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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.963.53470" ID-GBIF-Dataset="0831704f-4ecf-4989-8072-defde35921ec" ID-PMC="PMC7458944" ID-Pensoft-Pub="1313-2970-963-45" ID-Pensoft-UUID="FE8C7A0A85B45DF7848BFAB844F93C28" ID-PubMed="32922131" ID-ZooBank="9636C9F2C6BD4B34BCC6ED214C7B0D19" ModsDocID="1313-2970-963-45" checkinTime="1598316761636" checkinUser="pensoft" docAuthor="Englund, William F., Njoroge, Laban, Bistroem, Olof, Miller, Kelly B., Bilton, David T. &amp; Bergsten, Johannes" docDate="2020" docId="34A110FC1AAA506EAD44515DBF8E5B46" docLanguage="en" docName="ZooKeys 963: 45-79" docOrigin="ZooKeys 963" docSource="http://dx.doi.org/10.3897/zookeys.963.53470" docTitle="Agabus riberae Bilton, Englund &amp; Bergsten 2020, sp. nov." docType="treatment" docUuid="A391EC84-95B1-480F-B4DD-E7DEA2AE3076" docUuidSource="ZooBank" docVersion="4" id="FE8C7A0A85B45DF7848BFAB844F93C28" lastPageNumber="45" masterDocId="FE8C7A0A85B45DF7848BFAB844F93C28" masterDocTitle="Taxonomic revision of the Afrotropical Agabus raffrayi species group with the description of four new species (Coleoptera, Dytiscidae)" masterLastPageNumber="79" masterPageNumber="45" pageNumber="45" updateTime="1668169378299" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Taxonomic revision of the Afrotropical Agabus raffrayi species group with the description of four new species (Coleoptera, Dytiscidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Englund, William F.</mods:namePart>
<mods:affiliation>Swedish Museum of Natural History, Department of Zoology, Box 50007, SE- 10405 Stockholm, Sweden</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Njoroge, Laban</mods:namePart>
<mods:affiliation>National Museums of Kenya, Section of Invertebrate Zoology, Museum Hill, P. O. BOX 40658 - 00100, Nairobi, Kenya</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bistroem, Olof</mods:namePart>
<mods:affiliation>Finnish Museum of Natural History, Zoology Unit, P. O. Box 17, FI- 00014 University of Helsinki, Finland</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Miller, Kelly B.</mods:namePart>
<mods:affiliation>Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131 - 0001, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bilton, David T.</mods:namePart>
<mods:affiliation>Marine Biology and Ecology Research Centre, School of Biological and Marine Sciences, University of Plymouth, Drake Circus, Plymouth PL 4 8 AA, UK &amp; Department of Zoology, University of Johannesburg, PO Box 524, Auckland Park, Johannesburg 2006, South Africa</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bergsten, Johannes</mods:namePart>
<mods:affiliation>Swedish Museum of Natural History, Department of Zoology, Box 50007, SE- 10405 Stockholm, Sweden</mods:affiliation>
<mods:nameIdentifier type="email">johannes.bergsten@nrm.se</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="volume">
<mods:number>963</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>45</mods:start>
<mods:end>79</mods:end>
</mods:extent>
</mods:part>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.963.53470</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.963.53470</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-963-45</mods:identifier>
<mods:identifier type="ZooBank">9636C9F2C6BD4B34BCC6ED214C7B0D19</mods:identifier>
<mods:identifier type="Pensoft-UUID">FE8C7A0A85B45DF7848BFAB844F93C28</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="167293856" LSID="urn:lsid:zoobank.org:act:A391EC84-95B1-480F-B4DD-E7DEA2AE3076" httpUri="http://treatment.plazi.org/id/34A110FC1AAA506EAD44515DBF8E5B46" lastPageNumber="45" pageId="0" pageNumber="45">
<subSubSection pageId="0" pageNumber="45" type="nomenclature">
<paragraph pageId="0" pageNumber="45">
<taxonomicName LSID="34A110FC-1AAA-506E-AD44-515DBF8E5B46" authority="Bilton, Englund &amp; Bergsten" authorityName="Bilton, Englund &amp; Bergsten" authorityYear="2020" class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus riberae" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="species" species="riberae" status="sp. nov.">Agabus riberae Bilton, Englund &amp; Bergsten</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="45">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Distribution map of the Agabus raffrayi group. Relief is expressed in meters above sea level." figureDoi="10.3897/zookeys.963.53470.figure1" httpUri="https://binary.pensoft.net/fig/444398" pageId="0" pageNumber="45">Figures 1</figureCitation>
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. South African Agabus spp., aedeagal apices, lateral view A A. austellus sp. nov. holotype, Keurboom, Western Cape B A. austellus sp. nov., Gydo Pass, Western Cape C A. austellus sp. nov. Groote Swartberg, Western Cape D A. austellus sp. nov. Sentinel Peak, KZN Drakensberg E A. riberae sp. nov. holotype, Kamiesberg, Northern Cape; F A. riberae sp. nov. paratype, Kamiesberg, Northern Cape G A. agulhas sp. nov. holotype, Rattelrivier, Western Cape." figureDoi="10.3897/zookeys.963.53470.figure6" httpUri="https://binary.pensoft.net/fig/444403" pageId="0" pageNumber="45">, 6E, F</figureCitation>
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Agabus spp., aedeagus in lateral view A A. sjostedti B A. dytiscoides C A. ruwenzoricus D A. raffrayi E A. pallidus F A. anguluverpus sp. nov. G A. austellus sp. nov. H A. riberae sp. nov. I A. agulhas sp. nov." figureDoi="10.3897/zookeys.963.53470.figure8" httpUri="https://binary.pensoft.net/fig/444405" pageId="0" pageNumber="45">, 8H</figureCitation>
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. South African Agabus spp., microreticulation of male elytral disc A A. austellus sp. nov. holotype, Keurboom, Western Cape B A. austellus sp. nov. Groote Swartberg, Western Cape C A. austellus sp. nov. Sentinel Peak, KZN Drakensberg D A. austellus sp. nov. Harkerville Forest, Western Cape E A. riberae sp. nov. Kamiesberg, Northern Cape F A. agulhas sp. nov. holotype, Rattelrivier, Western Cape." figureDoi="10.3897/zookeys.963.53470.figure9" httpUri="https://binary.pensoft.net/fig/444406" pageId="0" pageNumber="45">, 9E</figureCitation>
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Agabus spp., habitus of males in dorsal view A, F A. sjostedti B, G A. dytiscoides; C, H A. ruwenzoricus D, I A. raffrayi E, J A. pallidus K, O A. anguluverpus sp. nov. L, P A. austellus sp. nov. M, Q A. riberae sp. nov. N, R A. agulhas sp. nov." figureDoi="10.3897/zookeys.963.53470.figure11" httpUri="https://binary.pensoft.net/fig/444408" pageId="0" pageNumber="45">, 11M</figureCitation>
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Agabus spp., habitus of males in dorsal view A, F A. sjostedti B, G A. dytiscoides; C, H A. ruwenzoricus D, I A. raffrayi E, J A. pallidus K, O A. anguluverpus sp. nov. L, P A. austellus sp. nov. M, Q A. riberae sp. nov. N, R A. agulhas sp. nov." figureDoi="10.3897/zookeys.963.53470.figure11" httpUri="https://binary.pensoft.net/fig/444408" pageId="0" pageNumber="45">, 11Q</figureCitation>
<figureCitation captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. The ratio between width of the metacoxal plate and metasternal wing (WC / WS) in the A. raffrayi group (incl. specimens of both sexes). Thick black line inside boxes represents medians, left and right box borders 25 th (Q 1) and 75 th (Q 3) percentiles respectively. Whiskers were calculated with the boxplot. stats function in R using the default coefficient value of 1.5 (drawn to the highest and lowest value within 1.5 * IQD (Inter Quartile Distance = Q 3 - Q 1) away from the 75 th and 25 th percentiles respectively). Note that this character fully separates A. raffrayi from A. pallidus." figureDoi="10.3897/zookeys.963.53470.figure12" httpUri="https://binary.pensoft.net/fig/444409" pageId="0" pageNumber="45">, 12</figureCitation>
<figureCitation captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. The ratio between pronotal width and interocular distance in males of the Agabus raffrayi group. Symbols as in Fig. 12. Note the very narrow pronotum of A. dytiscoides, a distinguishing feature of this species." figureDoi="10.3897/zookeys.963.53470.figure13" httpUri="https://binary.pensoft.net/fig/444410" pageId="0" pageNumber="45">, 13</figureCitation>
<figureCitation captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. The ratio between length of protarsal claw and length of protarsomere 4 in males of the Agabus raffrayi group. Symbols as in Fig. 12. Note the short claws of A. sjostedti and A. dytiscoides." figureDoi="10.3897/zookeys.963.53470.figure14" httpUri="https://binary.pensoft.net/fig/444411" pageId="0" pageNumber="45">, 14</figureCitation>
<figureCitation captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Stream at 1100 m on Witwater-Langkloof road ca 1 km S. of junction, Kamiesberg, Northern Cape Province, South Africa. Type locality of Agabus riberae sp. nov., with DTB. September 19, 2010. Photo Rebecca Bilton." figureDoi="10.3897/zookeys.963.53470.figure18" httpUri="https://binary.pensoft.net/fig/444415" pageId="0" pageNumber="45">, 18</figureCitation>
<figureCitation captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Tra-Tra River at 485 m at Wupperthal, Cederberg range, Western Cape Province, South Africa. Locality for Agabus riberae sp. nov. September 28, 2018. Photo Stacey DeAmicis." figureDoi="10.3897/zookeys.963.53470.figure19" httpUri="https://binary.pensoft.net/fig/444416" pageId="0" pageNumber="45">, 19</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="type locality">
<paragraph pageId="0" pageNumber="45">Type locality.</paragraph>
<paragraph pageId="0" pageNumber="45">South Africa, Northern Cape Province, Kamiesberg, stream on Witwater-Langkloof Road ca 1 km S. of junction, 30°23'41.30&quot;S 18°08'07.95&quot;E, 1100 m.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="type material">
<paragraph pageId="0" pageNumber="45">Type material.</paragraph>
<paragraph pageId="0" pageNumber="45">
<emphasis bold="true" italics="true" pageId="0" pageNumber="45">Holotype</emphasis>
♂ (AMG) labelled: &quot;19/ix/2010 South Africa NC Kamiesberg Stream on Witwater-Langkloof road ca 1 km S. of junction. D.T. Bilton leg.&quot;.
<emphasis bold="true" italics="true" pageId="0" pageNumber="45">Paratypes</emphasis>
: 5 ♂ 5 ♀ (CBP, SANC, ZSM) labelled: &quot;19/ix/2010 South Africa NC Kamiesberg Stream on Witwater-Langkloof road ca 1 km S. of junction. D.T. Bilton leg.&quot;; paratype 4 ♂ 4 ♀ (AMG, CBP, ISAM, NHRS) labelled: &quot;19/1x/2010 South Africa NC Kamiesberg - stream above Studer Pass ca 5 km W of Witwater
<geoCoordinate degrees="30" direction="south" minutes="23" orientation="latitude" precision="1" seconds="13.49" value="-30.387081">30°23'13.49&quot;S</geoCoordinate>
,
<geoCoordinate degrees="18" direction="east" minutes="07" orientation="longitude" precision="1" seconds="05.78" value="18.118273">18°07'05.78&quot;E</geoCoordinate>
1,105 m D T Bilton leg.&quot;; paratype 1 ♀ (IBE) labelled: &quot;22/viii/2004 South Africa NC Kamiesberg - stream at top of Studer Pass G. Challet leg.&quot;; paratype ♂ (CBP) labelled: &quot;19/ix/2014 South Africa NC Kamiesberg temporary stream in Kamiesberg Pass D T Bilton leg.&quot;; paratypes 2 ♂ (CBP) labelled: &quot;18/ix/2014 South Africa NC Kamiesberg spring pool on Rondefontein Farm
<geoCoordinate degrees="30" direction="south" minutes="30" orientation="latitude" precision="1" seconds="05.59" value="-30.501553">30°30'05.59&quot;S</geoCoordinate>
,
<geoCoordinate degrees="18" direction="east" minutes="08" orientation="longitude" precision="1" seconds="56.35" value="18.148987">18°08'56.35&quot;E</geoCoordinate>
1023 m D T Bilton leg.&quot;; paratypes 4 ♂ 1 ♀ (CBP, NHRS) labelled: &quot;17/ix/2014 South Africa NC Kamiesberg stream nr. Damsland on N. side of Rooiberg
<geoCoordinate degrees="30" direction="south" minutes="23" orientation="latitude" precision="1" seconds="36.33" value="-30.393425">30°23'36.33&quot;S</geoCoordinate>
,
<geoCoordinate degrees="18" direction="east" minutes="06" orientation="longitude" precision="1" seconds="32.12" value="18.108923">18°06'32.12&quot;E</geoCoordinate>
1111 m D T Bilton leg.&quot;; paratype 1 ♂ (CBP) labelled: 19/ix/2014 South Africa NC Kamiesberg stream nr. De Kuilen 30°10'44.94&quot;S, 18°04'37.71&quot;E 940 m D T Bilton leg.&quot;; paratypes 2 ♂ (CBP) labelled: &quot;18/ix/2014 South Africa NC Kamiesberg - stream at bottom of Langkloof
<geoCoordinate degrees="30" direction="south" minutes="33" orientation="latitude" precision="1" seconds="16.98" value="-30.554716">30°33'16.98&quot;S</geoCoordinate>
,
<geoCoordinate degrees="18" direction="east" minutes="08" orientation="longitude" precision="1" seconds="19.13" value="18.138647">18°08'19.13&quot;E</geoCoordinate>
594 m D T Bilton leg.&quot;; paratypes 2 ♂ 4 ♀ (AMG, CBP) labelled: &quot;28/ix/2018 South Africa WC Cederberg Tra-Tra river @ Wupperthal
<geoCoordinate degrees="32" direction="south" minutes="16" orientation="latitude" precision="1" seconds="45.35" value="-32.279263">32°16'45.35&quot;S</geoCoordinate>
,
<geoCoordinate degrees="19" direction="east" minutes="13" orientation="longitude" precision="1" seconds="04.32" value="19.217867">19°13'04.32&quot;E</geoCoordinate>
485 m D T Bilton leg.&quot;.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="diagnosis">
<paragraph pageId="0" pageNumber="45">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="45">
Very similar to
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov., differing from this species in having a relatively broad metasternal wing (see Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Morphological characters in the Agabus raffrayi group. WC / WS = width of metacoxal plate / width of metasternal wing, PW / ID = width of pronotum / interocular distance, TL = total body length, N = number of specimens, Min = minimum value, Max = maximum value, SD = standard deviation, NA = not applicable." httpUri="http://table.plazi.org/id/EC20EA9B30576BE179C66F15DBC0C3C6" pageId="0" pageNumber="45" tableUuid="EC20EA9B30576BE179C66F15DBC0C3C6">1</tableCitation>
), and an elytral microreticulation dominated by larger, more irregular meshes than seen in most
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov. specimens (see above). Metatarsomere 5 is also somewhat longer in this species than in most
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="description">
<paragraph pageId="0" pageNumber="45">Description.</paragraph>
<paragraph pageId="0" pageNumber="45">
Habitus as in Fig.
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Agabus spp., habitus of males in dorsal view A, F A. sjostedti B, G A. dytiscoides; C, H A. ruwenzoricus D, I A. raffrayi E, J A. pallidus K, O A. anguluverpus sp. nov. L, P A. austellus sp. nov. M, Q A. riberae sp. nov. N, R A. agulhas sp. nov." figureDoi="10.3897/zookeys.963.53470.figure11" httpUri="https://binary.pensoft.net/fig/444408" pageId="0" pageNumber="45">11M, Q</figureCitation>
.
</paragraph>
<paragraph pageId="0" pageNumber="45">
<emphasis bold="true" italics="true" pageId="0" pageNumber="45">Colour</emphasis>
: Head black with rufous interocular spots and an anterior rufous area. Pronotum black with rufous borders. Elytra rufopiceous to black. Ventral surface black, testaceous lines on abdominal segments rarely present, hypomeron and epipleuron rufotestaceous to rufous. Legs rufous to rufopiceous. Antennae and palpi testaceous.
</paragraph>
<paragraph pageId="0" pageNumber="45">
<taxonomicName genus="Microreticulation" lsidName="Microreticulation" pageId="0" pageNumber="45" rank="genus">
<emphasis bold="true" italics="true" pageId="0" pageNumber="45">Microreticulation</emphasis>
</taxonomicName>
: Relatively fine on both pronotum and elytra, and rather similarly impressed in both sexes. The microreticulation of the elytral disc is typically dominated by relatively large, somewhat irregular meshes (Fig.
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. South African Agabus spp., microreticulation of male elytral disc A A. austellus sp. nov. holotype, Keurboom, Western Cape B A. austellus sp. nov. Groote Swartberg, Western Cape C A. austellus sp. nov. Sentinel Peak, KZN Drakensberg D A. austellus sp. nov. Harkerville Forest, Western Cape E A. riberae sp. nov. Kamiesberg, Northern Cape F A. agulhas sp. nov. holotype, Rattelrivier, Western Cape." figureDoi="10.3897/zookeys.963.53470.figure9" httpUri="https://binary.pensoft.net/fig/444406" pageId="0" pageNumber="45">9E</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="45">
<taxonomicName genus="Structural" lsidName="Structural features" pageId="0" pageNumber="45" rank="species" species="features">
<emphasis bold="true" italics="true" pageId="0" pageNumber="45">Structural features</emphasis>
</taxonomicName>
: Body length: 7.21-8.24 mm (see Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Morphological characters in the Agabus raffrayi group. WC / WS = width of metacoxal plate / width of metasternal wing, PW / ID = width of pronotum / interocular distance, TL = total body length, N = number of specimens, Min = minimum value, Max = maximum value, SD = standard deviation, NA = not applicable." httpUri="http://table.plazi.org/id/EC20EA9B30576BE179C66F15DBC0C3C6" pageId="0" pageNumber="45" tableUuid="EC20EA9B30576BE179C66F15DBC0C3C6">1</tableCitation>
). Hypomeron marginally visible in strict lateral view, lateral bead of pronotum narrow and well defined. Metasternal wing broad, WC/WS 3.0 or less in most specimens (see Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Morphological characters in the Agabus raffrayi group. WC / WS = width of metacoxal plate / width of metasternal wing, PW / ID = width of pronotum / interocular distance, TL = total body length, N = number of specimens, Min = minimum value, Max = maximum value, SD = standard deviation, NA = not applicable." httpUri="http://table.plazi.org/id/EC20EA9B30576BE179C66F15DBC0C3C6" pageId="0" pageNumber="45" tableUuid="EC20EA9B30576BE179C66F15DBC0C3C6">1</tableCitation>
and Fig.
<figureCitation captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. The ratio between width of the metacoxal plate and metasternal wing (WC / WS) in the A. raffrayi group (incl. specimens of both sexes). Thick black line inside boxes represents medians, left and right box borders 25 th (Q 1) and 75 th (Q 3) percentiles respectively. Whiskers were calculated with the boxplot. stats function in R using the default coefficient value of 1.5 (drawn to the highest and lowest value within 1.5 * IQD (Inter Quartile Distance = Q 3 - Q 1) away from the 75 th and 25 th percentiles respectively). Note that this character fully separates A. raffrayi from A. pallidus." figureDoi="10.3897/zookeys.963.53470.figure12" httpUri="https://binary.pensoft.net/fig/444409" pageId="0" pageNumber="45">12</figureCitation>
). Pronotum broad, more than twice as broad as interocular distance (see Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Morphological characters in the Agabus raffrayi group. WC / WS = width of metacoxal plate / width of metasternal wing, PW / ID = width of pronotum / interocular distance, TL = total body length, N = number of specimens, Min = minimum value, Max = maximum value, SD = standard deviation, NA = not applicable." httpUri="http://table.plazi.org/id/EC20EA9B30576BE179C66F15DBC0C3C6" pageId="0" pageNumber="45" tableUuid="EC20EA9B30576BE179C66F15DBC0C3C6">1</tableCitation>
and Fig.
<figureCitation captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. The ratio between pronotal width and interocular distance in males of the Agabus raffrayi group. Symbols as in Fig. 12. Note the very narrow pronotum of A. dytiscoides, a distinguishing feature of this species." figureDoi="10.3897/zookeys.963.53470.figure13" httpUri="https://binary.pensoft.net/fig/444410" pageId="0" pageNumber="45">13</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="45">
<emphasis bold="true" italics="true" pageId="0" pageNumber="45">Legs</emphasis>
: Male protarsal claws long,&gt; 1.6
<normalizedToken originalValue="×">x</normalizedToken>
as long as protarsomere 4 (see Table
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Morphological characters in the Agabus raffrayi group. MT 2 L / MT 2 W = length / width of metatarsomere 2, MT 5 L / MT 5 W = length / width of metatarsomere 5, CL / PT 4 L = length of protarsal claw / length of protarsomere 4, N = number of specimens, Min = minimum value, Max = maximum value, SD = standard deviation, NA = not applicable." httpUri="http://table.plazi.org/id/59FBA533E2ADB4142B7B5363E75795AF" pageId="0" pageNumber="45" tableUuid="59FBA533E2ADB4142B7B5363E75795AF">2</tableCitation>
and Fig.
<figureCitation captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. The ratio between length of protarsal claw and length of protarsomere 4 in males of the Agabus raffrayi group. Symbols as in Fig. 12. Note the short claws of A. sjostedti and A. dytiscoides." figureDoi="10.3897/zookeys.963.53470.figure14" httpUri="https://binary.pensoft.net/fig/444411" pageId="0" pageNumber="45">14</figureCitation>
). Metatarsomere 2 short and broad, &lt;1.8
<normalizedToken originalValue="×">x</normalizedToken>
as long as broad (see Table
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Morphological characters in the Agabus raffrayi group. MT 2 L / MT 2 W = length / width of metatarsomere 2, MT 5 L / MT 5 W = length / width of metatarsomere 5, CL / PT 4 L = length of protarsal claw / length of protarsomere 4, N = number of specimens, Min = minimum value, Max = maximum value, SD = standard deviation, NA = not applicable." httpUri="http://table.plazi.org/id/59FBA533E2ADB4142B7B5363E75795AF" pageId="0" pageNumber="45" tableUuid="59FBA533E2ADB4142B7B5363E75795AF">2</tableCitation>
) in most specimens. Metatarsomere 5 long and slender,&gt; 3.3
<normalizedToken originalValue="×">x</normalizedToken>
as long as broad in most males (see Table
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Morphological characters in the Agabus raffrayi group. MT 2 L / MT 2 W = length / width of metatarsomere 2, MT 5 L / MT 5 W = length / width of metatarsomere 5, CL / PT 4 L = length of protarsal claw / length of protarsomere 4, N = number of specimens, Min = minimum value, Max = maximum value, SD = standard deviation, NA = not applicable." httpUri="http://table.plazi.org/id/59FBA533E2ADB4142B7B5363E75795AF" pageId="0" pageNumber="45" tableUuid="59FBA533E2ADB4142B7B5363E75795AF">2</tableCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="45">
<emphasis bold="true" italics="true" pageId="0" pageNumber="45">Male genitalia</emphasis>
: Tip of aedeagus short, lacking the prolongation of the area located between the subapical broadening and the apical and subapical tooth present in some species in the group (see Fig.
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Agabus spp., aedeagus in lateral view A A. sjostedti B A. dytiscoides C A. ruwenzoricus D A. raffrayi E A. pallidus F A. anguluverpus sp. nov. G A. austellus sp. nov. H A. riberae sp. nov. I A. agulhas sp. nov." figureDoi="10.3897/zookeys.963.53470.figure8" httpUri="https://binary.pensoft.net/fig/444405" pageId="0" pageNumber="45">8H</figureCitation>
). There is some variation in the shape and size of the subapical tooth (Fig.
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. South African Agabus spp., aedeagal apices, lateral view A A. austellus sp. nov. holotype, Keurboom, Western Cape B A. austellus sp. nov., Gydo Pass, Western Cape C A. austellus sp. nov. Groote Swartberg, Western Cape D A. austellus sp. nov. Sentinel Peak, KZN Drakensberg E A. riberae sp. nov. holotype, Kamiesberg, Northern Cape; F A. riberae sp. nov. paratype, Kamiesberg, Northern Cape G A. agulhas sp. nov. holotype, Rattelrivier, Western Cape." figureDoi="10.3897/zookeys.963.53470.figure6" httpUri="https://binary.pensoft.net/fig/444403" pageId="0" pageNumber="45">6E, F</figureCitation>
), this being relatively long and narrow in most specimens examined (Fig.
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. South African Agabus spp., aedeagal apices, lateral view A A. austellus sp. nov. holotype, Keurboom, Western Cape B A. austellus sp. nov., Gydo Pass, Western Cape C A. austellus sp. nov. Groote Swartberg, Western Cape D A. austellus sp. nov. Sentinel Peak, KZN Drakensberg E A. riberae sp. nov. holotype, Kamiesberg, Northern Cape; F A. riberae sp. nov. paratype, Kamiesberg, Northern Cape G A. agulhas sp. nov. holotype, Rattelrivier, Western Cape." figureDoi="10.3897/zookeys.963.53470.figure6" httpUri="https://binary.pensoft.net/fig/444403" pageId="0" pageNumber="45">6F</figureCitation>
), but with both narrow and broader teeth being observed within the same population.
</paragraph>
<paragraph pageId="0" pageNumber="45">
<emphasis bold="true" pageId="0" pageNumber="45">Female</emphasis>
: Externally similar to males. Some specimens with dorsal microreticulation slightly more strongly impressed.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="distribution">
<paragraph pageId="0" pageNumber="45">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="45">
To date known only from Kamiesberg Range in the Northern Cape Province, and the eastern fringes of the Cederberg, Western Cape Province, Republic of South Africa (see Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Distribution map of the Agabus raffrayi group. Relief is expressed in meters above sea level." figureDoi="10.3897/zookeys.963.53470.figure1" httpUri="https://binary.pensoft.net/fig/444398" pageId="0" pageNumber="45">1</figureCitation>
), material from both areas being confirmed from COI sequences. The Kamiesberg represents a northerly outlier of Fynbos and Renosterveld vegetation in predominantly arid Namaqualand, and consequently have a diverse flora with a number of localised endemics (
<bibRefCitation author="Helme, N" journalOrPublisher=")" pageId="0" pageNumber="45" publicationUrl="http://www.cepf.net/Documents/helme_desmet.pdf" refId="B8" refString="Helme, N, Desmet, PG, 2006. A description of the endemic flora and vegetation of the Kamiesberg Uplands, Namaqualand, South Africa. Report for CEPF/SKEP. http://www.cepf.net/Documents/helme_desmet.pdf" title="A description of the endemic flora and vegetation of the Kamiesberg Uplands, Namaqualand, South Africa. Report for CEPF / SKEP." url="http://www.cepf.net/Documents/helme_desmet.pdf" year="2006">Helme and Desmet 2006</bibRefCitation>
). The mountains support the bulk of the global population of the endemic dytiscid
<taxonomicName authorityName="Sharp" authorityYear="1882" class="Insecta" family="Dytiscidae" genus="Andex" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Andex insignis" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="species" species="insignis">
<emphasis italics="true" pageId="0" pageNumber="45">Andex insignis</emphasis>
</taxonomicName>
Sharp, 1882 and a number of new, apparently endemic, water beetles have been described from the area in recent years (
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.3626.4.13" author="Bilton, DT" journalOrPublisher="Zootaxa" pageId="0" pageNumber="45" pagination="589 - 592" refId="B1" refString="Bilton, DT, 2013. Crenitis bicolor sp. n. from the Kamiesberg of South Africa (Coleoptera: Hydrophilidae). Zootaxa 3626: 589 - 592, DOI: https://doi.org/10.11646/zootaxa.3626.4.13" title="Crenitis bicolor sp. n. from the Kamiesberg of South Africa (Coleoptera: Hydrophilidae)." url="https://doi.org/10.11646/zootaxa.3626.4.13" volume="3626" year="2013">Bilton 2013</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.3957.4.5" author="Bilton, DT" journalOrPublisher="Zootaxa" pageId="0" pageNumber="45" pagination="441 - 454" refId="B2" refString="Bilton, DT, 2015. A review of the Canthyporus exilis group, with the description of two new species (Coleoptera: Dytiscidae). Zootaxa 3957: 441 - 454, DOI: https://doi.org/10.11646/zootaxa.3957.4.5" title="A review of the Canthyporus exilis group, with the description of two new species (Coleoptera: Dytiscidae)." url="https://doi.org/10.11646/zootaxa.3957.4.5" volume="3957" year="2015">2015</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4137.4.11" author="Bilton, DT" journalOrPublisher="Zootaxa" pageId="0" pageNumber="45" pagination="585 - 591" refId="B3" refString="Bilton, DT, 2016. Two new water beetles from the South African Cape (Coleoptera, Hydraenidae). Zootaxa 4137: 585 - 591, DOI: https://doi.org/10.11646/zootaxa.4137.4.11" title="Two new water beetles from the South African Cape (Coleoptera, Hydraenidae)." url="https://doi.org/10.11646/zootaxa.4137.4.11" volume="4137" year="2016">2016</bibRefCitation>
).
<taxonomicName authorityName="Bilton, Englund &amp; Bergsten" authorityYear="2020" class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus riberae" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="species" species="riberae">
<emphasis italics="true" pageId="0" pageNumber="45">Agabus riberae</emphasis>
</taxonomicName>
sp. nov. appears to be the only
<taxonomicName class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="45">Agabus</emphasis>
</taxonomicName>
present in Kamiesberg, where it is abundant. In the Cederberg the species has been found close to Wupperthal, in the relatively dry northeastern fringes of the range.
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov. is the only species so far recorded from the wetter central areas of the Cederberg. All sites known to date experience predominantly winter rainfall.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="ecology">
<paragraph pageId="0" pageNumber="45">Ecology.</paragraph>
<paragraph pageId="0" pageNumber="45">
Found in streams and associated pools in the Kamiesberg and northeastern Cederberg ranges (Figs
<figureCitation captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Stream at 1100 m on Witwater-Langkloof road ca 1 km S. of junction, Kamiesberg, Northern Cape Province, South Africa. Type locality of Agabus riberae sp. nov., with DTB. September 19, 2010. Photo Rebecca Bilton." figureDoi="10.3897/zookeys.963.53470.figure18" httpUri="https://binary.pensoft.net/fig/444415" pageId="0" pageNumber="45">18</figureCitation>
,
<figureCitation captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Tra-Tra River at 485 m at Wupperthal, Cederberg range, Western Cape Province, South Africa. Locality for Agabus riberae sp. nov. September 28, 2018. Photo Stacey DeAmicis." figureDoi="10.3897/zookeys.963.53470.figure19" httpUri="https://binary.pensoft.net/fig/444416" pageId="0" pageNumber="45">19</figureCitation>
), in either Fynbos or Renosterveld vegetation between 480 and 1000 m elevation. Typically netted from marginal vegetation, including at the base of tussocks. Also found amongst grasses in a spring pool with cold water. Ecological differences between this species and
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov. are unclear, but may relate, at least in part, to rainfall.
</paragraph>
<caption doi="10.3897/zookeys.963.53470.figure18" httpUri="https://binary.pensoft.net/fig/444415" pageId="0" pageNumber="45" start="Figure 18" startId="F18">
<paragraph pageId="0" pageNumber="45">
<emphasis bold="true" pageId="0" pageNumber="45">Figure 18.</emphasis>
Stream at 1100 m on Witwater-Langkloof road ca 1 km S. of junction, Kamiesberg, Northern Cape Province, South Africa. Type locality of
<taxonomicName authorityName="Bilton, Englund &amp; Bergsten" authorityYear="2020" class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus riberae" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="species" species="riberae">
<emphasis italics="true" pageId="0" pageNumber="45">Agabus riberae</emphasis>
</taxonomicName>
sp. nov., with DTB. September 19, 2010. Photo Rebecca Bilton.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="etymology">
<paragraph pageId="0" pageNumber="45">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="45">Named after our late friend and colleague Ignacio Ribera, who will be sorely missed.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="comments">
<paragraph pageId="0" pageNumber="45">Comments.</paragraph>
<paragraph pageId="0" pageNumber="45">
Despite the relatively minor morphological differences between this species and
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov. we consider these two taxa distinct. COI sequences for four specimens of
<taxonomicName lsidName="A. riberae" pageId="0" pageNumber="45" rank="species" species="riberae">
<emphasis italics="true" pageId="0" pageNumber="45">A. riberae</emphasis>
</taxonomicName>
sp. nov. investigated differed by 4.5-4.7% from
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov., more than that observed between many well-established species in the subgenus
<taxonomicName lsidName="(Acatodes)" pageId="0" pageNumber="45" rank="subGenus" subGenus="Acatodes">Acatodes</taxonomicName>
Thomson, 1859 (I. Ribera pers. comm.). This is supportive of the recognition as a distinct taxon, particularly given the relative uniformity in COI sequence observed across the wide geographical range of
<taxonomicName lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
</subSubSection>
</treatment>
</document>