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<document id="3EC097315965707194A79E14B0B09542" ID-DOI="10.11646/zootaxa.5501.3.3" ID-ISSN="1175-5326" ID-Zenodo-Dep="13628197" ID-ZooBank="F6A55A77-C4C8-4BDD-8D36-482395A01311" IM.bibliography_approvedBy="karina" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="karina" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="karina" checkinTime="1725278975848" checkinUser="plazi" docAuthor="Nagel, Peter" docDate="2024" docId="03F01B1CFFF46D61A0EE5D0E1787FBF0" docLanguage="en" docName="zootaxa.5501.3.3.pdf" docOrigin="Zootaxa 5501 (3)" docSource="http://dx.doi.org/10.11646/zootaxa.5501.3.3" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Paussus lanxangensis Nagel 2009" docType="treatment" docVersion="3" lastPageNumber="455" masterDocId="FFC96364FFF36D69A0795F731654FFC5" masterDocTitle="The Paussus (Scaphipaussus) lanxangensis species group of Indomalayan ant nest beetles (Coleoptera, Carabidae, Paussinae, Paussini) with description of a new species" masterLastPageNumber="457" masterPageNumber="447" pageNumber="454" updateTime="1725501495294" updateUser="karina" zenodo-license-document="CLOSED">
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<mods:title id="AA98455B9284D98CF0F64AB1CE02B28A">The Paussus (Scaphipaussus) lanxangensis species group of Indomalayan ant nest beetles (Coleoptera, Carabidae, Paussinae, Paussini) with description of a new species</mods:title>
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<subSubSection id="C343F981FFF46D6EA0EE5D0E14DDFD52" box="[151,649,637,664]" pageId="7" pageNumber="454" type="nomenclature">
<paragraph id="8BE6AA0AFFF46D6EA0EE5D0E14DDFD52" blockId="7.[151,649,637,664]" box="[151,649,637,664]" pageId="7" pageNumber="454">
<heading id="D0AE1D66FFF46D6EA0EE5D0E14DDFD52" bold="true" box="[151,649,637,664]" fontSize="11" level="1" pageId="7" pageNumber="454" reason="1">
<emphasis id="B92D7618FFF46D6EA0EE5D0E14DDFD52" bold="true" box="[151,649,637,664]" pageId="7" pageNumber="454">
The
<taxonomicName id="4C59D189FFF46D6EA0B65D0D178EFD5D" ID-CoL="766YZ" authorityName="Nagel" authorityYear="2009" box="[207,474,637,664]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="lanxangensis">
<emphasis id="B92D7618FFF46D6EA0B65D0D178EFD5D" bold="true" box="[207,474,637,664]" italics="true" pageId="7" pageNumber="454">Paussus lanxangensis</emphasis>
</taxonomicName>
species group
</emphasis>
</heading>
</paragraph>
</subSubSection>
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<paragraph id="8BE6AA0AFFF46D6EA0EE5DB512DEFCAA" blockId="7.[151,1437,709,1168]" pageId="7" pageNumber="454">
Important morphological structures just as the ligula or the lacinia could not be studied because only a singleton (
<taxonomicName id="4C59D189FFF46D6EA0D85D99175DFCC6" authorityName="Nagel" authorityYear="2024" box="[161,265,746,771]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="agatae" status="sp.n.">
<emphasis id="B92D7618FFF46D6EA0D85D99175DFCC6" box="[161,265,746,771]" italics="true" pageId="7" pageNumber="454">P. agatae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21ECB63FFF46D6EA1685D9A1712FCC6" box="[273,326,745,771]" pageId="7" pageNumber="454" rank="species">sp.n.</taxonomicNameLabel>
) or very few specimens (
<taxonomicName id="4C59D189FFF46D6EA2115D991543FCC6" authorityName="Nagel" authorityYear="2009" box="[616,791,745,771]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="lanxangensis">
<emphasis id="B92D7618FFF46D6EA2115D991543FCC6" box="[616,791,745,771]" italics="true" pageId="7" pageNumber="454">P. lanxangensis</emphasis>
</taxonomicName>
male
<typeStatus id="54E214A8FFF46D6EA3245D9A15EBFCC6" box="[861,959,745,771]" pageId="7" pageNumber="454" type="holotype">holotype</typeStatus>
and one single female) were available for the present treatise. Destructive dissection was avoided. Other characters of the mouthparts (maxillary palps, labial palps) could not be used for all species in this study because the original descriptions of
<taxonomicName id="4C59D189FFF46D6EA4075C41171BFCAA" authority="(Maruyama &amp; Nakase 2023)" baseAuthorityName="Maruyama &amp; Nakase" baseAuthorityYear="2023" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="wakaharai">
<emphasis id="B92D7618FFF46D6EA4075C411346FC8E" box="[1150,1298,817,843]" italics="true" pageId="7" pageNumber="454">P. wakaharai</emphasis>
(
<bibRefCitation id="EFC8D7FBFFF46D6EA5585C421713FCAA" author="Maruyama, M. &amp; Nakase, Y." pageId="7" pageNumber="454" pagination="1 - 3" refId="ref7360" refString="Maruyama, M. &amp; Nakase, Y. (2023) A New Species of the Genus Paussus Linne, 1775 (Coleoptera: Carabidae: Paussinae) from Laos, Allied to P. lanxangensis Nagel, 2009. Japanese Journal of Systematic Entomology, 29 (1), 1 - 3." type="journal article" year="2023">Maruyama &amp; Nakase 2023</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName id="4C59D189FFF46D6EA1FC5C251542FCAA" authority="(Bednarik &amp; Bocak 2023)" baseAuthorityName="Bednarik &amp; Bocak" baseAuthorityYear="2023" box="[389,790,853,879]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="haucki">
<emphasis id="B92D7618FFF46D6EA1FC5C2517B8FCAA" box="[389,492,853,879]" italics="true" pageId="7" pageNumber="454">P. haucki</emphasis>
(
<bibRefCitation id="EFC8D7FBFFF46D6EA1835C26155BFCAA" author="Bednarik, M. &amp; Bocak, L." box="[506,783,853,879]" pageId="7" pageNumber="454" pagination="1 - 30" refId="ref6548" refString="Bednarik, M. &amp; Bocak, L. (2023) New species of Paussus, Subgenus Scaphipaussus (Coleoptera: Carabidae: Paussinae), from Southeast Asia Reveal Ambiguities in Species Group Limits and High Species Diversity in the Oriental Region. Insects, 14 (947), 1 - 30. https: // doi. org / 10.3390 / insects 14120947" type="journal article" year="2023">Bednařik &amp; Bocak 2023</bibRefCitation>
)
</taxonomicName>
did not provide this information.
</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE5C0A14EFFB82" blockId="7.[151,1437,709,1168]" pageId="7" pageNumber="454">
The text and illustration of the original description of
<taxonomicName id="4C59D189FFF46D6EA33B5C09159CFC56" authorityName="Maruyama" authorityYear="2014" box="[834,968,889,915]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="drumonti">
<emphasis id="B92D7618FFF46D6EA33B5C09159CFC56" box="[834,968,889,915]" italics="true" pageId="7" pageNumber="454">P. drumonti</emphasis>
</taxonomicName>
describes the maxillary palpomere II as “large, compressed” (
<bibRefCitation id="EFC8D7FBFFF46D6EA1F35CEE1413FC72" author="Maruyama, M." box="[394,583,925,951]" pageId="7" pageNumber="454" pagination="27 - 31" refId="ref7256" refString="Maruyama, M. (2014) Two New Species of the Genus Paussus Linne, 1775 (Coleoptera, Carabidae, Paussinae) from Thailand. Esakia, 54, 27 - 31. https: // doi. org / 10.5109 / 1516264" type="journal article" year="2014">Maruyama 2014</bibRefCitation>
) but does not inform if its “mesal margin ... is strongly rounded or more or less straight” as in
<taxonomicName id="4C59D189FFF46D6EA1125CB21444FC1E" authorityName="Fowler" authorityYear="1912" box="[363,528,961,987]" pageId="7" pageNumber="456" rank="subGenus" subGenus="Scaphipaussus">
<emphasis id="B92D7618FFF46D6EA1125CB21444FC1E" box="[363,528,961,987]" italics="true" pageId="7" pageNumber="454">Scaphipaussus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFC8D7FBFFF46D6EA2675CB21517FC1E" author="Robertson, J. A. &amp; Moore, W." box="[542,835,961,988]" pageId="7" pageNumber="454" pagination="134 - 170" refId="ref7784" refString="Robertson, J. A. &amp; Moore, W. (2016) Phylogeny of Paussus L. (Carabidae: Paussinae): unravelling morphological convergence associated with myrmecophilous life histories. Systematic Entomology, 42 (1), 134 - 170, figs. S 1 - S 2, tabs S 1 - S 5. [2017, DP 28 Sep 2016, version of record online] https: // doi. org / 10.1111 / syen. 12205" type="journal article" year="2016">Robertson &amp; Moore 2016</bibRefCitation>
, pp. 29, 32, 33). The importance of several characters became only known as a result of the recent revision of
<taxonomicName id="4C59D189FFF46D6EA34F5C9515C6FC3A" box="[822,914,998,1023]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="B92D7618FFF46D6EA34F5C9515C6FC3A" box="[822,914,998,1023]" italics="true" pageId="7" pageNumber="454">Paussus</emphasis>
</taxonomicName>
L. (
<bibRefCitation id="EFC8D7FBFFF46D6EA3B15C9612AFFC3A" author="Robertson, J. A. &amp; Moore, W." box="[968,1275,997,1024]" pageId="7" pageNumber="454" pagination="134 - 170" refId="ref7784" refString="Robertson, J. A. &amp; Moore, W. (2016) Phylogeny of Paussus L. (Carabidae: Paussinae): unravelling morphological convergence associated with myrmecophilous life histories. Systematic Entomology, 42 (1), 134 - 170, figs. S 1 - S 2, tabs S 1 - S 5. [2017, DP 28 Sep 2016, version of record online] https: // doi. org / 10.1111 / syen. 12205" type="journal article" year="2016">Robertson &amp; Moore 2016</bibRefCitation>
). Publications that were issued earlier or contemporarily with the revision could not yet consider these criteria and their authors described the standard traits of the taxon instead.
</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE5B21142FFB4A" blockId="7.[151,1437,709,1168]" pageId="7" pageNumber="454">I use the term “species group” for an informal, phenotypically denoted taxon. The term clade is reserved for a prospective proven monophylum (lineage).</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0EE5B9215F3FB3E" blockId="7.[151,935,1249,1275]" box="[151,935,1249,1275]" pageId="7" pageNumber="454">
<emphasis id="B92D7618FFF46D6EA0EE5B9215F3FB3E" bold="true" box="[151,935,1249,1275]" pageId="7" pageNumber="454">Description of the group (diagnostic, phenotypical characters):</emphasis>
</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0EE5A5A14DBFA81" blockId="7.[151,1437,1321,2032]" box="[151,655,1321,1348]" pageId="7" pageNumber="454">Small to very small species, BL 3.8–5.0 mm.</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE5A3E16B1FA75" blockId="7.[151,1437,1321,2032]" pageId="7" pageNumber="454">Head wider than long. Head approximately of same width as the pronotum, or little or considerably narrower than pronotum. Eyes small, temples large. Head with stigmatoid vestige of a short elongate opening at medial vertex.</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE5ACA1364FA32" blockId="7.[151,1437,1321,2032]" pageId="7" pageNumber="454">Antennal club strongly inflated to oblong lens-shaped, posteriorly with roundish to elliptic cavity or with narrow longitudinal furrow. Basal margin uninterrupted, without central incision or semi-transparent impression.</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE5972156DF9DE" blockId="7.[151,1437,1321,2032]" box="[199,825,1537,1564]" pageId="7" pageNumber="454">Terminal labial palpomeres not distinctly spade-shaped.</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE5955124AF942" blockId="7.[151,1437,1321,2032]" pageId="7" pageNumber="454">Pronotum with deep transverse furrow with small dorsolateral trichomes. Anterior part of pronotum broader and much higher than posterior part. Posterior part of pronotum deeply excavated in the middle. Two prominent tubercles behind the transverse excavation at both sides of a longitudinal furrow.</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE59E216B2F936" blockId="7.[151,1437,1321,2032]" pageId="7" pageNumber="454">
<taxonomicName id="4C59D189FFF46D6EA0BE59E21760F96E" box="[199,308,1681,1707]" class="Actinopterygii" family="Trichomycteridae" genus="Pygidium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Siluriformes" pageId="7" pageNumber="451" phylum="Chordata" rank="genus">Pygidium</taxonomicName>
excavated, with posterior dorsal or latero-dorsal margins explanate, sometimes blade-like and upturned, and bent inwardly in some species. Pygidial disc even, smooth, and glabrous. Marginal pygidial trichome absent.
</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE598E129EF8FE" blockId="7.[151,1437,1321,2032]" pageId="7" pageNumber="454">
Tibiae slightly compressed, not edged, margins rounded. Tibiae widened towards apex, exterior apical angle not or little pronounced, apically rounded, rarely rectangular (
<taxonomicName id="4C59D189FFF46D6EA35B585115F0F8FE" authorityName="Maruyama" authorityYear="2014" box="[802,932,1825,1851]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="drumonti">
<emphasis id="B92D7618FFF46D6EA35B585115F0F8FE" box="[802,932,1825,1851]" italics="true" pageId="7" pageNumber="454">P. drumonti</emphasis>
</taxonomicName>
excepted, see key above).
</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE58361561F89A" blockId="7.[151,1437,1321,2032]" box="[199,821,1861,1887]" pageId="7" pageNumber="454">Elytral lateral trichome (serially arranged setae) absent.</paragraph>
<paragraph id="8BE6AA0AFFF46D6EA0BE581A1571F82A" blockId="7.[151,1437,1321,2032]" pageId="7" pageNumber="454">
The group lacks falciform ridges or crests at the dorsal head. It also lacks elytral mirror markings (present at most species of the
<taxonomicName id="4C59D189FFF46D6EA1D858FD1542F862" authorityName="Westwood" authorityYear="1850" box="[417,790,1933,1959]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="hystrix" subGenus="Scaphipaussus">
<emphasis id="B92D7618FFF46D6EA1D858FD1542F862" box="[417,790,1933,1959]" italics="true" pageId="7" pageNumber="454">Paussus (Scaphipaussus) hystrix</emphasis>
</taxonomicName>
group (
<bibRefCitation id="EFC8D7FBFFF46D6EA30D58FE1262F862" author="Maruyama, M." box="[884,1078,1933,1959]" pageId="7" pageNumber="454" pagination="55 - 86" refId="ref7301" refString="Maruyama, M. (2016) Revision of the hystrix Westwood, 1850 Group of the Genus Paussus Linne, 1775 (Coleoptera: Carabidae: Paussinae). I. Descriptions of Nineteen New Species. Japanese Journal of Systematic Entomology, 22 (1), 55 - 86. [DP 30 May 2016]" type="journal article" year="2016">Maruyama 2016</bibRefCitation>
;
<bibRefCitation id="EFC8D7FBFFF46D6EA43D58FE1336F862" author="Bednarik, M. &amp; Bocak, L." box="[1092,1378,1933,1959]" pageId="7" pageNumber="454" pagination="1 - 30" refId="ref6548" refString="Bednarik, M. &amp; Bocak, L. (2023) New species of Paussus, Subgenus Scaphipaussus (Coleoptera: Carabidae: Paussinae), from Southeast Asia Reveal Ambiguities in Species Group Limits and High Species Diversity in the Oriental Region. Insects, 14 (947), 1 - 30. https: // doi. org / 10.3390 / insects 14120947" type="journal article" year="2023">Bednařik &amp; Bocak 2023</bibRefCitation>
) and the rounded elytral patches as well as the subapical trichome of antennomere 1 (scape) of the
<taxonomicName id="4C59D189FFF46D6EA4B058C1175EF82A" authorityName="Guerin-Meneville" authorityYear="1836" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="454" phylum="Arthropoda" rank="species" species="jousselinii" subGenus="Paussus">
<emphasis id="B92D7618FFF46D6EA4B058C1175EF82A" italics="true" pageId="7" pageNumber="454">Paussus (Paussus) jousselinii</emphasis>
</taxonomicName>
group (
<bibRefCitation id="EFC8D7FBFFF46D6EA11858A617BDF82A" author="Nagel, P." box="[353,489,2005,2031]" pageId="7" pageNumber="454" refId="ref7514" refString="Nagel, P. (2016) Paussus brancuccii sp. nov., an unusually abundant species of the Paussus jousselinii group from Laos and South China (Coleoptera, Carabidae, Paussinae, Paussini). In: Brancucci, M. †, Nagel, P., Kolibac, J. &amp; Geiser, M. (Eds.), Beetle diversity of Laos. Part II. Entomologica Basiliensia et Collectionis Frey. Vol. 35. Pro Entomologia, Basel, pp. 119 - 132. [DP 10 May 2016] https: // doi. org / 10.5169 / seals- 980956" type="book" year="2016">Nagel 2016</bibRefCitation>
;
<bibRefCitation id="EFC8D7FBFFF46D6EA18D58A6154DF82A" author="Robertson, J. A. &amp; Moore, W." box="[500,793,2005,2032]" pageId="7" pageNumber="454" pagination="134 - 170" refId="ref7784" refString="Robertson, J. A. &amp; Moore, W. (2016) Phylogeny of Paussus L. (Carabidae: Paussinae): unravelling morphological convergence associated with myrmecophilous life histories. Systematic Entomology, 42 (1), 134 - 170, figs. S 1 - S 2, tabs S 1 - S 5. [2017, DP 28 Sep 2016, version of record online] https: // doi. org / 10.1111 / syen. 12205" type="journal article" year="2016">Robertson &amp; Moore 2016</bibRefCitation>
).
</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5FE412D3FF74" blockId="8.[151,1437,151,1077]" box="[199,1159,151,177]" pageId="8" pageNumber="455">Distribution: Southeast Asian mainland, and most probably an Indomalayan endemic.</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5FC81585FF3C" blockId="8.[151,1437,151,1077]" pageId="8" pageNumber="455">
The group is characterized by the distinctive characters of the
<taxonomicName id="4C59D189FFFB6D61A3FE5FC815B7FF11" box="[903,995,187,212]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="B92D7618FFFB6D61A3FE5FC815B7FF11" box="[903,995,187,212]" italics="true" pageId="8" pageNumber="455">Paussus</emphasis>
</taxonomicName>
series II (see above) and the following possible common traits (candidates for common derived character states):
</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5E701252FED8" blockId="8.[151,1437,151,1077]" box="[199,1030,259,285]" pageId="8" pageNumber="455">- Head with stigmatoid vestige of short elongate opening at medial vertex.</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5E541316FE84" blockId="8.[151,1437,151,1077]" box="[199,1346,295,321]" pageId="8" pageNumber="455">- Pygidial disc smooth, glabrous and shining, positioned at the bottom of a hollow or large excavation.</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5E381443FE4C" blockId="8.[151,1437,151,1077]" pageId="8" pageNumber="455">- Maxillary palpomere II not subcylindrical but large, explanate, with mesal margin sinuate, faintly emarginate or almost straight, but not convex.</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5EE01718FE30" blockId="8.[151,1437,151,1077]" pageId="8" pageNumber="455">
The group is classified as part of genus
<taxonomicName id="4C59D189FFFB6D61A2EA5EE014BBFE69" box="[659,751,403,428]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="B92D7618FFFB6D61A2EA5EE014BBFE69" box="[659,751,403,428]" italics="true" pageId="8" pageNumber="455">Paussus</emphasis>
</taxonomicName>
L., and as a member of the clade “
<taxonomicName id="4C59D189FFFB6D61A4FF5EE012B6FE69" box="[1158,1250,403,428]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="B92D7618FFFB6D61A4FF5EE012B6FE69" box="[1158,1250,403,428]" italics="true" pageId="8" pageNumber="455">Paussus</emphasis>
</taxonomicName>
series II” in the latest revision (
<bibRefCitation id="EFC8D7FBFFFB6D61A1385EC4143CFE14" author="Robertson, J. A. &amp; Moore, W." box="[321,616,439,465]" pageId="8" pageNumber="455" pagination="134 - 170" refId="ref7784" refString="Robertson, J. A. &amp; Moore, W. (2016) Phylogeny of Paussus L. (Carabidae: Paussinae): unravelling morphological convergence associated with myrmecophilous life histories. Systematic Entomology, 42 (1), 134 - 170, figs. S 1 - S 2, tabs S 1 - S 5. [2017, DP 28 Sep 2016, version of record online] https: // doi. org / 10.1111 / syen. 12205" type="journal article" year="2016">Robertson &amp; Moore 2016</bibRefCitation>
). Most probably, it forms part of the subgenus
<taxonomicName id="4C59D189FFFB6D61A4085EC4133AFE14" authority="Fowler" authorityName="Fowler" authorityYear="1912" box="[1137,1390,439,465]" pageId="8" pageNumber="455" rank="subGenus" subGenus="Scaphipaussus">
<emphasis id="B92D7618FFFB6D61A4085EC41342FE14" box="[1137,1302,439,465]" italics="true" pageId="8" pageNumber="455">Scaphipaussus</emphasis>
Fowler
</taxonomicName>
, see checklist below.
</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5E8C1770FD08" blockId="8.[151,1437,151,1077]" pageId="8" pageNumber="455">
The recent revision (
<bibRefCitation id="EFC8D7FBFFFB6D61A1D55E8C1484FDDC" author="Robertson, J. A. &amp; Moore, W." box="[428,720,511,537]" pageId="8" pageNumber="455" pagination="134 - 170" refId="ref7784" refString="Robertson, J. A. &amp; Moore, W. (2016) Phylogeny of Paussus L. (Carabidae: Paussinae): unravelling morphological convergence associated with myrmecophilous life histories. Systematic Entomology, 42 (1), 134 - 170, figs. S 1 - S 2, tabs S 1 - S 5. [2017, DP 28 Sep 2016, version of record online] https: // doi. org / 10.1111 / syen. 12205" type="journal article" year="2016">Robertson &amp; Moore 2016</bibRefCitation>
) listed two taxa near or in Sg.
<taxonomicName id="4C59D189FFFB6D61A45A5E8C129CFDDC" authorityName="Fowler" authorityYear="1912" box="[1059,1224,511,537]" pageId="8" pageNumber="456" rank="subGenus" subGenus="Scaphipaussus">
<emphasis id="B92D7618FFFB6D61A45A5E8C129CFDDC" box="[1059,1224,511,537]" italics="true" pageId="8" pageNumber="455">Scaphipaussus</emphasis>
</taxonomicName>
for which an exact placement could not be demonstrated with sufficient certainty:
<taxonomicName id="4C59D189FFFB6D61A3085D50128CFDF8" authority="Luna de Carvalho" authorityName="Luna de Carvalho" authorityYear="1973" box="[881,1240,547,573]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="ghanensis">
<emphasis id="B92D7618FFFB6D61A3085D501254FDF8" box="[881,1024,547,573]" italics="true" pageId="8" pageNumber="455">P. ghanensis</emphasis>
Luna de Carvalho
</taxonomicName>
and
<emphasis id="B92D7618FFFB6D61A56D5D501304FDF9" box="[1300,1360,547,572]" italics="true" pageId="8" pageNumber="455">P. nr.</emphasis>
<taxonomicName id="4C59D189FFFB6D61A5205D501737FDA4" authority="Luna de Carvalho" authorityName="Luna de Carvalho" authorityYear="1968" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="leechi">
<emphasis id="B92D7618FFFB6D61A5205D5013C8FDF8" box="[1369,1436,547,573]" italics="true" pageId="8" pageNumber="455">leechi</emphasis>
Luna de Carvalho
</taxonomicName>
were both classified as
<emphasis id="B92D7618FFFB6D61A20E5D341566FDA4" box="[631,818,583,609]" italics="true" pageId="8" pageNumber="455">“incertae sedis”</emphasis>
of the
<emphasis id="B92D7618FFFB6D61A3FC5D341231FDA4" box="[901,1125,583,609]" italics="true" pageId="8" pageNumber="455">
“
<taxonomicName id="4C59D189FFFB6D61A3F65D341258FDA5" authority="II" authorityName="II" box="[911,1036,583,609]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="genus">Paussus II</taxonomicName>
series”
</emphasis>
. I checked specimens of
<taxonomicName id="4C59D189FFFB6D61A5F15D3417C2FD40" authority="Westwood" authorityName="Westwood" authorityYear="1838" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="cochlearius">
<emphasis id="B92D7618FFFB6D61A5F15D34174EFD40" italics="true" pageId="8" pageNumber="455">P. cochlearius</emphasis>
Westwood
</taxonomicName>
(
<collectingCountry id="F34EEA9AFFFB6D61A1DD5D181474FD40" box="[420,544,619,645]" name="Zimbabwe" pageId="8" pageNumber="455">Zimbabwe</collectingCountry>
, CN) (part of the same species group as
<taxonomicName id="4C59D189FFFB6D61A39A5D181215FD40" authorityName="Luna de Carvalho" authorityYear="1968" box="[995,1089,619,645]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="leechi">
<emphasis id="B92D7618FFFB6D61A39A5D181215FD40" box="[995,1089,619,645]" italics="true" pageId="8" pageNumber="455">P. leechi</emphasis>
</taxonomicName>
) and
<taxonomicName id="4C59D189FFFB6D61A4F95D18135FFD40" authorityName="Luna de Carvalho" authorityYear="1973" box="[1152,1291,619,645]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="ghanensis">
<emphasis id="B92D7618FFFB6D61A4F95D18135FFD40" box="[1152,1291,619,645]" italics="true" pageId="8" pageNumber="455">P. ghanensis</emphasis>
</taxonomicName>
(
<collectingCountry id="F34EEA9AFFFB6D61A5635D181335FD40" box="[1306,1377,619,645]" name="Benin" pageId="8" pageNumber="455">Benin</collectingCountry>
, CN) for characters that possibly indicate similarity beyond phenotypy with the
<taxonomicName id="4C59D189FFFB6D61A3935DFC12CDFD6C" authorityName="Nagel" authorityYear="2009" box="[1002,1177,655,681]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="lanxangensis">
<emphasis id="B92D7618FFFB6D61A3935DFC12CDFD6C" box="[1002,1177,655,681]" italics="true" pageId="8" pageNumber="455">P. lanxangensis</emphasis>
</taxonomicName>
species group, yet did not find any.
</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5DA41596FC98" blockId="8.[151,1437,151,1077]" pageId="8" pageNumber="455">
Equally small
<taxonomicName id="4C59D189FFFB6D61A11E5DA41797FD35" box="[359,451,727,752]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="B92D7618FFFB6D61A11E5DA41797FD35" box="[359,451,727,752]" italics="true" pageId="8" pageNumber="455">Paussus</emphasis>
</taxonomicName>
exist in other species groups and subgenera, such as the
<taxonomicName id="4C59D189FFFB6D61A44B5DA412EBFD34" authorityName="Guerin-Meneville" authorityYear="1836" box="[1074,1215,727,753]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="jousselinii">
<emphasis id="B92D7618FFFB6D61A44B5DA412EBFD34" box="[1074,1215,727,753]" italics="true" pageId="8" pageNumber="455">P. jousselinii</emphasis>
</taxonomicName>
group (Sg.
<taxonomicName id="4C59D189FFFB6D61A5395DA413C8FD35" box="[1344,1436,727,752]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="B92D7618FFFB6D61A5395DA413C8FD35" box="[1344,1436,727,752]" italics="true" pageId="8" pageNumber="455">Paussus</emphasis>
</taxonomicName>
L.) (
<taxonomicName id="4C59D189FFFB6D61A0B55D88147FFCD0" authority="Reichensperger, 1950" authorityName="Reichensperger" authorityYear="1950" box="[204,555,762,789]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="wittei">
<emphasis id="B92D7618FFFB6D61A0B55D88177CFCD1" box="[204,296,763,788]" italics="true" pageId="8" pageNumber="455">P. wittei</emphasis>
<bibRefCitation id="EFC8D7FBFFFB6D61A14B5D88147FFCD0" author="Reichensperger, A." box="[306,555,762,789]" pageId="8" pageNumber="455" pagination="1 - 12" refId="ref7734" refString="Reichensperger, A. (1950) Paussidae. In: Exploration du Parc National Albert, Mission G. F. de Witte (1933 - 1935). Fasc. 68. Institut des Parcs Nationaux du Congo Belge, Bruxelles, pp. 1 - 12." type="book chapter" year="1950">Reichensperger, 1950</bibRefCitation>
</taxonomicName>
), the
<taxonomicName id="4C59D189FFFB6D61A2145D8814ABFCD0" authorityName="Westwood" authorityYear="1850" box="[621,767,763,789]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="cucullatus">
<emphasis id="B92D7618FFFB6D61A2145D8814ABFCD0" box="[621,767,763,789]" italics="true" pageId="8" pageNumber="455">P. cucullatus</emphasis>
</taxonomicName>
group (Sg.
<taxonomicName id="4C59D189FFFB6D61A3F25D8812F3FCD0" authority="Dalman, 1823" authorityName="Dalman" authorityYear="1823" box="[907,1191,763,789]" class="Insecta" family="Carabidae" genus="Hylotorus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="B92D7618FFFB6D61A3F25D8815AFFCD0" box="[907,1019,763,789]" italics="true" pageId="8" pageNumber="455">Hylotorus</emphasis>
<bibRefCitation id="EFC8D7FBFFFB6D61A47C5D8812F3FCD0" author="Dalman, J. W." box="[1029,1191,763,789]" pageId="8" pageNumber="455" refId="ref6672" refString="Dalman, J. W. (1823) Analecta Entomologica. Typis Lindhianis, Holmiae, 7 + 104 + 4 pp. https: // doi. org / 10.5962 / bhl. title. 66069" type="book" year="1823">Dalman, 1823</bibRefCitation>
</taxonomicName>
) (
<taxonomicName id="4C59D189FFFB6D61A4BA5D881751FCFC" authority="Nagel &amp; Rasool" authorityName="Nagel &amp; Rasool" authorityYear="2019" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="minutulus">
<emphasis id="B92D7618FFFB6D61A4BA5D88131BFCD0" box="[1219,1359,763,789]" italics="true" pageId="8" pageNumber="455">P. minutulus</emphasis>
Nagel &amp; Rasool
</taxonomicName>
in
<bibRefCitation id="EFC8D7FBFFFB6D61A1555C6C1400FCFC" author="Abdel-Dayem, M. S. &amp; Elgharbawy, A. A. &amp; Rasool, I. &amp; Nagel, P. &amp; Aldhafer, H. M." box="[300,596,799,825]" pageId="8" pageNumber="455" pagination="93 - 131" refId="ref6470" refString="Abdel-Dayem, M. S., Elgharbawy, A. A., Rasool, I., Nagel, P. &amp; Aldhafer, H. M. (2019) The Carabidae (Coleoptera) of Shada Al- A'Ala Nature Reserve, Southwestern Saudi Arabia, with description of a new species of Paussinae. Zookeys, 812, 93 - 131. https: // doi. org / 10.3897 / zookeys. 812.30937" type="journal article" year="2019">
Abdel-Dayem
<emphasis id="B92D7618FFFB6D61A1AD5C6C145BFCFC" box="[468,527,799,825]" italics="true" pageId="8" pageNumber="455">et al.</emphasis>
2019
</bibRefCitation>
), or the
<taxonomicName id="4C59D189FFFB6D61A2CF5C6C1574FCFC" authorityName="Westwood" authorityYear="1850" box="[694,800,799,825]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="hystrix">
<emphasis id="B92D7618FFFB6D61A2CF5C6C1574FCFC" box="[694,800,799,825]" italics="true" pageId="8" pageNumber="455">P. hystrix</emphasis>
</taxonomicName>
group (Sg.
<taxonomicName id="4C59D189FFFB6D61A3D45C6C1206FCFC" authorityName="Fowler" authorityYear="1912" box="[941,1106,799,825]" pageId="8" pageNumber="456" rank="subGenus" subGenus="Scaphipaussus">
<emphasis id="B92D7618FFFB6D61A3D45C6C1206FCFC" box="[941,1106,799,825]" italics="true" pageId="8" pageNumber="455">Scaphipaussus</emphasis>
</taxonomicName>
) (
<taxonomicName id="4C59D189FFFB6D61A4175C6C13C5FCFC" authority="Maruyama, 2016" authorityName="Maruyama" authorityYear="2016" box="[1134,1425,799,825]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="kecil">
<emphasis id="B92D7618FFFB6D61A4175C6C1295FCFC" box="[1134,1217,799,825]" italics="true" pageId="8" pageNumber="455">P. kecil</emphasis>
<bibRefCitation id="EFC8D7FBFFFB6D61A4B35C6C13C5FCFC" author="Maruyama, M." box="[1226,1425,799,825]" pageId="8" pageNumber="455" pagination="55 - 86" refId="ref7301" refString="Maruyama, M. (2016) Revision of the hystrix Westwood, 1850 Group of the Genus Paussus Linne, 1775 (Coleoptera: Carabidae: Paussinae). I. Descriptions of Nineteen New Species. Japanese Journal of Systematic Entomology, 22 (1), 55 - 86. [DP 30 May 2016]" type="journal article" year="2016">Maruyama, 2016</bibRefCitation>
</taxonomicName>
). Therefore, the small size is not a good diagnostic character of the group.
</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5C1412A3FC0C" blockId="8.[151,1437,151,1077]" pageId="8" pageNumber="455">The group could not yet convincingly be established as a clade, because not all or most species could sufficiently be studied with regard to some of the more relevant morphological characters, such as the ligula or the lacinia. The small number of species and specimens also prevented the application of a proper cladistic analysis.</paragraph>
<paragraph id="8BE6AA0AFFFB6D61A0BE5CA01787FBF0" blockId="8.[151,1437,151,1077]" pageId="8" pageNumber="455">
It could, however, be demonstrated that the
<taxonomicName id="4C59D189FFFB6D61A2C35CA0153DFC28" authorityName="Nagel" authorityYear="2009" box="[698,873,979,1005]" class="Insecta" family="Carabidae" genus="Paussus" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="455" phylum="Arthropoda" rank="species" species="lanxangensis">
<emphasis id="B92D7618FFFB6D61A2C35CA0153DFC28" box="[698,873,979,1005]" italics="true" pageId="8" pageNumber="455">P. lanxangensis</emphasis>
</taxonomicName>
species group is a promising candidate for a new monophylum nested within the subgenus
<taxonomicName id="4C59D189FFFB6D61A2165C8412FAFBD4" authority="sensu Robertson and Moore (2016)" authorityName="Fowler sensu Robertson and Moore" authorityYear="2016" box="[623,1198,1015,1041]" pageId="8" pageNumber="455" rank="subGenus" subGenus="Scaphipaussus">
<emphasis id="B92D7618FFFB6D61A2165C841540FBD4" box="[623,788,1015,1041]" italics="true" pageId="8" pageNumber="455">Scaphipaussus</emphasis>
sensu
<bibRefCitation id="EFC8D7FBFFFB6D61A31B5C8412FAFBD4" author="Robertson, J. A. &amp; Moore, W." box="[866,1198,1015,1041]" pageId="8" pageNumber="455" pagination="134 - 170" refId="ref7784" refString="Robertson, J. A. &amp; Moore, W. (2016) Phylogeny of Paussus L. (Carabidae: Paussinae): unravelling morphological convergence associated with myrmecophilous life histories. Systematic Entomology, 42 (1), 134 - 170, figs. S 1 - S 2, tabs S 1 - S 5. [2017, DP 28 Sep 2016, version of record online] https: // doi. org / 10.1111 / syen. 12205" type="journal article" year="2016">Robertson and Moore (2016)</bibRefCitation>
</taxonomicName>
. Future research will have to consider this aspect.
</paragraph>
</subSubSection>
</treatment>
</document>
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