153 lines
18 KiB
XML
153 lines
18 KiB
XML
<document id="B404B05FF24F43BAD5DEB7C533E057AC" ID-DOI="10.1093/zoolinnean/zlad155" ID-ISSN="0024-4082" ID-Zenodo-Dep="13716099" IM.bibliography_approvedBy="karina" IM.illustrations_approvedBy="karina" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="karina" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="karina" checkinTime="1722845461810" checkinUser="plazi" docAuthor="Ntatsopoulos, Konstantinos, Nabozhenko, Maxim V, Jelinscaia Lagou, Loudmila, Chigray, Ivan A, Gagarina, Ludmila V, Alpagut Keskin, Nurşen, Keskin, Bekir & Papadopoulou, Anna" docDate="2023" docId="A67689422F44D5701554FACD8AD3D3D5" docLanguage="en" docName="zlad155.pdf" docOrigin="Zoological Journal of the Linnean Society 201 (3)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlad155" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-2023.journal_article" docStyleVersion="7" docTitle="Helopina" docType="treatment" docVersion="2" lastPageNumber="8" masterDocId="5A4FF13A2F43D5771400FFE28920D256" masterDocTitle="Beetles and lichens: tracing the origins and evolution of lichenophagy within the darkling beetle tribe Helopini (Coleoptera: Tenebrionidae)" masterLastPageNumber="8" masterPageNumber="1" pageNumber="1" updateTime="1725653545490" updateUser="ExternalLinkService">
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<mods:title id="DB5D90D67E2D46A4DF66EE10D1588A0B">Beetles and lichens: tracing the origins and evolution of lichenophagy within the darkling beetle tribe Helopini (Coleoptera: Tenebrionidae)</mods:title>
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<mods:namePart id="E224A27CBE70553B2FE1A6E80C0A5A4E">Ntatsopoulos, Konstantinos</mods:namePart>
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<mods:namePart id="F1C6437461F7B7013B190198C32B5DCE">Nabozhenko, Maxim V</mods:namePart>
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<mods:namePart id="17927BC5AE21D537BF1ED80F94848DC1">Jelinscaia Lagou, Loudmila</mods:namePart>
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<mods:namePart id="88F2597411E30584CAAB82974EA5D284">Chigray, Ivan A</mods:namePart>
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<mods:namePart id="30B361A2A9D4F19F9C8500916419FBE5">Gagarina, Ludmila V</mods:namePart>
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<mods:namePart id="054596A035451BFEDBDC0630DF96A66A">Alpagut Keskin, Nurşen</mods:namePart>
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<mods:namePart id="A7FA930DFC510178A9B36F2D59318DA1">Keskin, Bekir</mods:namePart>
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<mods:namePart id="CD4F39F226EC57F50B3C968A6C8DCB63">Papadopoulou, Anna</mods:namePart>
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<mods:title id="EE277BD404FAEA182D3C134530108425">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="D2074A0E1298180D15918C87159107BB">2023</mods:date>
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<mods:number id="C211B0D7C75D8FE3006D5A142D14803A">2023-10-27</mods:number>
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<mods:number id="3CAA9ABE3715515F99686162113F6D6F">201</mods:number>
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<mods:url id="90A6AB1296593F4243CA0A20E604A25D">http://dx.doi.org/10.1093/zoolinnean/zlad155</mods:url>
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<subSubSection id="66C56BDF2F44D5701554FACD8B38D71F" box="[340,536,1327,1353]" pageId="7" type="nomenclature">
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<paragraph id="2E6038542F44D5701554FACD8B38D71F" blockId="7.[340,536,1327,1353]" box="[340,536,1327,1353]" pageId="7">
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<heading id="75288F382F44D5701554FACD8B38D71F" box="[340,536,1327,1353]" centered="true" fontSize="9" level="2" pageId="7" reason="3">
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<emphasis id="1CABE4462F44D5701554FACD8B38D71F" bold="true" box="[340,536,1327,1353]" pageId="7">
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Subtribe
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<taxonomicName id="E9DF43D72F44D57015B4FACD8B38D71F" baseAuthorityName="Nabozhenko and Lobl" baseAuthorityYear="2008" box="[436,536,1327,1353]" class="Insecta" family="Tenebrionidae" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="subTribe" subTribe="Helopina">Helopina</taxonomicName>
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</emphasis>
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<paragraph id="2E6038542F44D5701471FAB48AD3D3D5" blockId="7.[113,763,1366,1985]" lastBlockId="7.[810,1459,144,387]" pageId="7">
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Most clades within
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<taxonomicName id="E9DF43D72F44D570153BFAB488BAD738" baseAuthorityName="Nabozhenko and Lobl" baseAuthorityYear="2008" box="[315,410,1366,1390]" class="Insecta" family="Tenebrionidae" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="subTribe" subTribe="Helopina">Helopina</taxonomicName>
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were supported by high statistical values, and the inferred relationships largely agreed with the current taxonomy, given that all sampled genera were recovered as monophyletic (
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<figureCitation id="B6E424D12F44D5701516FA568873D79A" box="[278,339,1460,1484]" captionStart="Figure 1" captionStartId="4.[129,194,1702,1726]" captionTargetBox="[130,1474,143,1672]" captionTargetId="graphics-86@4.[130,1194,147,1674]" captionTargetPageId="4" captionText="Figure 1. Maximum likelihood phylogenetic tree reconstruction based on the concatenated dataset.Branch colours indicate subtribal groupings (yellow, Enoplopodina; red, Cylindrinotina; and blue, Helopina). Circles in nodes represent values of bootstrap support (BT), posterior probabilities (PP), gene concordance factors (gCF), and site concordance factors (sCF) as indicated in the key.Clades not recovered in the BI analysis are depicted with white PP. Beetle illustrations correspond to terminal taxa as indicated by the leưering in brackets." figureDoi="http://doi.org/10.5281/zenodo.13716101" httpUri="https://zenodo.org/record/13716101/files/figure.png" pageId="7">Fig. 1</figureCitation>
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; Supporting Information,
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<figureCitation id="B6E424D12F44D5701671FA568B9FD79A" box="[625,703,1460,1484]" captionStart="Figure 2" captionStartId="6.[129,194,853,877]" captionTargetBox="[137,1470,151,825]" captionTargetId="graphics-619@6.[129,1473,144,794]" captionTargetPageId="6" captionText="Figure 2. Results of the ancestral state estimation analysis. Terminal taxa are colour-coded according to their state as illustrated in the key. Pie charts demonstrate the probability of the ancestral state at each node. In order to provide a time frame for the emergence of different ancestral modes, a time axis is also provided (based on the BEAST run calibrated by known substitution rates).A, evolution of feeding modes under the ER model. B, evolution of habitat preferences under the SYM model within the tribe Helopini." figureDoi="http://doi.org/10.5281/zenodo.13716103" httpUri="https://zenodo.org/record/13716103/files/figure.png" pageId="7">Figs S2</figureCitation>
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, S3). Two strongly supported monophyletic groups of genera were identified within
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<taxonomicName id="E9DF43D72F44D5701520FA1188A2D45D" baseAuthorityName="Nabozhenko and Lobl" baseAuthorityYear="2008" box="[288,386,1523,1547]" class="Insecta" family="Tenebrionidae" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="subTribe" subTribe="Helopina">Helopina</taxonomicName>
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, the first including
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<taxonomicName id="E9DF43D72F44D5701642FA118BB3D45D" authorityName="Boieldieu" authorityYear="1865" box="[578,659,1523,1547]" class="Insecta" family="Tenebrionidae" genus="Euboeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D5701642FA118BB3D45D" box="[578,659,1523,1547]" italics="true" pageId="7">Euboeus</emphasis>
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</taxonomicName>
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,
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<taxonomicName id="E9DF43D72F44D570169DFA118BD5D45C" authorityName="Allard" authorityYear="1876" box="[669,757,1523,1546]" class="Insecta" family="Tenebrionidae" genus="Catomus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D570169DFA118BD5D45C" box="[669,757,1523,1546]" italics="true" pageId="7">Catomus</emphasis>
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</taxonomicName>
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, and
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<taxonomicName id="E9DF43D72F44D57014A6F9F08838D47C" authorityName="Reiuer" authorityYear="1922" box="[166,280,1554,1578]" class="Insecta" family="Tenebrionidae" genus="Stenohelops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D57014A6F9F08838D47C" box="[166,280,1554,1578]" italics="true" pageId="7">Stenohelops</emphasis>
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</taxonomicName>
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, and the second
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<taxonomicName id="E9DF43D72F44D57015E6F9F18B4AD47C" authorityName="Solier" authorityYear="1848" box="[486,618,1555,1578]" class="Insecta" family="Tenebrionidae" genus="Entomogonus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D57015E6F9F18B4AD47C" box="[486,618,1555,1578]" italics="true" pageId="7">Entomogonus</emphasis>
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</taxonomicName>
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,
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<taxonomicName id="E9DF43D72F44D570167DF9F08BE0D47C" authorityName="Fabricius" authorityYear="1775" box="[637,704,1554,1578]" class="Insecta" family="Tenebrionidae" genus="Helops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D570167DF9F08BE0D47C" box="[637,704,1554,1578]" italics="true" pageId="7">Helops</emphasis>
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</taxonomicName>
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, and
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<taxonomicName id="E9DF43D72F44D5701471F9D389FBD41F" authorityName="Allard" authorityYear="1876" box="[113,219,1585,1609]" class="Insecta" family="Tenebrionidae" genus="Raiboscelis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D5701471F9D389FBD41F" box="[113,219,1585,1609]" italics="true" pageId="7">Raiboscelis</emphasis>
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</taxonomicName>
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, which seem to be in accordance with the ‘catomoid’ and ‘helopioid’ lineages, respectively, as previously recognized based on the structure of male terminalia. Taxa assigned to the first group are characterized by a median lobe rounded towards the apical part, by a very long phallobase, and by short parameres almost entirely covered with elongate, asperate punctuation and short setae, whereas ‘helopioid’ species possess a median lobe acute at the apex, ossen of gradually diminished thickness towards the end, and by long parameres, only apically covered with short, spiniform bristles (
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<bibRefCitation id="4A4E45A52F44D57015AFF8A98B59D535" author="Nabozhenko MV" box="[431,633,1867,1891]" pageId="7" pagination="1024 - 72" refId="ref9307" refString="Nabozhenko MV. A revision of the genus Catomus Allard, 1876 and the allied genera (Coleoptera, Tenebrionidae) from the Caucasus, Middle Asia, and China. Entomological Review 2006 b; 86: 1024 - 72." type="journal article" year="2006">Nabozhenko 2006b</bibRefCitation>
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). Ŋe genus
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<taxonomicName id="E9DF43D72F44D5701471F889899AD5D4" authorityName="Allard" authorityYear="1876" box="[113,186,1899,1922]" class="Insecta" family="Asilidae" genus="Nesotes" kingdom="Animalia" order="Diptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D5701471F889899AD5D4" box="[113,186,1899,1922]" italics="true" pageId="7">Nesotes</emphasis>
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</taxonomicName>
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(traditionally assigned to the ‘catomoid’ group) was recovered as sister lineage to all other
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<taxonomicName id="E9DF43D72F44D57015E4F8688B63D5F4" baseAuthorityName="Nabozhenko and Lobl" baseAuthorityYear="2008" box="[484,579,1930,1954]" class="Insecta" family="Tenebrionidae" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="subTribe" subTribe="Helopina">Helopina</taxonomicName>
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in the BI analysis (Fig. 1; Supporting Information Fig. S3), although its placement
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<pageTitle id="6E40E0332F44D570172AFF728C93D2FE" box="[810,1459,144,168]" pageId="7">with regard to the other two clades remains rather uncertain</pageTitle>
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given the weak bootstrap support and concordance factors in the ML analysis (
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<figureCitation id="B6E424D12F44D57017DAFF2C8D32D2B1" box="[986,1042,206,231]" captionStart="Figure 1" captionStartId="4.[129,194,1702,1726]" captionTargetBox="[130,1474,143,1672]" captionTargetId="graphics-86@4.[130,1194,147,1674]" captionTargetPageId="4" captionText="Figure 1. Maximum likelihood phylogenetic tree reconstruction based on the concatenated dataset.Branch colours indicate subtribal groupings (yellow, Enoplopodina; red, Cylindrinotina; and blue, Helopina). Circles in nodes represent values of bootstrap support (BT), posterior probabilities (PP), gene concordance factors (gCF), and site concordance factors (sCF) as indicated in the key.Clades not recovered in the BI analysis are depicted with white PP. Beetle illustrations correspond to terminal taxa as indicated by the leưering in brackets." figureDoi="http://doi.org/10.5281/zenodo.13716101" httpUri="https://zenodo.org/record/13716101/files/figure.png" pageId="7">Fig. 1</figureCitation>
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||
; Supporting Information,
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||
<figureCitation id="B6E424D12F44D570111EFF2C8C43D2B1" box="[1310,1379,206,231]" captionStart="Figure 2" captionStartId="6.[129,194,853,877]" captionTargetBox="[137,1470,151,825]" captionTargetId="graphics-619@6.[129,1473,144,794]" captionTargetPageId="6" captionText="Figure 2. Results of the ancestral state estimation analysis. Terminal taxa are colour-coded according to their state as illustrated in the key. Pie charts demonstrate the probability of the ancestral state at each node. In order to provide a time frame for the emergence of different ancestral modes, a time axis is also provided (based on the BEAST run calibrated by known substitution rates).A, evolution of feeding modes under the ER model. B, evolution of habitat preferences under the SYM model within the tribe Helopini." figureDoi="http://doi.org/10.5281/zenodo.13716103" httpUri="https://zenodo.org/record/13716103/files/figure.png" pageId="7">Fig. S2</figureCitation>
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). Apart from the uncertainty regarding the placement of
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<taxonomicName id="E9DF43D72F44D5701139FF0D8CA2D350" authorityName="Allard" authorityYear="1876" box="[1337,1410,239,262]" class="Insecta" family="Asilidae" genus="Nesotes" kingdom="Animalia" order="Diptera" pageId="7" phylum="Arthropoda" rank="genus">
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<emphasis id="1CABE4462F44D5701139FF0D8CA2D350" box="[1337,1410,239,262]" italics="true" pageId="7">Nesotes</emphasis>
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</taxonomicName>
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, the results of our phylogenetic reconstruction within
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<taxonomicName id="E9DF43D72F44D570112DFEEF8CACD373" baseAuthorityName="Nabozhenko and Lobl" baseAuthorityYear="2008" box="[1325,1420,269,293]" class="Insecta" family="Tenebrionidae" kingdom="Animalia" order="Coleoptera" pageId="7" phylum="Arthropoda" rank="subTribe" subTribe="Helopina">Helopina</taxonomicName>
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are largely in agreement with the existing morphological studies, corroborating the currently recognized intergeneric groupings within the subtribe.
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</paragraph>
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</treatment>
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