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<document ID-CLB-Dataset="275942" ID-DOI="http://dx.doi.org/10.3897/asp.81.e107579" ID-GBIF-Dataset="742ed870-c756-4a81-8324-e95c9808643a" ID-Pensoft-Pub="1864-8312-81-819" ID-Pensoft-UUID="70148F72D847500A930DA76D2196759A" ID-ZooBank="50EE94AAD90A43E1B54E4FADE22F4668" ModsDocID="1864-8312-81-819" checkinTime="1698951166303" checkinUser="pensoft" docAuthor="Ge, Si-Xun, Jiang, Zhuo-Heng, Ren, Li-Li, van Achterberg, Cornelis &amp; Tan, Jiang-Li" docDate="2023" docId="8F85EE450C7F5ECBBDFD514C3F4DCBAD" docLanguage="en" docName="ArthropodSystPhyl 81: 819-844" docOrigin="Arthropod Systematics &amp; amp; Phylogeny 81" docPubDate="2023-11-02" docSource="http://dx.doi.org/10.3897/asp.81.e107579" docTitle="Lagenostephaninae Ge and Tan 2023, subf. nov." docType="treatment" docUuid="2231D702-9DC0-4277-8E5B-75611E0CBA24" docUuidSource="ZooBank" docVersion="2" id="70148F72D847500A930DA76D2196759A" lastPageNumber="819" masterDocId="70148F72D847500A930DA76D2196759A" masterDocTitle="New insights into the phylogeny of Stephanidae (Hymenoptera: Apocrita), with a revision of the fossil species" masterLastPageNumber="844" masterPageNumber="819" pageNumber="819" updateTime="1698998894021" updateUser="ExternalLinkService">
<mods:mods id="3D4D6BF9EC633A29D09C0C2711A14373" xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:title id="64A56E60C07EDF6208D775150FC99223">New insights into the phylogeny of Stephanidae (Hymenoptera: Apocrita), with a revision of the fossil species</mods:title>
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<mods:namePart id="7B13F8896002CC2C8DDCED1ECFAB1D99">Ge, Si-Xun</mods:namePart>
<mods:nameIdentifier id="37CE755F95107D3ACABDCBAC0C0CEEEF" type="ORCID">https://orcid.org/0000-0003-3769-1530</mods:nameIdentifier>
<mods:affiliation id="B073544F0A54C184A004DA2C024B823A">College of Forestry, Beijing Forestry University, Beijing 100083, China</mods:affiliation>
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<mods:namePart id="ADF3F40404A9BC6A543B4524926839FF">Jiang, Zhuo-Heng</mods:namePart>
<mods:affiliation id="CA04345C425367477191F424FC5A3288">School of Science, Westlake University, Hangzhou, China</mods:affiliation>
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<mods:namePart id="DC98950E2F12F4478822AECFBF7A5DF9">Ren, Li-Li</mods:namePart>
<mods:affiliation id="4B63D310DF28643EAD6D5332E01C799E">College of Forestry, Beijing Forestry University, Beijing 100083, China</mods:affiliation>
<mods:nameIdentifier id="E96F664DCA0C72F2F36C263ED69E2818" type="email">lily_ren@bjfu.edu.cn</mods:nameIdentifier>
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<mods:namePart id="6A1E6FBD5F32758020272F62F64A16E3">van Achterberg, Cornelis</mods:namePart>
<mods:nameIdentifier id="6804279E383BD4FA1B54FDAA6976F569" type="ORCID">https://orcid.org/0000-0002-6495-4853</mods:nameIdentifier>
<mods:affiliation id="66670D367EDD4130FB8D226E37FA1DA4">Shaanxi Key Laboratory for Animal Conservation / Key Laboratory of Resource Biology and Biotechnology in Western China, College of Life Sciences, Northwest University, 229 North Taibai Road, Xi' an, Shaanxi 710069, China</mods:affiliation>
<mods:nameIdentifier id="6BB0C20D1F3C8A0681ABE0B668EF808A" type="email">kees@vanachterberg.org</mods:nameIdentifier>
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<mods:namePart id="328F97481509B1A393727FA152B2B922">Tan, Jiang-Li</mods:namePart>
<mods:affiliation id="86440140F43FAC42355DAE1C16E657D2">Shaanxi Key Laboratory for Animal Conservation / Key Laboratory of Resource Biology and Biotechnology in Western China, College of Life Sciences, Northwest University, 229 North Taibai Road, Xi' an, Shaanxi 710069, China</mods:affiliation>
<mods:nameIdentifier id="808688391974313C9FE5EAF246BDABEE" type="email">tanjiangli@sina.com</mods:nameIdentifier>
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<mods:title id="9949EABAEE59734FBA343B6472B66F18">Arthropod Systematics &amp; amp; Phylogeny</mods:title>
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<mods:part id="026CB599FCF9157B2C273D06BF743774">
<mods:date id="622987CE607532DCAB1E45C8C5CD2ACB">2023</mods:date>
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<mods:number id="A128FCEA58050D324BC539619D3290AC">2023-11-02</mods:number>
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<mods:classification id="353A3268F5EF1531AA8C399E1C69685C">journal article</mods:classification>
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<treatment id="8F85EE450C7F5ECBBDFD514C3F4DCBAD" ID-GBIF-Taxon="216248482" LSID="urn:lsid:zoobank.org:act:2231D702-9DC0-4277-8E5B-75611E0CBA24" httpUri="http://treatment.plazi.org/id/8F85EE450C7F5ECBBDFD514C3F4DCBAD" lastPageNumber="819" pageId="0" pageNumber="819">
<subSubSection id="A5A844471AE133D0BB6F1D3CFA7D2389" pageId="0" pageNumber="819" type="nomenclature">
<paragraph id="65D5EB3239857B858EA7335430B6FEE6" pageId="0" pageNumber="819">
Subfamily †
<taxonomicName id="391DFB50F4B4E1C075E21923C06295F3" LSID="https://zoobank.org/2231D702-9DC0-4277-8E5B-75611E0CBA24" authority="Ge and Tan" authorityName="Ge and Tan" authorityYear="2023" family="Stephanidae" genus="Lagenostephaninae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Lagenostephaninae" order="Hymenoptera" pageId="0" pageNumber="819" rank="genus" status="subf. nov.">Lagenostephaninae Ge and Tan</taxonomicName>
<taxonomicNameLabel id="9DF4F806E617D0355EFA118FA89BC858" pageId="0" pageNumber="819">subf. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="437C88A2B92A93A1551CCA0AD4F72120" pageId="0" pageNumber="819" type="type genus">
<paragraph id="90E6C7280B9CAD2E84E747BDBEB2045D" pageId="0" pageNumber="819">Type genus.</paragraph>
<paragraph id="087A6D71F09B83E0EE357327407303FB" pageId="0" pageNumber="819">
†
<taxonomicName id="0B4D42288F1533352B7F7052C39334DE" class="Insecta" family="Stephanidae" genus="Lagenostephanus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Lagenostephanus" order="Hymenoptera" pageId="0" pageNumber="819" phylum="Arthropoda" rank="genus">
<emphasis id="5ADBFC75FB7E57379A1AE37F90E3F304" italics="true" pageId="0" pageNumber="819">Lagenostephanus</emphasis>
</taxonomicName>
Li, Rasnitsyn, Shih and Ren, 2017
</paragraph>
</subSubSection>
<subSubSection id="54F81FDCB609FA887960DE5AE6039AF5" pageId="0" pageNumber="819" type="diagnosis">
<paragraph id="6DA4C643816EA62DDB09AC53FB42962C" pageId="0" pageNumber="819">
Diagnosis (see also Table
<tableCitation id="9CE94E442C08A82FD34E29FCA56D8F66" captionStart="Table 1" captionStartId="T1" captionText="Table 1. Comparisons of some key morphological characteristics between subfamilies of Stephanidae." httpUri="http://table.plazi.org/id/F8F78DED151EC131D4B4FD929A16212B" pageId="0" pageNumber="819" tableUuid="F8F78DED151EC131D4B4FD929A16212B">1</tableCitation>
).
</paragraph>
<paragraph id="99E5325BA60E94E983137097EB67A402" pageId="0" pageNumber="819">Pronotum elongated with neck differentiated. Pronotal fold present. Forewing with vein 1-M arched; vein r-rs as long as 1-Rs; vein Rs+M and 1Cu nonparallel; vein 2Rs + M differentiated, not connect with free abscissa of vein M and vein 2-Rs. Hind coxa largely smooth, without transverse rugose. Hind femur robust and coriaceous. Tergite I almost as long as tergite II. Ovipositor sheath about as long as metasoma.</paragraph>
<caption id="F8F78DED151EC131D4B4FD929A16212B" ID-Table-UUID="F8F78DED151EC131D4B4FD929A16212B" httpUri="http://table.plazi.org/id/F8F78DED151EC131D4B4FD929A16212B" pageId="0" pageNumber="819" start="Table 1" startId="T1">
<paragraph id="6352732A911967C3927D329D6F9991BD" pageId="0" pageNumber="819">
<emphasis id="8F67C1F5938F7A1D9AC3DA5F214F1BBA" bold="true" pageId="0" pageNumber="819">Table 1.</emphasis>
Comparisons of some key morphological characteristics between subfamilies of
<taxonomicName id="8B1E08932F8D1DCD72B62D64257E24EA" class="Insecta" family="Stephanidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="819" phylum="Arthropoda" rank="family">Stephanidae</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="CFF368E0A802389F7CFE12F678414BE4" pageId="0" pageNumber="819">
<table id="50F134843A44EE4933312341FC22CC06" pageId="0" pageNumber="819">
<tr id="525173E45994FB060DD72598767D17DC" pageId="0" pageNumber="819">
<td id="EE0560B27F76F3E4E0B7E34501F8842B" colspan="1" pageId="0" pageNumber="819" rowspan="1">
<emphasis id="F8E5C3FA89BFA1CA4345F6F3A9FD8A46" bold="true" pageId="0" pageNumber="819">Characters/subfamilies</emphasis>
</td>
<td id="64064672CF2B949480012543BC56DAA5" colspan="1" pageId="0" pageNumber="819" rowspan="1">
†
<taxonomicName id="87D6AF62F573294120A6A62CDE93582D" authorityName="Ge and Tan" authorityYear="2023" family="Stephanidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="819" rank="subFamily" subFamily="Lagenostephaninae">
<emphasis id="427B35C3216FCAF7124018BACA7E149A" bold="true" pageId="0" pageNumber="819">Lagenostephaninae</emphasis>
</taxonomicName>
</td>
<td id="BD790063C7AB2E2E26B9B464C07F1BAE" colspan="1" pageId="0" pageNumber="819" rowspan="1">
<taxonomicName id="3522A08C7A41029A17894F191B645DDD" lsidName="" pageId="0" pageNumber="819" rank="subFamily" subFamily="Schletteriinae">
<emphasis id="49D1F45D9DAD5365BF748BF3D4110748" bold="true" pageId="0" pageNumber="819">Schletteriinae</emphasis>
</taxonomicName>
</td>
<td id="D6DE8E370D4CAA5F3915AA8BBACA0E49" colspan="1" pageId="0" pageNumber="819" rowspan="1">
<taxonomicName id="B3A397A6572A6F3E29167D142782175E" authorityName="Leach" authorityYear="1815" lsidName="" pageId="0" pageNumber="819" rank="subFamily" subFamily="Stephaninae">
<emphasis id="E7188E067538B711CB73E2F05885BFAD" bold="true" pageId="0" pageNumber="819">Stephaninae</emphasis>
</taxonomicName>
</td>
</tr>
<tr id="1CCD44393629B6E89CB9F21165F2EE65" pageId="0" pageNumber="819">
<td id="FAA40EDF45A8620C6862FEA8D9F5AC14" colspan="1" pageId="0" pageNumber="819" rowspan="1">Anterior part of pronotum</td>
<td id="C2EEBB3BB22AA2B3B9FB5F9A20A4546A" colspan="1" pageId="0" pageNumber="819" rowspan="1">
Elongated as
<normalizedToken id="97D050BBB547C34FD936ED636330E49C" originalValue="“neck”">&quot;neck&quot;</normalizedToken>
</td>
<td id="39E486D8CF40251FB3757CEF750A61C9" colspan="1" pageId="0" pageNumber="819" rowspan="1">
Not elongated as
<normalizedToken id="03DFB6C05D50E6285D22EE5F0F8D1252" originalValue="“neck”">&quot;neck&quot;</normalizedToken>
</td>
<td id="7D103541174C4E06BBA21C0ED81A6D21" colspan="1" pageId="0" pageNumber="819" rowspan="1">
Elongated as
<normalizedToken id="A7AE7C57894D6802B333AA427DEEE4D3" originalValue="“neck”">&quot;neck&quot;</normalizedToken>
</td>
</tr>
<tr id="48F5DBD18BB66E370F46DDF054DDBC7B" pageId="0" pageNumber="819">
<td id="9955C52052A0C07CD0E51075F0E990BD" colspan="1" pageId="0" pageNumber="819" rowspan="1">Hindwing with vein Cu-a</td>
<td id="65F50546EBAA4E7B5566859A4BE01215" colspan="1" pageId="0" pageNumber="819" rowspan="1">Absent</td>
<td id="D7FD857219D8E8B669ACE4B5B9167DB4" colspan="1" pageId="0" pageNumber="819" rowspan="1">Present</td>
<td id="F65AE284ACAC14D6D3CC93162EAD22B8" colspan="1" pageId="0" pageNumber="819" rowspan="1">Absent</td>
</tr>
<tr id="71CC77B4A70334E6607FE299BDB3CF81" pageId="0" pageNumber="819">
<td id="EB69536FDE53313963F0AE1667248BD0" colspan="1" pageId="0" pageNumber="819" rowspan="1">Forewing with the junction of 2-Rs, M, and 2Rs+M</td>
<td id="CADBA1F311459E08B311ECB109C7DF55" colspan="1" pageId="0" pageNumber="819" rowspan="1">Distinctly separated</td>
<td id="7B964E5CA0DBAD13C10A0E7ED5576FA5" colspan="1" pageId="0" pageNumber="819" rowspan="1">Distinctly separated</td>
<td id="B2C638158B953AD4BA7425FE277373B8" colspan="1" pageId="0" pageNumber="819" rowspan="1">Variable</td>
</tr>
<tr id="31C760000647EDFC2E76FB5CF7315655" pageId="0" pageNumber="819">
<td id="5345DDAB1288AF6113ED277BFC64E427" colspan="1" pageId="0" pageNumber="819" rowspan="1">Hind coxa with surface sculpture</td>
<td id="937C4271BF06B78EBF3D3E398FE68D4C" colspan="1" pageId="0" pageNumber="819" rowspan="1">Largely smooth</td>
<td id="2E1F4D6052100E82AB71E075DA7D95A1" colspan="1" pageId="0" pageNumber="819" rowspan="1">Rugose</td>
<td id="DEDA783A7B20A17ECFC347BD48248A26" colspan="1" pageId="0" pageNumber="819" rowspan="1">Variable</td>
</tr>
<tr id="CB8BF2F8CC4D01CC98C07EB75DD2F525" pageId="0" pageNumber="819">
<td id="A198188A28DD76E0829102A2000D160E" colspan="1" pageId="0" pageNumber="819" rowspan="1">Hind coxa with dorsal teeth</td>
<td id="F9F444E38A5C4D59C028FB89F0DB5529" colspan="1" pageId="0" pageNumber="819" rowspan="1">Absent</td>
<td id="29D4396B79FC339F51519201CBEB468D" colspan="1" pageId="0" pageNumber="819" rowspan="1">Present</td>
<td id="C016EA69E025043D67EE2A8A0FF45306" colspan="1" pageId="0" pageNumber="819" rowspan="1">Absent</td>
</tr>
<tr id="11CBA5185866952D02DE77F2D4B788CE" pageId="0" pageNumber="819">
<td id="628EFB5F0C3E02AE33526753A51DA951" colspan="1" pageId="0" pageNumber="819" rowspan="1">Hind tarsus of females</td>
<td id="6772C764336A73669788E17E95564451" colspan="1" pageId="0" pageNumber="819" rowspan="1">5-segmented</td>
<td id="1DC4E67CBCB882E034433EF914FD942E" colspan="1" pageId="0" pageNumber="819" rowspan="1">5-segmented</td>
<td id="30F3002E4E7756A6FC6D14E32DA0DA1C" colspan="1" pageId="0" pageNumber="819" rowspan="1">Variable</td>
</tr>
<tr id="753ECB3BC20DE19404AEB259D62D2C97" pageId="0" pageNumber="819">
<td id="4F17C7F67A60001497013F34C42C0557" colspan="1" pageId="0" pageNumber="819" rowspan="1">Metasoma with tergum I and sternum I</td>
<td id="3952545478F9E7611875EE40ABC25F93" colspan="1" pageId="0" pageNumber="819" rowspan="1">Variable</td>
<td id="FE86537BE221D91BF9B61E422FDEE25F" colspan="1" pageId="0" pageNumber="819" rowspan="1">Separated</td>
<td id="A4425FC5B4652BA5B9089663DC1F6664" colspan="1" pageId="0" pageNumber="819" rowspan="1">Variable</td>
</tr>
<tr id="78F7A370D42A7F2752C843E4623B214F" pageId="0" pageNumber="819">
<td id="2FB80175305566385BA5002CFD0AE4A8" colspan="1" pageId="0" pageNumber="819" rowspan="1">Length of ovipositor</td>
<td id="773A2D46B128E07F6629EF8C4B7B28FB" colspan="1" pageId="0" pageNumber="819" rowspan="1">Approximately as long as metasoma</td>
<td id="03A7AD65D6D3C4C84338FA5A2E041172" colspan="1" pageId="0" pageNumber="819" rowspan="1">Approximately as long as body</td>
<td id="95328A0F695D9F68239DBB2F316A2F5F" colspan="1" pageId="0" pageNumber="819" rowspan="1">Approximately as long as body</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection id="1F8E967B5FE798EF9EE1FE76D9E5904B" pageId="0" pageNumber="819" type="comments">
<paragraph id="D8A4D8673170B07229EA9889B2018C81" pageId="0" pageNumber="819">Comments.</paragraph>
<paragraph id="9F278EDC158DF3FD9622E91D58BEDF38" pageId="0" pageNumber="819">
The phylogenetic results indicated that the new subfamily forms a monophyletic group that sister to
<taxonomicName id="DBF9BE8157CB68F6C838DBACEACDCCCC" authorityName="Orfila" authorityYear="1949" lsidName="" pageId="0" pageNumber="819" rank="subFamily" subFamily="Schlettereriinae">Schlettereriinae</taxonomicName>
. When the characters are mapped on the tree, the new subfamily does not possess any unique synapomorphy, while its sister relationship to
<taxonomicName id="5DDA393F637372D14E934FE67E1047D6" authorityName="Orfila" authorityYear="1949" lsidName="" pageId="0" pageNumber="819" rank="subFamily" subFamily="Schlettereriinae">Schlettereriinae</taxonomicName>
is supported by a synapomorphy (character 34: 0). This is caused by the male †
<taxonomicName id="B55105923AB212505B66ADBDB0D92A28" family="Stephanidae" genus="Neurastephanus" kingdom="Animalia" lsidName="Neurastephanus neovenatus" order="Hymenoptera" pageId="0" pageNumber="819" rank="species" species="neovenatus">
<emphasis id="36B1A55DE5715A4BBE8B94016EB440A5" italics="true" pageId="0" pageNumber="819">Neurastephanus neovenatus</emphasis>
</taxonomicName>
which is revealed as member of †
<taxonomicName id="A2131998146DD24965FF6D0265BF3C6B" authorityName="Ge and Tan" authorityYear="2023" family="Stephanidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="819" rank="subFamily" subFamily="Lagenostephaninae">Lagenostephaninae</taxonomicName>
based on forewing with vein 1-M 1.4-1.7
<normalizedToken id="B47DC9B9568F8C6BB433C72E299A80F9" originalValue="×">x</normalizedToken>
as long as vein 1m-cu (character 38: 1), the largely smooth hind coxa (character 45: 0),the forewing with vein 2Rs+M non-connecting to 2-Rs (character 34: 0), the apical abscissa of vein M (character 23: 1), and vein Rs+M of fore wing converging to vein 1Cu distally (character 36: 1). All above characters are homoplastic within
<taxonomicName id="6E587E31291F3793A3F6097E031B291F" class="Insecta" family="Stephanidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="819" phylum="Arthropoda" rank="family">Stephanidae</taxonomicName>
. The length of ovipositor (characters 64), which is short in †
<taxonomicName id="494D860717EE47476ABF80C7D2F039BD" authorityName="Ge and Tan" authorityYear="2023" family="Stephanidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="819" rank="subFamily" subFamily="Lagenostephaninae">Lagenostephaninae</taxonomicName>
and represents the unique synapomorphy of the subfamily, cannot be coded for †
<taxonomicName id="2B7AF8CDF1529656197438600AB4CDDA" family="Stephanidae" genus="Neurastephanus" kingdom="Animalia" lsidName="Neurastephanus neovenatus" order="Hymenoptera" pageId="0" pageNumber="819" rank="species" species="neovenatus">
<emphasis id="A308359086A35DA4D810E8EF90DED6FB" italics="true" pageId="0" pageNumber="819">Neurastephanus neovenatus</emphasis>
</taxonomicName>
and hence cannot be mapped in Fig.
<figureCitation id="1696A624F578658CFC59496053DE81CA" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A strict consensus tree was obtained by implied weighting (IW). Solid bullets (●) indicate non-homoplastic synapomorphies; open bullets (○) indicate homoplastic characters. Bootstrap values (BS, shown as ' - ' if absent) and Bremer support values (BR, shown as ' * ' if absent) are separated by a slash ' / ' and marked beside each node. A O. robusta Jouault, Rasnitsyn and Perrichot; B T. hui Engel; C L. lii Li, Rasnitsyn, Shih and Ren; D S. chundanae Tan and van Achterberg; E S. anijimensis Watanabe and van Achterberg; F P. notiochinensis Tan and van Achterberg; G F. achterbergi Gupta and Gawas." figureDoi="10.3897/asp.81.e107579.figure6" httpUri="https://binary.pensoft.net/fig/925966" pageId="0" pageNumber="819">6</figureCitation>
. Intriguingly, based on the comparatively short ovipositor and small body sizes of †
<taxonomicName id="4A6E408E502046B4AFFF80E209746004" authorityName="Ge and Tan" authorityYear="2023" family="Stephanidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="819" rank="subFamily" subFamily="Lagenostephaninae">Lagenostephaninae</taxonomicName>
, we speculate that these may reflect a unique ecological niche of the lineage (e.g. possible parasitism of bark beetles) different from
<taxonomicName id="CCC2E91656AE7809FB3B38E30E7DF51C" authorityName="Orfila" authorityYear="1949" lsidName="" pageId="0" pageNumber="819" rank="subFamily" subFamily="Schlettereriinae">Schlettereriinae</taxonomicName>
that are parasitoids of xylem borers. We believe it is hence reasonable to establish this lineage as a new subfamily rather than combine it with
<taxonomicName id="54C9BC84B719AA74E0461E89D435C05D" authorityName="Orfila" authorityYear="1949" lsidName="" pageId="0" pageNumber="819" rank="subFamily" subFamily="Schlettereriinae">Schlettereriinae</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>
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