477 lines
74 KiB
XML
477 lines
74 KiB
XML
<document ID-CLB-Dataset="284558" ID-DOI="http://dx.doi.org/10.3897/subtbiol.48.114404" ID-GBIF-Dataset="d0238341-4407-4d11-b1f7-0a4f7d4b22e4" ID-Pensoft-Pub="1314-2615-48-1" ID-Pensoft-UUID="768E4D6A499F5E8DB5FA59ACA071AC36" ID-ZooBank="989B4A14F7D54805822A2181DE5223A4" ModsDocID="1314-2615-48-1" checkinTime="1704767870154" checkinUser="pensoft" docAuthor="Harden, Curt W., Davidson, Robert L., Malabad, Thomas E., Caterino, Michael S. & Maddison, David R." docDate="2024" docId="8AFCA86F53155A0AAACA7564D17925B8" docLanguage="en" docName="SubterBiol 48: 1-49" docOrigin="Subterranean Biology 48" docPubDate="2024-01-08" docSource="http://dx.doi.org/10.3897/subtbiol.48.114404" docTitle="Horologion hubbardi Harden & Davidson 2024, sp. nov." docType="treatment" docUuid="F34C3E90-0136-4D25-A5B0-3E1E384BAF12" docUuidSource="ZooBank" docVersion="2" id="768E4D6A499F5E8DB5FA59ACA071AC36" lastPageNumber="1" masterDocId="768E4D6A499F5E8DB5FA59ACA071AC36" masterDocTitle="Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia" masterLastPageNumber="49" masterPageNumber="1" pageNumber="1" updateTime="1704768481253" updateUser="ExternalLinkService">
|
||
<mods:mods id="9AE62122F70AB6D2BE789F62099FA9CD" xmlns:mods="http://www.loc.gov/mods/v3">
|
||
<mods:titleInfo id="3D10E22AE37689B0B1D0EF9DA3BC26E6">
|
||
<mods:title id="503F9D7659721CCDEF23402EFF011CA8">Phylogenetic systematics of the enigmatic genus Horologion Valentine, 1932 (Coleoptera, Carabidae, Trechinae, Horologionini), with description of a new species from Bath County, Virginia</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:name id="21E11123AAE5D10138A1B2A6B4EEDFB2" type="personal">
|
||
<mods:role id="6BC970831EDC169C70761AB2830FEB29">
|
||
<mods:roleTerm id="C1AF219F86547D2CD96B6766913BAA9D">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="B568B239D3F2317C6A73F1A11B51B01C">Harden, Curt W.</mods:namePart>
|
||
<mods:nameIdentifier id="4D5D6D753974B01BB7E6416B0D8B0663" type="ORCID">https://orcid.org/0000-0002-5226-8725</mods:nameIdentifier>
|
||
<mods:affiliation id="8923368308123FD5D9578B3E40A6B037">Department of Plant and Environmental Sciences, 277 Poole Agricultural Center, Clemson University, Clemson, South Carolina 29634, USA</mods:affiliation>
|
||
<mods:nameIdentifier id="9837B0055325FCE1DF327DA1A951AF40" type="email">c_har@fastmail.com</mods:nameIdentifier>
|
||
</mods:name>
|
||
<mods:name id="174143171EBADABCDF9BF685FCBA1E11" type="personal">
|
||
<mods:role id="20594C19B5F407E95E3CDF3D02310221">
|
||
<mods:roleTerm id="3B3A8EB69F5C8C10D6F5BD16081B7BE0">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="425E15BF8785CE3A7FA27BDBB8CBE437">Davidson, Robert L.</mods:namePart>
|
||
<mods:affiliation id="2265CEDB649DEE1FE96296CAD9F03814">Department of Invertebrate Zoology, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213, USA</mods:affiliation>
|
||
</mods:name>
|
||
<mods:name id="F14F851707DE80C7061E0BCF57A4EBD9" type="personal">
|
||
<mods:role id="3B6558E54BEC4DEB3B3074005A5BE1C6">
|
||
<mods:roleTerm id="8BD77713119520E0967133AE0D37C7BD">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="27689BB546869CFBEF0CF39B287991F6">Malabad, Thomas E.</mods:namePart>
|
||
<mods:affiliation id="A7BEE12F43E34AABB7D77C064A27217F">Virginia Department of Conservation and Recreation, Division of Natural Heritage, Karst Program, 16 th Floor, 600 East Main Street, Richmond, Virginia, USA</mods:affiliation>
|
||
</mods:name>
|
||
<mods:name id="A61B76B8ECDBBFC48D1A99BBA50866A1" type="personal">
|
||
<mods:role id="ECA21D17179EDAB613FDEC4D949B7883">
|
||
<mods:roleTerm id="B56AB36CF55785A5960A5C2132B12CA9">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="56FBB146600E5BA0DF787E0221B9B226">Caterino, Michael S.</mods:namePart>
|
||
<mods:nameIdentifier id="5E05265A2F8DD7764B32594F5CF99595" type="ORCID">https://orcid.org/0000-0002-2597-5707</mods:nameIdentifier>
|
||
<mods:affiliation id="7C5EBB99F8B268D193C4DD0806425C45">Department of Plant and Environmental Sciences, 277 Poole Agricultural Center, Clemson University, Clemson, South Carolina 29634, USA</mods:affiliation>
|
||
</mods:name>
|
||
<mods:name id="E5A53C80E4CCE9ACF4B0CF53D6CE961B" type="personal">
|
||
<mods:role id="AA0662D2A0E2734463C62A2A0F5DDF18">
|
||
<mods:roleTerm id="6014FA2CBF107F112E8CA75281C2CA8D">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="F6A0F40F4651957213CFB4DD94771B63">Maddison, David R.</mods:namePart>
|
||
<mods:nameIdentifier id="D802B057975E891205E12ABCEEE0FE0C" type="ORCID">https://orcid.org/0000-0002-7152-3824</mods:nameIdentifier>
|
||
<mods:affiliation id="A05A913F3F80633D1E2BE133D35631F5">Department of Integrative Biology, Oregon State University, Corvallis, Oregon 97331, USA</mods:affiliation>
|
||
</mods:name>
|
||
<mods:typeOfResource id="8891ED66CC7FB299EF0AB7380A2F3198">text</mods:typeOfResource>
|
||
<mods:relatedItem id="503C95793706E3024D29AEFFA7477181" type="host">
|
||
<mods:titleInfo id="E0AD6E690323A287DDCA8647E1D2BFED">
|
||
<mods:title id="1BBD145CC8C71CB433AE972FAC6FA7C9">Subterranean Biology</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:part id="492B15C8706402B8088BDD97B9CC7B3E">
|
||
<mods:date id="5D9F1951120BEC0B2D4DB45E2048531B">2024</mods:date>
|
||
<mods:detail id="E6A22C670FB298A11F8B3D09DD6EE1B6" type="pubDate">
|
||
<mods:number id="BF158BC8B57802D0CEDF6B9110387125">2024-01-08</mods:number>
|
||
</mods:detail>
|
||
<mods:detail id="D0107240651BFC7E011F5F72DF63C5DD" type="volume">
|
||
<mods:number id="0A1FDD6F13F04832F09E0A34A756E81F">48</mods:number>
|
||
</mods:detail>
|
||
<mods:extent id="7341C368AD04BC1933EE2309396007C3" unit="page">
|
||
<mods:start id="9E39E754BB550233BACB0E3CAC4AC73F">1</mods:start>
|
||
<mods:end id="1FFA56959A34F61BB036AE887C7F3C05">49</mods:end>
|
||
</mods:extent>
|
||
</mods:part>
|
||
</mods:relatedItem>
|
||
<mods:location id="A0489AFC4C33762E5DD20D8E8057DB8D">
|
||
<mods:url id="F2E3DA0E0355BB02C116680C83355DAA">http://dx.doi.org/10.3897/subtbiol.48.114404</mods:url>
|
||
</mods:location>
|
||
<mods:classification id="3EC46C755746ED6C5EF1718A30885604">journal article</mods:classification>
|
||
<mods:identifier id="9942F41220D0C09606D6929955F0E7CC" type="DOI">http://dx.doi.org/10.3897/subtbiol.48.114404</mods:identifier>
|
||
<mods:identifier id="A3359A53F3EC4AC9A2697BD769FE800A" type="Pensoft-Pub">1314-2615-48-1</mods:identifier>
|
||
<mods:identifier id="A669EBB63C0A3504E337A75FB35EF41B" type="ZooBank">989B4A14F7D54805822A2181DE5223A4</mods:identifier>
|
||
<mods:identifier id="69E5B8A910799E799C2D49576265BB3A" type="Pensoft-UUID">768E4D6A499F5E8DB5FA59ACA071AC36</mods:identifier>
|
||
</mods:mods>
|
||
<subSection id="14601D856B539E30A9EC02DA417190E7" pageId="0" pageNumber="1" type="results">
|
||
<treatment id="8AFCA86F53155A0AAACA7564D17925B8" LSID="urn:lsid:zoobank.org:act:F34C3E90-0136-4D25-A5B0-3E1E384BAF12" httpUri="http://treatment.plazi.org/id/8AFCA86F53155A0AAACA7564D17925B8" lastPageNumber="1" pageId="0" pageNumber="1">
|
||
<subSubSection id="115CEDB342A3D4D9E3B54D9EC22C5FCF" pageId="0" pageNumber="1" type="nomenclature">
|
||
<paragraph id="5A6B4E3EFF24A275CC4E36CDD259B624" pageId="0" pageNumber="1">
|
||
<taxonomicName id="FA1AA579A555B05B7635D7B7DE0A8465" LSID="https://zoobank.org/F34C3E90-0136-4D25-A5B0-3E1E384BAF12" authority="Harden & Davidson" authorityName="Harden & Davidson" authorityYear="2024" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion hubbardi" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="hubbardi" status="sp. nov.">Horologion hubbardi Harden & Davidson</taxonomicName>
|
||
<taxonomicNameLabel id="54EE3507D78DDEA89DA263EA0D8BDE2E" pageId="0" pageNumber="1">sp. nov.</taxonomicNameLabel>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="3ADF3BB2734EB0E1BE9DFAAE2FD74BA6" pageId="0" pageNumber="1" type="description">
|
||
<paragraph id="F4900ABA933AF43310DCB5A119A1E705" pageId="0" pageNumber="1">
|
||
<figureCitation id="D5828C63A0A65971E30FA04496EC513A" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Horologion hubbardi female paratype, voucher CWH- 452 (VMNH), dorsal habitus. Abdomen removed for DNA extraction. Scale bar: 1 mm." figureDoi="10.3897/subtbiol.48.114404.figure2" httpUri="https://binary.pensoft.net/fig/963502" pageId="0" pageNumber="1">Figs 2</figureCitation>
|
||
<figureCitation id="407B486C8CA68E3C01122351949176E9" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Habitat of species of Horologion A surface habitat at Williams Cave, Bath Co., VA B C. Harden (yellow helmet) and T. Malabad (red helmet) in microhabitat of H. hubbardi voucher CWH- 452 in Williams Cave C C. Harden examining live H. hubbardi holotype in Williams Cave D live holotype male of H. hubbardi on surface of drip pool in Williams Cave E surface habitat at Arbuckle Cave, Greenbrier Co., WV F T. Malabad examining flowstone in Arbuckle Cave. Photo credit: C. Harden (A, E), K. Kosic Ficco (B-D), M. Miller (F)." figureDoi="10.3897/subtbiol.48.114404.figure3" httpUri="https://binary.pensoft.net/fig/963503" pageId="0" pageNumber="1">, 3D</figureCitation>
|
||
<figureCitation id="A6F49BB2D0AE34848D282361DD2C162B" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Protarsi of Horologion A H. hubbardi male holotype, dorsal aspect B H. hubbardi female paratype, dorsal aspect C H. speokoites male holotype, dorsal aspect D H. hubbardi male holotype, ventral aspect E H. hubbardi male holotype, dorsal aspect. Scale bars: 0.10 mm." figureDoi="10.3897/subtbiol.48.114404.figure8" httpUri="https://binary.pensoft.net/fig/963508" pageId="0" pageNumber="1">, 8</figureCitation>
|
||
<figureCitation id="76AEB9B02028B9A5CF3BA225DEF4B1E0" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">, 9D-H</figureCitation>
|
||
<figureCitation id="C088150C672D09B5783F66E3594690CB" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">, 10</figureCitation>
|
||
<figureCitation id="6543B6520B9DEB38013C88532CD86A7A" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Elytral characters of Horologion species A H. speokoites holotype, elytra, oblique left lateral aspect B H. hubbardi paratype (CMNH), ditto C H. hubbardi paratype (VMNH), SEM detail of apex of left elytron, ventral aspect D H. hubbardi paratype (VMNH), SEM photo of right elytron, dorsal aspect E H. hubbardi paratype (VMNH), SEM detail of right elytron (area indicated by black box in D). Black arrows: discal setae, white arrows: plica." figureDoi="10.3897/subtbiol.48.114404.figure11" httpUri="https://binary.pensoft.net/fig/963511" pageId="0" pageNumber="1">, 11B-E</figureCitation>
|
||
<figureCitation id="0389CFB1E280824ED6FAFE6601B06584" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Scanning electron micrographs of the forebody of Horologion hubbardi A head of voucher CWH- 452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH- 452, ventral aspect." figureDoi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1">, 12</figureCitation>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="7A6DCCDA13C5E106D7A112B04C1ABD8E" pageId="0" pageNumber="1" type="type material">
|
||
<paragraph id="39F5275596ACD5C631F4E36F6E0C267C" pageId="0" pageNumber="1">Type material.</paragraph>
|
||
<paragraph id="896738E4296F9BE8480FF2612EA969B5" pageId="0" pageNumber="1">
|
||
<emphasis id="61551ED4A6FBADF7D1BFE5F685D9C96E" bold="true" pageId="0" pageNumber="1">
|
||
<emphasis id="75BE825565D1BC481CDE3884C6206D58" italics="true" pageId="0" pageNumber="1">Holotype</emphasis>
|
||
male
|
||
</emphasis>
|
||
(Suppl. material 3: fig. S1B) (CMNH), point mounted, abdominal ventrites and right protibia and protarsus glued to point, genitalia in plastic glycerin microvial pinned beneath specimen labels. Original labels: "USA: VIRGINIA, Bath Co. Williams Cave. 29.March.2023. T. Malabad, C. Harden, K.
|
||
<normalizedToken id="C4A8503D75858312AD0BC10EB25999C8" originalValue="Kosič">Kosic</normalizedToken>
|
||
Ficco. Found floating on pool surface." "Harden DNA Voucher CWH-484
|
||
<emphasis id="92B0C32DDB57B5B9F4518E80832F7B2F" italics="true" pageId="0" pageNumber="1">H.</emphasis>
|
||
m Ext. 12/April/2023 [green-bordered cardstock]" "[QR code] CMNH-IZ 769,132" "HOLOTYPE
|
||
<taxonomicName id="18AA8BD4CA7057016ECEFED25277A5ED" authorityName="Harden & Davidson" authorityYear="2024" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion hubbardi" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="hubbardi">
|
||
<emphasis id="7C3993661730E3E16890D1FC2FDD84F5" italics="true" pageId="0" pageNumber="1">Horologion hubbardi</emphasis>
|
||
</taxonomicName>
|
||
♂ Harden & Davidson [computer printed on red cardstock]".
|
||
<emphasis id="55CC8CE2E0C4AB535FA57C7D8EAB9764" italics="true" pageId="0" pageNumber="1">COI</emphasis>
|
||
GenBank accession: OR500887.
|
||
</paragraph>
|
||
<paragraph id="D4BFD36893A1F225BD929E3AE59CB21C" pageId="0" pageNumber="1">
|
||
<emphasis id="23C57F3EA815EE2D12B6312A816A58BA" bold="true" pageId="0" pageNumber="1">
|
||
<emphasis id="F927170EF6DFE69221BBA553E5B7FC95" italics="true" pageId="0" pageNumber="1">Paratypes</emphasis>
|
||
(n = 4)
|
||
</emphasis>
|
||
: One female (VMNH), point mounted, abdominal ventrites and genitalia in glass glycerin vial pinned beneath specimen, labeled "USA: VIRGINIA, Bath Co. Williams Cave. 2.August.2022. T. Malabad, D. Hubbard, C. Harden. Active on ground near drip pool." "Harden DNA Voucher CWH-452 H. Williams F Ext. 7/August/2022" [green-bordered cardstock]. GenBank: OR505843, OR505933, OR500886, OR503053, OR503061, OR503098, OR503071, OR503063, OR503052.
|
||
</paragraph>
|
||
<paragraph id="72F408FBDDB5346B0E39B88DDD9BA902" pageId="0" pageNumber="1">
|
||
One female (CMNH), point mounted, not dissected, labeled "VIRGINIA: Bath County,
|
||
<normalizedToken id="265BFC812D7001C4158237CA7461B8CB" originalValue="Williams’">Williams'</normalizedToken>
|
||
Cave, Sep 8 1991" "D.A. Hubbard" "THOMAS C. BARR COLLECTION 2011 Acc. No. 38.014" "VANHP #: Hubbard VA: Co: Bath Loc: Williams Cave Date: 8 Sept 91." "[QR code] CMNH-IZ 769,133".
|
||
</paragraph>
|
||
<paragraph id="4DFA0664DCCB0201C7E92BA48AD0D17B" pageId="0" pageNumber="1">
|
||
Two males (VMNH), dry mounted with genitalia in glycerin, missing most of antennae and legs, labeled "USA: VIRGINIA, Bath Co. Williams Cave. 28.March.2023. T Malabad, K.
|
||
<normalizedToken id="C068E28B8670651FAC470931D7FB7382" originalValue="Kosič">Kosic</normalizedToken>
|
||
Ficco, CW Harden. Found dead.".
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="7764F76AA5D2ABC9CFEF8BB068F90615" pageId="0" pageNumber="1" type="materials_examined">
|
||
<paragraph id="D67BA2848AC0E81D4BDE57B18675D7D7" pageId="0" pageNumber="1">Other material.</paragraph>
|
||
<paragraph id="9A4B95CCF2FECDC46C5E0FA330741DB2" pageId="0" pageNumber="1">
|
||
<materialsCitation id="229CDECFA354681B7C76F0E206089AA7" collectingDate="2023-03-29" location="Found" municipality="Williams Cave" specimenCount="three" specimenCount-female="1" specimenCount-male="1">
|
||
Fragments of
|
||
<specimenCount id="2DA24AC5652EAA12C66F5315CE19CFBF" count="3" type="generic">three specimens</specimenCount>
|
||
,
|
||
<specimenCount id="80CB8D4CBACA27034BCD6112542710CD" count="1" type="male">one male</specimenCount>
|
||
,
|
||
<specimenCount id="FEFFCC153ED9E0B14CC743727703F478" count="1" type="female">one female</specimenCount>
|
||
, and one unknown sex, in alcohol vials (VMNH):
|
||
<location id="51AAC0E6B7D47C24DF0BFAA951581F95" LSID="urn:lsid:plazi:treatment:8AFCA86F53155A0AAACA7564D17925B8:51AAC0E6B7D47C24DF0BFAA951581F95" municipality="Williams Cave" name="Found">Found</location>
|
||
dead in or near small pools,
|
||
<collectingMunicipality id="B4EAEB4655579E6AD1209B05C638F44B">Williams Cave</collectingMunicipality>
|
||
,
|
||
<collectingDate id="95D2AE4180FB516426B5DD9FDC11D1E1" value="2023-03-29">29 March 2023</collectingDate>
|
||
</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="4D2938D2FDC1ECBF478B677C943E5A34" pageId="0" pageNumber="1" type="diagnosis">
|
||
<paragraph id="6E6E2F1CC0AAF0096981EDD763E0392A" pageId="0" pageNumber="1">Diagnosis.</paragraph>
|
||
<paragraph id="58D15C9AA1F8AA29879DF34460A8FBCE" pageId="0" pageNumber="1">
|
||
From
|
||
<taxonomicName id="77CEC60826F7EEC2B46896D714EB4869" authorityName="Valentine" authorityYear="1932" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion speokoites" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="speokoites">
|
||
<emphasis id="CA30EEC125E501CBF56E647ACA323144" italics="true" pageId="0" pageNumber="1">Horologion speokoites</emphasis>
|
||
</taxonomicName>
|
||
, this species differs in the following external characters: the elytral humeri have longer carinal shelves that terminate in a sharp, curved spine (Fig.
|
||
<figureCitation id="2FCCF470FFC4C32ECD2E4E8EABF6407B" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">10D</figureCitation>
|
||
); the elytra are flatter (Fig.
|
||
<figureCitation id="6558A9C8AD97AA4C2C0A14329BE4DE7B" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Elytral characters of Horologion species A H. speokoites holotype, elytra, oblique left lateral aspect B H. hubbardi paratype (CMNH), ditto C H. hubbardi paratype (VMNH), SEM detail of apex of left elytron, ventral aspect D H. hubbardi paratype (VMNH), SEM photo of right elytron, dorsal aspect E H. hubbardi paratype (VMNH), SEM detail of right elytron (area indicated by black box in D). Black arrows: discal setae, white arrows: plica." figureDoi="10.3897/subtbiol.48.114404.figure11" httpUri="https://binary.pensoft.net/fig/963511" pageId="0" pageNumber="1">11</figureCitation>
|
||
); and the two basal protarsomeres of males are asymmetrically dilated and spinose on their inner margin (Fig.
|
||
<figureCitation id="C5BF0CD95D241289837ACED60A31C03D" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Protarsi of Horologion A H. hubbardi male holotype, dorsal aspect B H. hubbardi female paratype, dorsal aspect C H. speokoites male holotype, dorsal aspect D H. hubbardi male holotype, ventral aspect E H. hubbardi male holotype, dorsal aspect. Scale bars: 0.10 mm." figureDoi="10.3897/subtbiol.48.114404.figure8" httpUri="https://binary.pensoft.net/fig/963508" pageId="0" pageNumber="1">8A</figureCitation>
|
||
). The male genitalia (Fig.
|
||
<figureCitation id="B57131DF9C7B0A8AB39E8B3695B732C5" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9D-G</figureCitation>
|
||
) also differ from those of
|
||
<taxonomicName id="7D6B6E56C891D42CB1FAC52970733257" lsidName="H. speokoites" pageId="0" pageNumber="1" rank="species" species="speokoites">
|
||
<emphasis id="C711997442E616401DF0BA9484F75F1C" italics="true" pageId="0" pageNumber="1">H. speokoites</emphasis>
|
||
</taxonomicName>
|
||
(Fig.
|
||
<figureCitation id="23BE4447704EB76C2FB54FF68376BAF7" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9A-C</figureCitation>
|
||
): the parameres are smaller and each bear three apical setae, the median lobe is straighter ventrally, not twisted from plane of basal lobes, with a smaller and more symmetrical apex, and the flagellum of the internal sac is slightly longer and without a distinct sinuation.
|
||
</paragraph>
|
||
<caption id="FC508ADB793AE3971093A24138F4AD38" doi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1" start="Figure 10" startId="F10">
|
||
<paragraph id="1F1F2F933EB37D65491FF535CF4E27BE" pageId="0" pageNumber="1">
|
||
<emphasis id="59E8AA4B002B03262625623E4EC837B9" bold="true" pageId="0" pageNumber="1">Figure 10.</emphasis>
|
||
Scanning electron microscope images of
|
||
<taxonomicName id="9143331785B179D966221560A8CE578F" authorityName="Harden & Davidson" authorityYear="2024" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion hubbardi" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="hubbardi">
|
||
<emphasis id="E36026199E913E95B84C1B4B86B14BEE" italics="true" pageId="0" pageNumber="1">Horologion hubbardi</emphasis>
|
||
</taxonomicName>
|
||
female paratype, voucher CWH-452 (VMNH)
|
||
<emphasis id="485406B121A33F87E47BFDD131D05102" bold="true" pageId="0" pageNumber="1">A</emphasis>
|
||
detail of right side of pterothorax, ventral aspect
|
||
<emphasis id="3BA0954BD24163F53ACDE106601B6410" bold="true" pageId="0" pageNumber="1">B</emphasis>
|
||
mesoventrite, ventral aspect
|
||
<emphasis id="67914E3F1902FB99F2BA5340FAE02E4D" bold="true" pageId="0" pageNumber="1">C</emphasis>
|
||
metaventrite, ventral aspect, abdomen removed
|
||
<emphasis id="C075B1EC02F2B60E7E9AA6E62427C682" bold="true" pageId="0" pageNumber="1">D</emphasis>
|
||
detail of humeral region of right elytron, dorsal aspect.
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
<subSubSection id="7DCC053E83AEE75E9A1961E95FDC1A5A" pageId="0" pageNumber="1" type="description">
|
||
<paragraph id="A9388B849925BD575A7267870B603A3E" pageId="0" pageNumber="1">Description.</paragraph>
|
||
<paragraph id="544E4F33A71FFFE569136194B370C8C5" pageId="0" pageNumber="1">
|
||
<emphasis id="9AC43E0E80C96960242ED94091D83F48" bold="true" italics="true" pageId="0" pageNumber="1">Habitus</emphasis>
|
||
: Average sized for
|
||
<taxonomicName id="D38CBB1A0143AC7B37497B09B502C00E" authorityName="Bonelli" authorityYear="1810" lsidName="" pageId="0" pageNumber="1" rank="subFamily" subFamily="Trechinae">Trechinae</taxonomicName>
|
||
(
|
||
<emphasis id="BAF7B0B2F924E139E7AA586F62D049CF" bold="true" pageId="0" pageNumber="1">ABL</emphasis>
|
||
= 3.16-3.20 mm), pubescent, without trace of eyes. Variable in color, dark castaneous in the 1991 specimen (Suppl. material 3: fig. S1A) (possibly stained due to unusual ethanol preservation) and lighter in fresher specimens (Fig.
|
||
<figureCitation id="CA2CC63A2835FA3EF23523D926EA6781" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Horologion hubbardi female paratype, voucher CWH- 452 (VMNH), dorsal habitus. Abdomen removed for DNA extraction. Scale bar: 1 mm." figureDoi="10.3897/subtbiol.48.114404.figure2" httpUri="https://binary.pensoft.net/fig/963502" pageId="0" pageNumber="1">2</figureCitation>
|
||
, Suppl. material 3: fig. S1B); integument strongly sclerotized; proportions delicate, with pedunculate pro-mesothoracic junction; elytra vase shaped, with prominent humeral carinae ending in curved spines. Appendages relatively short; body flattened dorsoventrally.
|
||
</paragraph>
|
||
<paragraph id="36D92587C6B3D766F8D2042C1DC96FBC" pageId="0" pageNumber="1">
|
||
<emphasis id="DD9D83455E071C599CB56CEC775750E1" bold="true" italics="true" pageId="0" pageNumber="1">Head</emphasis>
|
||
: Relatively large (
|
||
<emphasis id="65A3A9D3BB621CCC6EFE2265EEDB31C2" bold="true" pageId="0" pageNumber="1">HW/PW</emphasis>
|
||
= 0.84-0.86); temples rounded; eyes entirely absent. Dorsal surface evenly covered with short, light-colored setae set in coarse circular pits. Microsculpture consisting of weakly impressed, irregular scalelike sculpticells, except a subtriangular patch on vertex where the sculpticels are coarse and conspicuous. Occipital region (concealed by pronotum) smooth, demarcated from rough vertex by a curved marginal line. Vertex with anterior supraorbital seta present on each side (Fig.
|
||
<figureCitation id="2F60297C0ED4DB4B2B3ABAC8801A6018" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Scanning electron micrographs of the forebody of Horologion hubbardi A head of voucher CWH- 452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH- 452, ventral aspect." figureDoi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1">12A</figureCitation>
|
||
); posteriorly with 3 to 5 pairs of moderately long inward-facing setae in a transverse row, none of which arise from a pore of comparable size to the anterior supraorbital setae. Frontal grooves weakly defined, shallow and short, ending at level of anterior supraorbital setae. Frontoclypeal suture weakly impressed, without carinae or horn like projection. Clypeus transverse, subrectangular, with four large fixed setae, outer pair erect and longer than inner pair; inner pair appressed, arising from smaller pores than outer pair (Suppl. material 3: fig. S3C); in addition to scattered background setae, two thin setae the same length as inner pair are present near anterior angles (Fig.
|
||
<figureCitation id="41975AC13B6025F1514A67523751D7C0" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Scanning electron micrographs of the forebody of Horologion hubbardi A head of voucher CWH- 452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH- 452, ventral aspect." figureDoi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1">12A</figureCitation>
|
||
). Labrum transverse, similar in size and shape to clypeus; anterior margin slightly crenulate, protruding slightly forward at each setiferous pore insertion; six fixed apical setae present, decreasing in length from outer to inner pairs. Ventral surface of head with a long suborbital seta on each side, set just anterior to the arcuate gular impression; tentorial pits present at anterior end of gular sutures, with small slit like openings; ventral surfaces pubescent anterior to gular impression, except strip between gular sutures, which is also strongly microsculptured with coarse, small sculpticels; microsculpture also strong within gular impression, and along margins of maxillary grooves, weak elsewhere.
|
||
</paragraph>
|
||
<paragraph id="DB296DB197BC93C1C3831E0E32A8A467" pageId="0" pageNumber="1">
|
||
<emphasis id="01BC226DF78EFB203D0D2FE0267EFC42" bold="true" italics="true" pageId="0" pageNumber="1">Antennae</emphasis>
|
||
: Length approximately half of body length (
|
||
<emphasis id="CEC38BC20741C90D24418665B41BF7A2" bold="true" pageId="0" pageNumber="1">AntL/ABL</emphasis>
|
||
= 0.51-0.53). All antennomeres pubescent, filiform; antennomeres I-X with a subapical ring of long setae, antennomere XI with a ring of long setae just beyond middle, and a crown of long setae at apex. Several small, circular pores scattered in apical half of antennomere XI, concentrated near apex. Antennomere I shorter and thicker than antennomeres II-X. Four apical antennomeres gradually increasing in width; antennomere XI largest, slightly longer than antennomere II and clearly longer than all other antennomeres; gradually tapered apically. Antennae similar in both sexes.
|
||
</paragraph>
|
||
<paragraph id="99FD0CDA00E762FAC4AA14BCA136345A" pageId="0" pageNumber="1">
|
||
<emphasis id="6CE3D01C1D2E111F8AA0667BCE66CA87" bold="true" italics="true" pageId="0" pageNumber="1">Mouthparts</emphasis>
|
||
: Mandibles with scrobal seta present; narrow and elongate, both similar in size and shape but differing in dentation: right mandible with prominent anterior retinacular tooth, terebral tooth, posterior retinacular tooth and molar tooth; left mandible without anterior retinacular tooth, with small terebral, posterior retinacular and molar teeth (Suppl. material 3: fig. S3A, B). Mentum and submentum separated by suture; submentum generally setose, with two pairs of long fixed setae, inner pair very long (the longest ventral setae of head); mentum transverse, surface glabrous except for two pairs of fixed setae, inner pair situated well behind mentum tooth; mentum shallowly biconcave, each concavity with a small irregular pit with numerous small pores; mentum tooth carinate, entire, long and acute (Fig.
|
||
<figureCitation id="8BAA80F8FC4454BB11B3DB35B63E7270" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Scanning electron micrographs of the forebody of Horologion hubbardi A head of voucher CWH- 452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH- 452, ventral aspect." figureDoi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1">12C</figureCitation>
|
||
). Labial palps glabrous except for penultimate palpomere, which has four long setae; apical palpomere long, much narrower than penultimate. Ligula carinate medially, with distinct paraglossae; anterior margin between paraglossae with six setae, an outer pair of very short setae, a submedial pair of moderately long fixed setae, and a medial pair of long fixed setae that are conjoined, arising from adjacent pores, appearing as one long seta except in SEM images; short outer pair not visible under a stereoscope at 100
|
||
<normalizedToken id="F0F0F66C2437B856D5A42208382CBCAB" originalValue="×">x</normalizedToken>
|
||
, but visible in SEM images (Suppl. material 3: fig. S4A). Maxillary palps glabrous except second palpomere, which bears two setae on outer surface near apex; second and third palpomeres somewhat globular; apical palpomere narrow and elongate, but not truly subulate (basal width subequal to apical width of penultimate palpomere) (Fig.
|
||
<figureCitation id="1AE4325A769C232298145E2DFCC53A66" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Scanning electron micrographs of the forebody of Horologion hubbardi A head of voucher CWH- 452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH- 452, ventral aspect." figureDoi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1">12A</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="D35F48811CD4657C78D54D63CBB2BE7C" pageId="0" pageNumber="1">
|
||
<emphasis id="F5435AF4599599BDE6737549074A5FC5" bold="true" italics="true" pageId="0" pageNumber="1">Prothorax</emphasis>
|
||
: Pronotum small, narrower than elytra (
|
||
<emphasis id="A17A0A06CF2742A50757E7601BEE5F4B" bold="true" pageId="0" pageNumber="1">PW/EW</emphasis>
|
||
= 0.67) and less than one fourth body length (
|
||
<emphasis id="A2598A9AC7079221ECCC703E02FEFD78" bold="true" pageId="0" pageNumber="1">PL/ABL</emphasis>
|
||
= 0.23); greatly narrowed posteriorly (
|
||
<emphasis id="15447EFB120B29B1A5D499DDFF816C41" bold="true" pageId="0" pageNumber="1">PbW/PW</emphasis>
|
||
= 0.39). Surface densely covered in light-colored setae, each set in a circular pore; setae whorled along midline: facing posteriorly in posterior half, medially in middle, and anteriorly in anterior half. Median longitudinal sulcus well impressed, but not reaching anterior or posterior margins. Lateral marginal bead lacking except for a short distance near lateral setae; otherwise, dorsal surface and hypomeron continuous. Posterior angles obsolete, without lateral setae; posterior impressions lacking. Posterior margin without bead, dorsal surface curved beneath itself, forming a smooth shelf that overhangs the mesothoracic pedicel. Prosternum (Fig.
|
||
<figureCitation id="C18CDD44F1F55DDC37A76CCBCDA40861" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Scanning electron micrographs of the forebody of Horologion hubbardi A head of voucher CWH- 452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH- 452, ventral aspect." figureDoi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1">12C</figureCitation>
|
||
) shorter than pronotum, ending anteriorly and posteriorly well before pronotal extent; setose medially; propleuron glabrous. Pleurosternal suture meeting hypomeron anteriorly behind anterior angles of prosternum, which are produced forward, overlapping hypomeron. Procoxal cavities bordered by raised margin anteriorly; closed posteriorly by propleuron narrowly joining intercoxal process (Fig.
|
||
<figureCitation id="C4632DC2D2D0607C3B5095AD11A33A82" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Scanning electron micrographs of the forebody of Horologion hubbardi A head of voucher CWH- 452, dorsal aspect, clypeal setae indicated by white arrows B prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed) C forebody of voucher CWH- 452, ventral aspect." figureDoi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1">12B</figureCitation>
|
||
). Intercoxal process elongate, acuminate posteriorly. Procoxae coarsely microsculptured, glabrous; protrochanters small, setose, with single large fixed seta near apex; femora and tibiae slender and setose, tibiae strigose on outer margin; outer margin declivitous in dorsal view, but without distinct notch (Fig.
|
||
<figureCitation id="13DC67A84D0E499699CB3BA4D454E24D" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Protarsi of Horologion A H. hubbardi male holotype, dorsal aspect B H. hubbardi female paratype, dorsal aspect C H. speokoites male holotype, dorsal aspect D H. hubbardi male holotype, ventral aspect E H. hubbardi male holotype, dorsal aspect. Scale bars: 0.10 mm." figureDoi="10.3897/subtbiol.48.114404.figure8" httpUri="https://binary.pensoft.net/fig/963508" pageId="0" pageNumber="1">8A</figureCitation>
|
||
). Inner margin of tibiae with large antenna cleaner of typical "grade B" (
|
||
<bibRefCitation id="6DD170E917906AC7C95D86BB2B0E4802" DOI="https://doi.org/10.1155/1971/927545" author="Hlavac, TF" journalOrPublisher="Psyche: A Journal of Entomology" pageId="0" pageNumber="1" pagination="51 - 66" refId="B50" refString="Hlavac, TF, 1971. Differentiation of the carabid antenna cleaner. Psyche: A Journal of Entomology 78: 51 - 66, DOI: https://doi.org/10.1155/1971/927545" title="Differentiation of the carabid antenna cleaner." url="https://doi.org/10.1155/1971/927545" volume="78" year="1971">Hlavac 1971</bibRefCitation>
|
||
), i.e. with a sinuate longitudinal band of tightly packed setae within the channel; arrangement of terminal spurs anisochaetus, i.e. situated at opposite ends of the setal band; anterior spur stouter than posterior; one large clip seta present. Tarsi densely setose and very short; protarsomeres 2-IV distinctly transverse, wider than long, each with a pair of long setae ventrally, ventral setae of protarsomere III longest and conspicuous; protarsomere IV with a thick medial ribbonlike seta that surpasses apex of tarsus; tarsal claws simple and evenly curved, without basal tooths or serrations, relatively elongate, longer than protarsomere V; males (Fig.
|
||
<figureCitation id="39DF244CF7A716FD057C911EA8396EE7" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Protarsi of Horologion A H. hubbardi male holotype, dorsal aspect B H. hubbardi female paratype, dorsal aspect C H. speokoites male holotype, dorsal aspect D H. hubbardi male holotype, ventral aspect E H. hubbardi male holotype, dorsal aspect. Scale bars: 0.10 mm." figureDoi="10.3897/subtbiol.48.114404.figure8" httpUri="https://binary.pensoft.net/fig/963508" pageId="0" pageNumber="1">8A, C-D</figureCitation>
|
||
) with protarsomeres I and II asymmetrically expanded and dentate on inner margin, with a single row of at least seven adhesive setae on venter of inner dentate expansions; females with protarsomeres I and II symmetrical, not dentate and without ventral adhesive vestiture, protarsomere I slightly longer than wide, protarsomere II transverse, subequal to protarsomere III.
|
||
</paragraph>
|
||
<paragraph id="CC0E4A1AAD65D35904FE5B04E5E5E20F" pageId="0" pageNumber="1">
|
||
<emphasis id="E3D860FD92C8D970C45F9E967821FC4B" bold="true" italics="true" pageId="0" pageNumber="1">Pterothorax</emphasis>
|
||
: Elytra moderately long, length slightly more than half of
|
||
<emphasis id="6A9E4B20F42AD4E2BEB69D3C6CB26532" bold="true" pageId="0" pageNumber="1">ABL</emphasis>
|
||
; scutellum very narrow and elongate (Fig.
|
||
<figureCitation id="9A7A17DE90CD0DD23F6BEF238F702747" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">10D</figureCitation>
|
||
). Elytra fused along suture for most of their length, narrowly separated in apical fourth. Dorsal surface evenly setose, with short light-colored setae each set in a deep circular pit; microsculpture weakly impressed, sculpticels irregularly shaped, scale like and longitudinally stretched. Humeri each bearing an angulate shelf, flanked proximally by a strong carina and ending in a prominent curved spine; lateral bead of elytra moderately crenulate beyond humeral spine for a short distance and smooth beyond that (Fig.
|
||
<figureCitation id="AE237ADFD17729BA3E7BF0FFBD245B52" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">10D</figureCitation>
|
||
). Each elytron with a parascutellar seta, four subhumeral lateral setae, two submedial lateral setae, three apical setae (two lateral and one discal, in the position of the "subapical seta" of
|
||
<bibRefCitation id="1C16640C8575AE673625EB1E448250F1" DOI="https://doi.org/10.3897/dez.68.66181" author="Schmidt, J" journalOrPublisher="Deutsche Entomologische Zeitschrift" pageId="0" pageNumber="1" pagination="207 - 224" refId="B92" refString="Schmidt, J, Scholz, S, Maddison, DR, 2021. Balticeler kerneggeri gen. nov., sp. nov., an enigmatic Baltic amber fossil of the ground beetle subfamily Trechinae (Coleoptera, Carabidae). Deutsche Entomologische Zeitschrift 68: 207 - 224, DOI: https://doi.org/10.3897/dez.68.66181" title="Balticeler kerneggeri gen. nov., sp. nov., an enigmatic Baltic amber fossil of the ground beetle subfamily Trechinae (Coleoptera, Carabidae)." url="https://doi.org/10.3897/dez.68.66181" volume="68" year="2021">Schmidt et al. (2021)</bibRefCitation>
|
||
, umbilicate pore
|
||
<normalizedToken id="F2B846919FCEF15D4198262F10151BA8" originalValue="“8”">"8"</normalizedToken>
|
||
of anilline taxonomists (
|
||
<bibRefCitation id="EAA4DA20FCD272A5EF5B1080A5814EFA" author="Giachino, PM" journalOrPublisher="Biodiversity Journal Monograph" pageId="0" pageNumber="1" pagination="1 - 112" refId="B43" refString="Giachino, PM, Vailati, D, 2011. Review of the Anillina of Greece (Coleoptera, Carabidae, Bembidiini). Biodiversity Journal Monograph 1: 1 - 112" title="Review of the Anillina of Greece (Coleoptera, Carabidae, Bembidiini)." volume="1" year="2011">Giachino and Vailati 2011</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="9BF62AB6089F5F85CA8BADD1AE740097" DOI="https://doi.org/10.3897/zookeys.352.6052" author="Sokolov, IM" journalOrPublisher="ZooKeys" pageId="0" pageNumber="1" pagination="51 - 92" refId="B97" refString="Sokolov, IM, 2013. A new genus and eight new species of the subtribe Anillina (Carabidae, Trechinae, Bembidiini) from Mexico, with a cladistic analysis and some notes on the evolution of the genus. ZooKeys 352: 51 - 92, DOI: https://doi.org/10.3897/zookeys.352.6052" title="A new genus and eight new species of the subtribe Anillina (Carabidae, Trechinae, Bembidiini) from Mexico, with a cladistic analysis and some notes on the evolution of the genus." url="https://doi.org/10.3897/zookeys.352.6052" volume="352" year="2013">Sokolov 2013</bibRefCitation>
|
||
)), and one discal seta in third interval at about the level of fifth lateral seta (Fig.
|
||
<figureCitation id="757B845C0B9AB95A1C9F9F27A763A8D6" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Elytral characters of Horologion species A H. speokoites holotype, elytra, oblique left lateral aspect B H. hubbardi paratype (CMNH), ditto C H. hubbardi paratype (VMNH), SEM detail of apex of left elytron, ventral aspect D H. hubbardi paratype (VMNH), SEM photo of right elytron, dorsal aspect E H. hubbardi paratype (VMNH), SEM detail of right elytron (area indicated by black box in D). Black arrows: discal setae, white arrows: plica." figureDoi="10.3897/subtbiol.48.114404.figure11" httpUri="https://binary.pensoft.net/fig/963511" pageId="0" pageNumber="1">11B, D</figureCitation>
|
||
); discal seta indistinguishable from background pubescence in low-magnification dorsal view but distinctly visible in oblique or lateral views (Fig.
|
||
<figureCitation id="C6DB9B7D74226DBD4244A1005FB3C6B1" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Elytral characters of Horologion species A H. speokoites holotype, elytra, oblique left lateral aspect B H. hubbardi paratype (CMNH), ditto C H. hubbardi paratype (VMNH), SEM detail of apex of left elytron, ventral aspect D H. hubbardi paratype (VMNH), SEM photo of right elytron, dorsal aspect E H. hubbardi paratype (VMNH), SEM detail of right elytron (area indicated by black box in D). Black arrows: discal setae, white arrows: plica." figureDoi="10.3897/subtbiol.48.114404.figure11" httpUri="https://binary.pensoft.net/fig/963511" pageId="0" pageNumber="1">11B</figureCitation>
|
||
) or at higher magnification (Fig.
|
||
<figureCitation id="92957C42807A67556DED663FBE3A4AF1" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Elytral characters of Horologion species A H. speokoites holotype, elytra, oblique left lateral aspect B H. hubbardi paratype (CMNH), ditto C H. hubbardi paratype (VMNH), SEM detail of apex of left elytron, ventral aspect D H. hubbardi paratype (VMNH), SEM photo of right elytron, dorsal aspect E H. hubbardi paratype (VMNH), SEM detail of right elytron (area indicated by black box in D). Black arrows: discal setae, white arrows: plica." figureDoi="10.3897/subtbiol.48.114404.figure11" httpUri="https://binary.pensoft.net/fig/963511" pageId="0" pageNumber="1">11D, E</figureCitation>
|
||
); second subhumeral, second submedial, and posterior-most apical seta greatly elongate and filamentous. Ventral surface of elytra each with a well-developed lateral plica near apex, its surface strongly microsculptured with scale-like sculpticells (Fig.
|
||
<figureCitation id="40E014C57EAE934A9F1E1A198843A057" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Elytral characters of Horologion species A H. speokoites holotype, elytra, oblique left lateral aspect B H. hubbardi paratype (CMNH), ditto C H. hubbardi paratype (VMNH), SEM detail of apex of left elytron, ventral aspect D H. hubbardi paratype (VMNH), SEM photo of right elytron, dorsal aspect E H. hubbardi paratype (VMNH), SEM detail of right elytron (area indicated by black box in D). Black arrows: discal setae, white arrows: plica." figureDoi="10.3897/subtbiol.48.114404.figure11" httpUri="https://binary.pensoft.net/fig/963511" pageId="0" pageNumber="1">11E</figureCitation>
|
||
). Mesoventrite (Fig.
|
||
<figureCitation id="D389542AD1B81782A3F71A3170198DE9" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">10B</figureCitation>
|
||
) with coarse isodiametric microsculpture; narrow, much longer than metaventrite; extended anteriorly as a parallel-sided pedicel that extends beyond posterior extent of pronotum; surface of pedicel strongly rugose, with numerous transverse furrows; posterior half of mesoventrite with a medial longitudinal depression flanked by low, parallel carinae that coalesce anteriorly, each carina bearing a pair of long setae; medial depression extending posteriorly onto intercoxal process. Mesanepisternum and mesoventrite apparently fused, without discernible suture; mesocoxae conjunct, i.e., entirely enclosed by mesoventrite and metaventrite, mesepimeron not meeting mesocoxae (Fig.
|
||
<figureCitation id="67C8F47EE5A2341E800E425B20316A6E" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">10A</figureCitation>
|
||
). Mesocoxae with coarse scale-like microsculpture and sparse setae, each with a well-developed knob on inner margin; mesotrochanters densely setose but without apparent macrosetae; mesofemora and mesotibiae slender, setose except for glabrous area on posterior face of femora; inner face of mesotibiae strigose; apical half of mesotibiae with dense brush of coarse setae; apex of mesotibiae with adjacent pair of short spurs on posterior margin, barely extending past length of first mesotarsomere; mesotarsi of both sexes similar in form to female protarsi. Metaventrite (Fig.
|
||
<figureCitation id="B15E266DE4CD6DE0160918C7E3FF2F50" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">10C</figureCitation>
|
||
) short, setose and coarsely microsculptured, with shallow medial depression; intercoxal process cleft posteriorly; metanepisternum and metaventrite separated by suture; metepimeron visible (Fig.
|
||
<figureCitation id="806D5F39709FEA587B908683C402B889" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Scanning electron microscope images of Horologion hubbardi female paratype, voucher CWH- 452 (VMNH) A detail of right side of pterothorax, ventral aspect B mesoventrite, ventral aspect C metaventrite, ventral aspect, abdomen removed D detail of humeral region of right elytron, dorsal aspect." figureDoi="10.3897/subtbiol.48.114404.figure10" httpUri="https://binary.pensoft.net/fig/963510" pageId="0" pageNumber="1">10A</figureCitation>
|
||
), overlapping first abdominal ventrite. Metacoxae setose, without apparent macrosetae; well separated, distance between them approximately equal to width of one mesocoxa; metatrochanters small, approximately equal in length to metacoxae, gradually narrowed apically, not strongly pointed; metafemora and metatibiae similar to those of mesothoracic legs, except metatibiae lack dense brush of setae; metatarsi more elongate than tarsi of other legs, metatarsomeres slightly longer than wide, gradually increasing in length from I to IV, V slightly longer than combined length of III and IV; medial ribbonlike setae on apex of metatarsomere IV narrower than on pro- and mesotarsi.
|
||
</paragraph>
|
||
<caption id="60553A9247E76833146C9B1BCC21647F" doi="10.3897/subtbiol.48.114404.figure11" httpUri="https://binary.pensoft.net/fig/963511" pageId="0" pageNumber="1" start="Figure 11" startId="F11">
|
||
<paragraph id="51E27A7BD8375A15F17D8D9366ACE64B" pageId="0" pageNumber="1">
|
||
<emphasis id="A9BBF3D6349D69F97C0FA95706F5935D" bold="true" pageId="0" pageNumber="1">Figure 11.</emphasis>
|
||
Elytral characters of
|
||
<taxonomicName id="6C0AE7DADE2B1113DA1DD992329D406D" authorityName="Valentine" authorityYear="1932" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="E467D7EFD678F46FA9E0624F685F6BAD" italics="true" pageId="0" pageNumber="1">Horologion</emphasis>
|
||
</taxonomicName>
|
||
species
|
||
<emphasis id="78BA37575F360A0819BF2D167CB465F8" bold="true" pageId="0" pageNumber="1">A</emphasis>
|
||
<taxonomicName id="BA7409046529370383D452BAF6782A7F" lsidName="H. speokoites" pageId="0" pageNumber="1" rank="species" species="speokoites">
|
||
<emphasis id="9DEFB4033A35C5045E7DABAD9A406A83" italics="true" pageId="0" pageNumber="1">H. speokoites</emphasis>
|
||
</taxonomicName>
|
||
holotype, elytra, oblique left lateral aspect
|
||
<emphasis id="234107740058056C302D06F99C75BEDB" bold="true" pageId="0" pageNumber="1">B</emphasis>
|
||
<taxonomicName id="3077740B68F26E97E5A054663B65BD00" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="C687D7C5BA199E5C37D5C658D3CDCC79" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
paratype (CMNH), ditto
|
||
<emphasis id="079D60CCCF0A5BF2EDA943F850184F25" bold="true" pageId="0" pageNumber="1">C</emphasis>
|
||
<taxonomicName id="5F3A7DF2C95F668C5852065507CFB120" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="63C8B6D401C39EE133C6EEF56367350A" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
paratype (VMNH), SEM detail of apex of left elytron, ventral aspect
|
||
<emphasis id="DFC8C35DAA2F3CDB09E8902161B19011" bold="true" pageId="0" pageNumber="1">D</emphasis>
|
||
<taxonomicName id="11B1C076AC0F74498BC9A05AF793E55C" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="D5252688F199B74D13F416F67B5BD77F" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
paratype (VMNH), SEM photo of right elytron, dorsal aspect
|
||
<emphasis id="37580374524EC56A08E0D3851B135B9E" bold="true" pageId="0" pageNumber="1">E</emphasis>
|
||
<taxonomicName id="E0FABADB69C5B450A25188C074B29F52" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="5409149F9F539A563FE2EF4A5AEE787A" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
paratype (VMNH), SEM detail of right elytron (area indicated by black box in
|
||
<emphasis id="C55EEB250B5D3098FFB68D6D58C9D18C" bold="true" pageId="0" pageNumber="1">D</emphasis>
|
||
). Black arrows: discal setae, white arrows: plica.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="C6070AB959D4AE9F27379A1F77BA3B6B" pageId="0" pageNumber="1">
|
||
<emphasis id="ECBC40112D3B9CED52D50D5C85A85532" bold="true" italics="true" pageId="0" pageNumber="1">Male genitalia</emphasis>
|
||
: Relatively small (Length of ring sclerite /
|
||
<emphasis id="6371F8DC1FD1390CFD8C03E4190F8B42" bold="true" pageId="0" pageNumber="1">ABL</emphasis>
|
||
= 0.16); ring sclerite similar to that of
|
||
<taxonomicName id="32480259EED741B925EB1DBE82E05CD7" lsidName="H. speokoites" pageId="0" pageNumber="1" rank="species" species="speokoites">
|
||
<emphasis id="F374BAA35F07051356584E60E369B4D2" italics="true" pageId="0" pageNumber="1">H. speokoites</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="2D81DB95E0836D501F0EE17514C1850D" DOI="https://doi.org/10.1093/aesa/25.1.1" author="Valentine, JM" journalOrPublisher="Annals of the Entomological Society of America" pageId="0" pageNumber="1" pagination="1 - 11" refId="B114" refString="Valentine, JM, 1932. Horologion, a New Genus of Cave Beetles. Annals of the Entomological Society of America 25: 1 - 11, DOI: https://doi.org/10.1093/aesa/25.1.1" title="Horologion, a New Genus of Cave Beetles." url="https://doi.org/10.1093/aesa/25.1.1" volume="25" year="1932">Valentine 1932</bibRefCitation>
|
||
, fig. 13): yoke shaped, posterior margin produced as an obtuse angulation, narrowed anteriorly where sides join to form a short, flattened extension that is curved ventrally at its slightly asymmetrical apex. Median lobe of aedeagus (Fig.
|
||
<figureCitation id="12CA40425CCE43A2FA3E84E2995903DE" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9D</figureCitation>
|
||
) broad and lightly sclerotized, with entire dorsal margin and most of ventral face membranous; basal lobes and sides of ventral margin sclerotized and giving the organ a trough-like shape; ventral margin in lateral aspect slightly curved in proximal half, with membranous portion sagging below; apex small, extended a short distance past membranous dorsal margin and appearing evenly rounded and symmetrical in dorsal aspect; median lobe not twisted from plane of basal lobes (Fig.
|
||
<figureCitation id="DC4ACA8013256084B3FE98A14B6FE1FE" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9F</figureCitation>
|
||
). Left paramere (Fig.
|
||
<figureCitation id="D9A0DD27CDA9E7AFB7C4855B9F25D112" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9E</figureCitation>
|
||
) relatively large and subtriangular, with numerous pores on dorsoapical margin and three apical setae. Right paramere (Fig.
|
||
<figureCitation id="8D6F0663BB50743D28F9DFE22B5163BE" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9G</figureCitation>
|
||
) slightly smaller than left paramere, and more styliform, with numerous pores on dorsoapical margin and three apical setae. Internal sac of median lobe with well sclerotized flagellum surrounding a small spine and a ventral field of small sclerotized scales; flagellum rotated dorsally, so that in lateral aspect it appears as a complex folded structure (Fig.
|
||
<figureCitation id="742243A2D66EA04A127260EC0A33C2B3" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9D</figureCitation>
|
||
); in dorsal aspect strongly curved (Fig.
|
||
<figureCitation id="5652BA3AD5BDD36031019E1ED7403D0E" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9F</figureCitation>
|
||
), swollen and spiraled proximally at junction with sperm duct, abruptly narrowed beyond this region, very gradually tapering toward apex.
|
||
</paragraph>
|
||
<caption id="C684899EA42CD068271F1838762915A5" doi="10.3897/subtbiol.48.114404.figure12" httpUri="https://binary.pensoft.net/fig/963512" pageId="0" pageNumber="1" start="Figure 12" startId="F12">
|
||
<paragraph id="244DA12EB6321C916CC86C4DAB7F9D97" pageId="0" pageNumber="1">
|
||
<emphasis id="44DC8CC58ADABE239D41502E53731DB0" bold="true" pageId="0" pageNumber="1">Figure 12.</emphasis>
|
||
Scanning electron micrographs of the forebody of
|
||
<taxonomicName id="E75FB57F19926C5C1BD350EED9BEB82F" authorityName="Harden & Davidson" authorityYear="2024" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion hubbardi" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="hubbardi">
|
||
<emphasis id="252B279CD8DF12D2C21AAE650FC4E2E3" italics="true" pageId="0" pageNumber="1">Horologion hubbardi</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="493726AFB870DEE1C6F75DCDF6C2B793" bold="true" pageId="0" pageNumber="1">A</emphasis>
|
||
head of voucher CWH-452, dorsal aspect, clypeal setae indicated by white arrows
|
||
<emphasis id="CB310D55B3B505C5EB8BCE6ABFAA0C4F" bold="true" pageId="0" pageNumber="1">B</emphasis>
|
||
prosternum of non-type specimen, posterior aspect (head, pronotum and most of leg segments removed)
|
||
<emphasis id="505CA636AB7C0C8D0703C6F98726BD00" bold="true" pageId="0" pageNumber="1">C</emphasis>
|
||
forebody of voucher CWH-452, ventral aspect.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC9A6F64431DF02951618E09B5DD55E1" pageId="0" pageNumber="1">
|
||
<emphasis id="84BF3BB336FAAA274BE6C44AEE423AD1" bold="true" italics="true" pageId="0" pageNumber="1">Female genitalia</emphasis>
|
||
: (Fig.
|
||
<figureCitation id="B4D36BEB4856E5B59200680AFB74FAFB" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Genitalia of Horologion species A H. speokoites median lobe, left lateral aspect B H. speokoites left paramere, left lateral aspect C H. speokoites median lobe, dorsal aspect D H. hubbardi median lobe, left lateral aspect E H. hubbardi left paramere, left lateral aspect F H. hubbardi median lobe, dorsal aspect G H. hubbardi right paramere, right lateral aspect H H. hubbardi female genitalia, ventral aspect A-C after Valentine (1932). Scale bar: 0.1 mm." figureDoi="10.3897/subtbiol.48.114404.figure9" httpUri="https://binary.pensoft.net/fig/963509" pageId="0" pageNumber="1">9H</figureCitation>
|
||
) Gonocoxite 2 narrow, moderately long and weakly curved, bearing numerous pores and a single short preapical seta on inner margin. Tergite X well-sclerotized and forming a subtriangular bridge behind gonocoxites. Spermatheca and spermathecal gland present; spermathecal duct relatively wide and moderately long, with an abrupt U-shaped bend proximally; spermatheca small and pouchlike; spermathecal gland long and narrow, arising from base of spermatheca near junction with duct.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="076B07AA3CF29635D30E3CD02C508601" pageId="0" pageNumber="1" type="distribution">
|
||
<paragraph id="2077D279EAFA11BC546FD066A8FA5474" pageId="0" pageNumber="1">Distribution.</paragraph>
|
||
<paragraph id="B2477D8BAE697895DF1607515FBCF56D" pageId="0" pageNumber="1">Known only from Williams Cave, in Bath County, Virginia. In the database of the Virginia Speleological Survey (VSS, https://www.virginiacaves.org/), this cave is number 2779.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C2DC716B6AAE964F18FCDE440142D64E" pageId="0" pageNumber="1" type="sympatry">
|
||
<paragraph id="5EBC147B08AF578211D34FA1EF1DBFB3" pageId="0" pageNumber="1">Sympatry.</paragraph>
|
||
<paragraph id="19036ABDFD31921F1EFA9F3C3BC32E02" pageId="0" pageNumber="1">
|
||
Williams Cave is also home to the eyeless trechine
|
||
<taxonomicName id="D5F0409B4C1A1075B1FC5799FA547DF8" authorityName="Barr" authorityYear="1965" class="Insecta" family="Carabidae" genus="Pseudanophthalmus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudanophthalmus intersectus" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="intersectus">
|
||
<emphasis id="E343D437BAF5963A77598D6E428C301C" italics="true" pageId="0" pageNumber="1">Pseudanophthalmus intersectus</emphasis>
|
||
</taxonomicName>
|
||
Barr, which also occurs in two other nearby caves in Bath County (Virginia DCR-DNH data). An individual of this species was found in the same microhabitat as the female paratype of
|
||
<taxonomicName id="DE9BC5316DCE1F6B9E3815E27A9CD95B" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="D0F9FB39F45F334D5DEDA13E0AC89E49" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
, and members of the two species presumably occur in syntopy. The only other carabid beetle known from Williams Cave is the surface tachyine
|
||
<taxonomicName id="C678AD91B174A501AC3DC35BDC2BF23C" authorityName="LeConte" authorityYear="1848" class="Insecta" family="Carabidae" genus="Paratachys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paratachys scitulus" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="scitulus">
|
||
<emphasis id="C2526E3E25246D55AEE890A3E07A05EC" italics="true" pageId="0" pageNumber="1">Paratachys scitulus</emphasis>
|
||
</taxonomicName>
|
||
(LeConte), a common and widespread species in eastern North America; one specimen was found in organic debris just inside the entrance in August 2022.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="1E34C60C8CF4C1804FB7F7845D6EE272" pageId="0" pageNumber="1" type="natural history">
|
||
<paragraph id="FD3940A642F8D407530D8C1AB4252BD4" pageId="0" pageNumber="1">Natural history.</paragraph>
|
||
<paragraph id="DDD077D80A4E5F3F0A79CA3B44BF6CDC" pageId="0" pageNumber="1">
|
||
Williams Cave is a large cave, with a surveyed length of 5.39 km (VSS data). The cave is shallow in relation to the overlying surface topology and is damp in places, with numerous ceiling drips and small pools. Most water in the cave is recharged through these ceiling drips. All specimens of
|
||
<taxonomicName id="2B41AA1B1B4D28CFA8FEF1069EDCD98A" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="5A03A66858F8681B497DFC83A82526F6" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
were found in or near small pools of water. Specimens were collected in March, August and September. Specimens from 2022 and 2023 were collected in somewhat distant sections of the cave, but
|
||
<emphasis id="A931B2080DAFB92F9F1D2FC419D60B9A" italics="true" pageId="0" pageNumber="1">COI</emphasis>
|
||
sequences of the two are identical, suggesting they are not isolated. Immature life stages are unknown.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="D89F97305B099140F97D6DC9181C5CEE" pageId="0" pageNumber="1" type="species status justification">
|
||
<paragraph id="219BB353A49AD7F7433EA4F027E63286" pageId="0" pageNumber="1">Species status justification.</paragraph>
|
||
<paragraph id="85B9E8EAA582CCDD73D1E2D425783A1A" pageId="0" pageNumber="1">
|
||
The differences in male protarsi (first and second protarsomeres asymmetrically expanded and dentate in
|
||
<taxonomicName id="0FB1FAF58EF033F6C1ED2F1FA65CF92F" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="A5E9299814DDFAC25AFD5AF5FC6E08D6" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
, only first protarsomere weakly dentate in
|
||
<taxonomicName id="0EE124957F4A4864C9E9E748654BD17C" lsidName="H. speokoites" pageId="0" pageNumber="1" rank="species" species="speokoites">
|
||
<emphasis id="A684561DFC2792C2715D2480D02D2311" italics="true" pageId="0" pageNumber="1">H. speokoites</emphasis>
|
||
</taxonomicName>
|
||
), the form of the elytra (flattened, with prominent curved spines on the humeri in
|
||
<taxonomicName id="4D5F8A537873947A3DCAFEACC4D1C86E" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="E5576B583A0701D4326AD52EA70740B1" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
, convex and with a small humeral carinal shelf without spines in
|
||
<taxonomicName id="67E1E429FA6B59C7658A7E145FBFD5D1" lsidName="H. speokoites" pageId="0" pageNumber="1" rank="species" species="speokoites">
|
||
<emphasis id="DF36912205213D3FCEE35D26DF14F8A3" italics="true" pageId="0" pageNumber="1">H. speokoites</emphasis>
|
||
</taxonomicName>
|
||
), and the male genitalia, particularly the parameres (tapered and with 3 apical setae in
|
||
<taxonomicName id="96679B5C99671FAE74AA9DE3137F096F" lsidName="H. hubbardi" pageId="0" pageNumber="1" rank="species" species="hubbardi">
|
||
<emphasis id="1AF7A1C03C9145D7D5BD5DE0FE006DEA" italics="true" pageId="0" pageNumber="1">H. hubbardi</emphasis>
|
||
</taxonomicName>
|
||
, broad and blunt with 6 apical setae in
|
||
<taxonomicName id="2B000F1D8F66DA87F30E7A53D8C9BAD8" lsidName="H. speokoites" pageId="0" pageNumber="1" rank="species" species="speokoites">
|
||
<emphasis id="D9F51D43E1417BE32AC0B9252D82B824" italics="true" pageId="0" pageNumber="1">H. speokoites</emphasis>
|
||
</taxonomicName>
|
||
) are great enough to warrant recognition of the two as distinct taxa that are reproductively isolated. The two species are also geographically isolated, occurring 70 air km distant and on the opposite side of several large ridges of noncarbonate rock with numerous peaks above 1200 m, indicating complete isolation of these blind subterranean beetles (Fig.
|
||
<figureCitation id="318FD1DEA037300CF62C2507D8CA5916" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Distribution map of species of Horologion. Black triangle: Horologion hubbardi, black circle: Horologion speokoites. Grey shaded areas represent exposed karst (Weary and Doctor 2014)." figureDoi="10.3897/subtbiol.48.114404.figure13" httpUri="https://binary.pensoft.net/fig/963513" pageId="0" pageNumber="1">13</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="5E832136A61E90D789CC647F3A00CBD4" doi="10.3897/subtbiol.48.114404.figure13" httpUri="https://binary.pensoft.net/fig/963513" pageId="0" pageNumber="1" start="Figure 13" startId="F13">
|
||
<paragraph id="99862E1B7EC3EFCB3EE66CABDD556631" pageId="0" pageNumber="1">
|
||
<emphasis id="4A141AB04D78906D84CDEEE6FDE6B60E" bold="true" pageId="0" pageNumber="1">Figure 13.</emphasis>
|
||
Distribution map of species of
|
||
<taxonomicName id="8862E523E5C19743A78889805E89B5F6" authorityName="Valentine" authorityYear="1932" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="0DAAB944EDEAF1440D5D44D9EAF53F37" italics="true" pageId="0" pageNumber="1">Horologion</emphasis>
|
||
</taxonomicName>
|
||
. Black triangle:
|
||
<taxonomicName id="9C368AFC329317B15466C72D2E7DD538" authorityName="Harden & Davidson" authorityYear="2024" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion hubbardi" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="hubbardi">
|
||
<emphasis id="01FD83FCFD65E47502D3133582887A96" italics="true" pageId="0" pageNumber="1">Horologion hubbardi</emphasis>
|
||
</taxonomicName>
|
||
, black circle:
|
||
<taxonomicName id="A1B083C3877F0C9598BE08CA4DAD9E8F" authorityName="Valentine" authorityYear="1932" class="Insecta" family="Carabidae" genus="Horologion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Horologion speokoites" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="speokoites">
|
||
<emphasis id="E5FAA79AD99E509356AC61AFE5330ADB" italics="true" pageId="0" pageNumber="1">Horologion speokoites</emphasis>
|
||
</taxonomicName>
|
||
. Grey shaded areas represent exposed karst (
|
||
<bibRefCitation id="223CFE4DD466E4B50F45D52E14E5F665" DOI="https://doi.org/10.3133/ofr20141156" author="Weary, DJ" journalOrPublisher="Systematic Entomology" pageId="0" pageNumber="1" refId="B118" refString="Weary, DJ, Doctor, DH, 2014. Karst in the United States: A digital map compilation and database. U.S. Geological Survey Open-File Report, 23 pp. https://doi.org/10.3133/ofr20141156" title="Karst in the United States: A digital map compilation and database. U. S. Geological Survey Open-File Report, 23 pp." url="https://doi.org/10.3133/ofr20141156" year="2014">Weary and Doctor 2014</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
<subSubSection id="369EE1DC8562CF3C64D5BCA9F413E101" pageId="0" pageNumber="1" type="derivation of name">
|
||
<paragraph id="29841CC4911C9F8AB9A00CA12C1599AF" pageId="0" pageNumber="1">Derivation of name.</paragraph>
|
||
<paragraph id="1CAB534A0EE9E18E8C8B68B890AB140B" pageId="0" pageNumber="1">
|
||
This species is named in honor of its discoverer, David A. Hubbard, Jr., in recognition of his important contributions to cave biodiversity and conservation. In addition to many significant collections of cave carabids in Virginia, notable discoveries by Hubbard include the single known specimen of the Chinese stygobiontic dytiscid genus
|
||
<taxonomicName id="6E1A9676A3A636E24CC118EE90F0DB2C" authorityName="Spangler" authorityYear="1996" class="Insecta" family="Dytiscidae" genus="Sinodytes" kingdom="Animalia" lsidName="Sinodytes" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="2F0E27737A4EEB8677AC78FBF59997B7" italics="true" pageId="0" pageNumber="1">Sinodytes</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="E6A2F765225FF8D0DA8807E2F54E3C4F" author="Spangler, PJ" journalOrPublisher="Insecta Mundi" pageId="0" pageNumber="1" pagination="241 - 259" refId="B102" refString="Spangler, PJ, 1996. Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae). Insecta Mundi 10: 241 - 259" title="Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae)." volume="10" year="1996">Spangler 1996</bibRefCitation>
|
||
) and a highly modified species of the pselaphine rove beetle genus
|
||
<taxonomicName id="654DABFAA8C071A133DB08E13AAB0812" class="Insecta" family="Staphylinidae" genus="Mipseltyrus" kingdom="Animalia" lsidName="Mipseltyrus" order="Coleoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="BA14861832425E1D92767D914F7CD89F" italics="true" pageId="0" pageNumber="1">Mipseltyrus</emphasis>
|
||
</taxonomicName>
|
||
that remains undescribed (C. Harden, personal observation).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="05B97C1230C9F69F1870A6C8544CD99D" pageId="0" pageNumber="1" type="suggested vernacular name">
|
||
<paragraph id="588A7314EE9E74A16E1832523F20E37E" pageId="0" pageNumber="1">Suggested vernacular name.</paragraph>
|
||
<paragraph id="3BF3E16EAB8140550B61E3E93346EFA1" pageId="0" pageNumber="1">
|
||
<normalizedToken id="4A9C82C63C64EB7CCC77E4A8D723F346" originalValue="“Hubbard’s">"Hubbard's</normalizedToken>
|
||
Hourglass Beetle".
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</subSection>
|
||
</document> |