A72304A</object-id>
        <label>Supplementary material 1</label>
        <caption>
          <p>Courtship in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Myrmeleotettix">Myrmeleotettix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palpalis">palpalis</tp:taxon-name-part></tp:taxon-name></italic></p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>mpg</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>The male produces high-amplitude strokes with hind legs, moves his body from side to side, generates low-amplitude movements with antennae, and conspicuously moves with palps up and down. In this movie and some others, the pieces of reflecting foil glued to the distal part of each hind femur remained from the leg-movement recordings.</p>
        </statement>
        <media xlink:href="zookeys-1200-001_article-118422__-s001.mpg" mimetype="video" mime-subtype="mpeg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040941.mpg">
          <uri content-type="original_file">https://binary.pensoft.net/file/1040941</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Varvara Vedenina, Nikita Sevastianov, Evgenia Kovalyova</attrib>
      </supplementary-material>
      <supplementary-material id="S2" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.suppl2</object-id>
        <object-id content-type="arpha">E15493A6-E794-5C15-B79E-FC678493A86E</object-id>
        <label>Supplementary material 2</label>
        <caption>
          <p>Courtship in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stenobothrus">Stenobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="newskii">newskii</tp:taxon-name-part></tp:taxon-name></italic></p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>mpg</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>The male generates sound by both femoral-tegminal stridulation and wing clapping. The female responds by stridulation. The male attempts to copulate.</p>
        </statement>
        <media xlink:href="zookeys-1200-001_article-118422__-s002.mpg" mimetype="video" mime-subtype="mpeg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040942.mpg">
          <uri content-type="original_file">https://binary.pensoft.net/file/1040942</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Varvara Vedenina, Nikita Sevastianov, Evgenia Kovalyova</attrib>
      </supplementary-material>
      <supplementary-material id="S3" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.suppl3</object-id>
        <object-id content-type="arpha">8C358310-86E1-5D35-9CF0-7D99C5EABD18</object-id>
        <label>Supplementary material 3</label>
        <caption>
          <p>Courtship in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pullus">pullus</tp:taxon-name-part></tp:taxon-name></italic></p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>mpg</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>The first whizzing element is produced by the low-amplitude leg movements, after which the legs are moved asynchronously with the high amplitude, which produces almost no sound.</p>
        </statement>
        <media xlink:href="zookeys-1200-001_article-118422__-s003.mpg" mimetype="video" mime-subtype="mpeg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040943.mpg">
          <uri content-type="original_file">https://binary.pensoft.net/file/1040943</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Varvara Vedenina, Nikita Sevastianov, Evgenia Kovalyova</attrib>
      </supplementary-material>
      <supplementary-material id="S4" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.suppl4</object-id>
        <object-id content-type="arpha">3DD4DAFD-2F69-5EC5-9B43-174164C629C8</object-id>
        <label>Supplementary material 4</label>
        <caption>
          <p>Courtship in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorthippus">Chorthippus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrocerus">macrocerus</tp:taxon-name-part></tp:taxon-name></italic></p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>mpg</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>Immediately after the song generation, the male conspicuously moves with antennae and attempts to copulate.</p>
        </statement>
        <media xlink:href="zookeys-1200-001_article-118422__-s004.mpg" mimetype="video" mime-subtype="mpeg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040944.mpg">
          <uri content-type="original_file">https://binary.pensoft.net/file/1040944</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Varvara Vedenina, Nikita Sevastianov, Evgenia Kovalyova</attrib>
      </supplementary-material>
      <supplementary-material id="S5" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.suppl5</object-id>
        <object-id content-type="arpha">97F85880-DB52-5899-A832-9BE5613817BF</object-id>
        <label>Supplementary material 5</label>
        <caption>
          <p>Calling in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic></p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>mpg</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>The male produces two elements, one element when the legs are maintained at the higher position, the second element when the legs are kept at the lower position.</p>
        </statement>
        <media xlink:href="zookeys-1200-001_article-118422__-s005.mpg" mimetype="video" mime-subtype="mpeg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040945.mpg">
          <uri content-type="original_file">https://binary.pensoft.net/file/1040945</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Varvara Vedenina, Nikita Sevastianov, Evgenia Kovalyova</attrib>
      </supplementary-material>
      <supplementary-material id="S6" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/zookeys.1200.118422.suppl6</object-id>
        <object-id content-type="arpha">09D62008-F68B-5AF0-899E-D68AA7382B90</object-id>
        <label>Supplementary material 6</label>
        <caption>
          <p>Courtship in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaulacobothrus">Megaulacobothrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethalinus">aethalinus</tp:taxon-name-part></tp:taxon-name></italic></p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>mpg</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>Three elements of the courtship song are produced: element 1 is produced by synchronous leg movements kept at the higher position, element 2 is produced by also synchronous leg movements but kept at the lower position, the short element 3 is produced by asynchronous leg movements.</p>
        </statement>
        <media xlink:href="zookeys-1200-001_article-118422__-s006.mpg" mimetype="video" mime-subtype="mpeg" position="float" orientation="portrait" xlink:type="simple" id="oo_1040946.mpg">
          <uri content-type="original_file">https://binary.pensoft.net/file/1040946</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Varvara Vedenina, Nikita Sevastianov, Evgenia Kovalyova</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
