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<document ID-CLB-Dataset="21142" 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="49160C6A22265941BC5E7342B0895666" docLanguage="en" docName="ZooKeys 963: 45-79" docOrigin="ZooKeys 963" docSource="http://dx.doi.org/10.3897/zookeys.963.53470" docTitle="Agabus agulhas Englund &amp; Njoroge &amp; Biström &amp; Miller &amp; Bilton &amp; Bergsten 2020, sp. nov." docType="treatment" docUuid="FD6B9312-065E-4B96-9FDF-9B1C45EA8D69" docUuidSource="ZooBank" docVersion="6" 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="1732748813246" updateUser="ExternalLinkService">
<mods:mods id="518E31E7DF5AC33E842709958C90D208" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="CB6708B7847BBA52888EAA3818D2EE41">
<mods:title id="42D9CC3A8AE49E55B1FA4D9A62435007">Taxonomic revision of the Afrotropical Agabus raffrayi species group with the description of four new species (Coleoptera, Dytiscidae)</mods:title>
</mods:titleInfo>
<mods:name id="FFDE6A65F61C0D9B7C1F729E6F82A98A" type="personal">
<mods:role id="B1212B9722AA1850D5FBE295EFCBF23F">
<mods:roleTerm id="4D2A4DEB785A556B96F9E4380D4202D8">Author</mods:roleTerm>
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<mods:namePart id="73BE56D7E4372B7805AC97F91838BE10">Englund, William F.</mods:namePart>
<mods:affiliation id="7870F60459548865F65BBF63BCC358F0">Swedish Museum of Natural History, Department of Zoology, Box 50007, SE- 10405 Stockholm, Sweden</mods:affiliation>
</mods:name>
<mods:name id="AB83EBC37DFDCCA5F785FC647E5A3292" type="personal">
<mods:role id="2AA7A4A019131F44BB1188B24748F09F">
<mods:roleTerm id="7F2B22CFC985B9C76AEE80B5F42342F8">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="93A95D9C1035FBF0E02E209CE5A0C64B">Njoroge, Laban</mods:namePart>
<mods:affiliation id="17517605BACC40C4C400BD8CCD6FFA8E">National Museums of Kenya, Section of Invertebrate Zoology, Museum Hill, P. O. BOX 40658 - 00100, Nairobi, Kenya</mods:affiliation>
</mods:name>
<mods:name id="394EFF264C317F677A0CD8131CFC2798" type="personal">
<mods:role id="C2EC202A15BBBBAC54076041B735574D">
<mods:roleTerm id="FB8384DB52482B77F695271DC8B74225">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="6D46E913BACCBF3E07C78453F41327F8">Bistroem, Olof</mods:namePart>
<mods:affiliation id="966F8A98BA166D5437C76A5351F01456">Finnish Museum of Natural History, Zoology Unit, P. O. Box 17, FI- 00014 University of Helsinki, Finland</mods:affiliation>
</mods:name>
<mods:name id="CE84F4329BC938C774877E421F49FAD9" type="personal">
<mods:role id="2B9363FFD0CB6DFAA1814C6056176D37">
<mods:roleTerm id="FFA98651A9B7F84C7CE05CE6D70D6A08">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="9AEF5DAAB60105657398C0673D53E874">Miller, Kelly B.</mods:namePart>
<mods:affiliation id="5B1EF7864CE1640E0652A636EF4B5251">Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131 - 0001, USA</mods:affiliation>
</mods:name>
<mods:name id="B536FD1FA33B2D9E0927B26C81D8135F" type="personal">
<mods:role id="D2C2465838095264789414830DC2C1A3">
<mods:roleTerm id="8FF6546B4F0B929ED627D2A5A9122812">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="247C011BA7BB46E2FFB85924D090AA18">Bilton, David T.</mods:namePart>
<mods:affiliation id="84CCBE789CD11C023C5020C12FC40B9D">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 id="964D7A7D5EC68FD9C9FE29A1E016A695" type="personal">
<mods:role id="A4A299150A2C8D9090361C2DE9FB7C4F">
<mods:roleTerm id="22ED7E4BF0D6012CBA831C408D445DF5">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="A7E2D1B48B059B99E7AFBE3F97F69D0C">Bergsten, Johannes</mods:namePart>
<mods:affiliation id="3FE8FB622C374972499D191BDD5051CE">Swedish Museum of Natural History, Department of Zoology, Box 50007, SE- 10405 Stockholm, Sweden</mods:affiliation>
<mods:nameIdentifier id="A4BB59D8C53F10298A7BA35BB0E04ABF" type="email">johannes.bergsten@nrm.se</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource id="A63D57DFB3B71E758C0C649723C325CC">text</mods:typeOfResource>
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<mods:titleInfo id="C688647919B1BE0199E7885C8BFBAC97">
<mods:title id="F0642B9D42FC3E99E66E7F2814B491EA">ZooKeys</mods:title>
</mods:titleInfo>
<mods:part id="D1A724160D338003CE4B9FBA32FF4759">
<mods:date id="F0D994683470377FB24D859CC7FED944">2020</mods:date>
<mods:detail id="7EEBDC926A54770498D9CC85B2C864CE" type="volume">
<mods:number id="34CD99D8E0062524501C599855B58AC3">963</mods:number>
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<mods:start id="DD86FFF85E68270CA05AB95FCD541594">45</mods:start>
<mods:end id="9908EB6412CF4239028C4E9BBA71A622">79</mods:end>
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<mods:location id="65E2634CF0E08AFEA0F96F25D9888F23">
<mods:url id="5D8D98060A40D14571527E3053AAE122">http://dx.doi.org/10.3897/zookeys.963.53470</mods:url>
</mods:location>
<mods:classification id="BEAD4F2E3681940009E0CEE620357B12">journal article</mods:classification>
<mods:identifier id="3E84DC16AA1F003AC4C74CADA899F84A" type="DOI">http://dx.doi.org/10.3897/zookeys.963.53470</mods:identifier>
<mods:identifier id="7904E3DA48FA3A2753B38B26E05AC544" type="Pensoft-Pub">1313-2970-963-45</mods:identifier>
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<treatment id="49160C6A22265941BC5E7342B0895666" ID-GBIF-Taxon="167293854" LSID="urn:lsid:zoobank.org:act:FD6B9312-065E-4B96-9FDF-9B1C45EA8D69" httpUri="http://treatment.plazi.org/id/49160C6A22265941BC5E7342B0895666" lastPageNumber="45" pageId="0" pageNumber="45" scope_family="Dytiscidae" scope_order="Coleoptera">
<subSubSection id="E9413C6F03741FAFCD10BC4E0F3CCCCD" pageId="0" pageNumber="45" type="nomenclature">
<paragraph id="93F4A39AC84EE3E9866C5492A1174DB7" pageId="0" pageNumber="45">
<taxonomicName id="EEEDFEBB8F6F6C50979061B17F7F9599" ID-CoL="8LZV9" LSID="49160C6A-2226-5941-BC5E-7342B0895666" authority="Bilton, Englund &amp; Bergsten" authorityName="Englund &amp; Njoroge &amp; Biström &amp; Miller &amp; Bilton &amp; Bergsten" authorityYear="2020" class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus agulhas" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="species" species="agulhas" status="sp. nov.">Agabus agulhas Bilton, Englund &amp; Bergsten</taxonomicName>
<taxonomicNameLabel id="9068B3B24226FE7BCA401513D99745DF" pageId="0" pageNumber="45">sp. nov.</taxonomicNameLabel>
<figureCitation id="D24544DEB51E8002F915A75463FAF611" 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 id="DB36324CD46E66F2630FDF690B3EDA85" 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">, 6G</figureCitation>
<figureCitation id="04E8C87C8312489F8F3FEBE487AC2A04" 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">, 8I</figureCitation>
<figureCitation id="54135B00DC79C18C8133DB56EFDF600D" 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">, 9F</figureCitation>
<figureCitation id="04941D2FDC44E9E1C81FBD53E19C3FD3" 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">, 11N</figureCitation>
<figureCitation id="BCD48CB5B6E5DAA391B59D23B6125704" 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">, 11R</figureCitation>
<figureCitation id="EDC277021F21036E180196F66CA0D087" 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 id="8ADEEC1264D59B7F8BF3E57A6B9466C7" 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 id="7419A204D1842C577075CB14F2A7EC60" 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>
</paragraph>
</subSubSection>
<subSubSection id="A18C667BE7910F4A3435A7451F0B311F" pageId="0" pageNumber="45" type="type locality">
<paragraph id="5F443BCC311FBAAEBA765DE1AD5B00B5" pageId="0" pageNumber="45">Type locality.</paragraph>
<paragraph id="ADB3A95D1BFEE68F6DA3731939CC979F" pageId="0" pageNumber="45">
South Africa, Western Cape Province, Rooistrandveld, Bredasdorp, natural viei beside road to Die Dam at Ratelrivier
<geoCoordinate id="462970B751ED142E0A20AE855105A702" degrees="34" direction="south" minutes="43" orientation="latitude" precision="1" seconds="00.47" value="-34.716797">34°43'00.47&quot;S</geoCoordinate>
,
<geoCoordinate id="74080F3E81444DBD983697320668D9AF" degrees="19" direction="east" minutes="41" orientation="longitude" precision="1" seconds="53.81" value="19.69828">19°41'53.81&quot;E</geoCoordinate>
, 31 m.
</paragraph>
</subSubSection>
<subSubSection id="B353CAD38EC8FE183CA8D5311AAB47AF" pageId="0" pageNumber="45" type="type material">
<paragraph id="7058AB9DEF6A5C4AE13295ADB2A79F3E" pageId="0" pageNumber="45">Type material.</paragraph>
<paragraph id="9E65A09C0260AE505AAD73063AC8A24C" pageId="0" pageNumber="45">
<emphasis id="2A2A9FC32643D682C4C6F6F55A4D4333" bold="true" italics="true" pageId="0" pageNumber="45">Holotype</emphasis>
♂ (AMG) labelled: &quot;26/ix/2010 South Africa WC Rooistrandveld, Bredasdorp natural viei beside road to Die Dam @ Ratelrivier FW marsh with tussocks etc. D. T. Bilton leg.&quot;.
<emphasis id="CECBE2BEEA5E0EEE4C22D70DD2A56953" bold="true" italics="true" pageId="0" pageNumber="45">Paratypes</emphasis>
3 ♂ 2 ♀ (AMG, CBP, NHRS, ZSM) labelled: &quot;26/ix/2010 South Africa WC Rooistrandveld, Bredasdorp natural viei beside road to Die Dam @ Ratelrivier FW marsh with tussocks etc. D. T. Bilton leg.&quot;.
</paragraph>
</subSubSection>
<subSubSection id="E6F3DA95D750410944C27EB776C8EA84" pageId="0" pageNumber="45" type="diagnosis">
<paragraph id="6C8DF76CD35A689E51A91999FDA93F85" pageId="0" pageNumber="45">Diagnosis.</paragraph>
<paragraph id="B1D4925697A92FED692C4755FF43A73C" pageId="0" pageNumber="45">
Very similar to
<taxonomicName id="D272006D3037259C175003752F52F3CA" lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis id="54DCFAE8FD3BEEE6E31D4D7B111DB888" italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName id="A15D6DB854081416D92ED83E49CFE216" lsidName="A. riberae" pageId="0" pageNumber="45" rank="species" species="riberae">
<emphasis id="4DB7313F8CF99C15DBA77DBB831CB266" italics="true" pageId="0" pageNumber="45">A. riberae</emphasis>
</taxonomicName>
sp. nov., but distinguishable by the distinctly curved base of the aedeagal subapical tooth (compare Fig.
<figureCitation id="972104501C9F67FB7A95972F398D5D4C" 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">8I</figureCitation>
with Fig.
<figureCitation id="B375A4FF89CCF0A34C05A112847072C4" 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">8G, H</figureCitation>
and see Fig.
<figureCitation id="C88966E905FF858D5B897A28EA9FBC31" 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">6G</figureCitation>
), the scutellum being lighter than the elytra and its relatively narrow metasternal wing (see Table
<tableCitation id="0C4BFCC11FBA8285EACBECFE507E21EA" 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 id="D7E76E46B574275B8AEF631979FFE78D" 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>
).
</paragraph>
</subSubSection>
<subSubSection id="258AC5C4B360372E46921D6267A06D81" pageId="0" pageNumber="45" type="description">
<paragraph id="79C25EAB804869BE97B4098B61090771" pageId="0" pageNumber="45">Description.</paragraph>
<paragraph id="05A4864FDD94568248DE9D2060772001" pageId="0" pageNumber="45">
Habitus as in Fig.
<figureCitation id="7550692EAECF0C4C2B4BBC4D065AE15F" 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">11N, R</figureCitation>
.
</paragraph>
<paragraph id="0C7C785D953E3BC55DC3B785F8F6415C" pageId="0" pageNumber="45">
<emphasis id="691F8BEB8C50B617BA81D88897D09F30" bold="true" italics="true" pageId="0" pageNumber="45">Colour</emphasis>
: Head black with weak rufous interocular spots and an anterior rufous area. Pronotum black with slightly rufous margins. Elytra blackish brown to black, with a lighter scutellum. Ventral surface black, testaceous lines on abdominal segments reduced or absent, hypomeron and epipleuron rufotestaceous to rufous. Legs rufopiceous to rufous. Antennae and palpi testacous.
</paragraph>
<paragraph id="40DB369586B2086255B6766BAC9257A8" pageId="0" pageNumber="45">
<taxonomicName id="CF1061A2B6645DD6FC2B00E9F4F91246" genus="Microreticulation" lsidName="Microreticulation" pageId="0" pageNumber="45" rank="genus">
<emphasis id="593CCECB6F55E80AC6DB5933CF2AEECA" bold="true" italics="true" pageId="0" pageNumber="45">Microreticulation</emphasis>
</taxonomicName>
: Relatively fine on both pronotum and elytra, and slightly more impressed in females. The microreticulation of the elytral disc is composed of a mix of small and larger, somewhat irregular meshes (Fig.
<figureCitation id="E5D49A9666D97599F29105030705BEB1" 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">9F</figureCitation>
).
</paragraph>
<paragraph id="865D37F80C69F92598565B82C095BEC4" pageId="0" pageNumber="45">
<taxonomicName id="2B5EB7A74114A571613AB4DF99524675" genus="Structural" lsidName="Structural features" pageId="0" pageNumber="45" rank="species" species="features">
<emphasis id="DEB02F2D5875083D25DED758B691EF44" bold="true" italics="true" pageId="0" pageNumber="45">Structural features</emphasis>
</taxonomicName>
: Body length: 7.60-8.00 mm (see Table
<tableCitation id="7B96793D954569289327C627E01E92E9" 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 very narrow, WC/WS&gt; 3.6 in all specimens (see Table
<tableCitation id="2113CC22EF39F4612FE1FD8291D438FE" 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 id="4C54D192E9190AAFEA8434E4FDF8CD33" 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 id="1EE948091AC4AB26716AFF29119BBD89" 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 id="5CB417FE2C40EB3BDB446CF7C772BB1E" 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 id="C440FD3D6B384C71DE70F6F0A73284C6" pageId="0" pageNumber="45">
<emphasis id="8DCDA4C6B652FD0A12729F5F54357E92" bold="true" italics="true" pageId="0" pageNumber="45">Legs</emphasis>
: Protarsal claws very long,&gt; 1.8
<normalizedToken id="667A3759EFD6664110E954CBB7699456" originalValue="×">x</normalizedToken>
as long as protarsomere 4 in all males (see Table
<tableCitation id="125D096A5D7A4E7DD562E5A5EE59E605" 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>
&amp; Fig.
<figureCitation id="C9541921658CB34CCECEE9F2B3EE3EDF" 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>
). Metatarsomeres short and broad; metatarsomere 2 &lt;1.8
<normalizedToken id="01A4680E74A6D0406B8620CF2670A7C8" originalValue="×">x</normalizedToken>
as long as broad (see Table
<tableCitation id="E325DEC8758745BFAD3995E19E9F31CB" 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>
), metatarsomere 5 &lt;3
<normalizedToken id="76F274CDC621435BEFC39EEEC08B1328" originalValue="×">x</normalizedToken>
as long as broad in all specimens (see Table
<tableCitation id="D8A7B23C33E3C0A75BA54491407BAFD4" 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 id="3057B3A40AD566EB3DD1E8576CA01725" pageId="0" pageNumber="45">
<emphasis id="F5D8B5BB440F75A7FD05428E9056DB00" bold="true" italics="true" pageId="0" pageNumber="45">Male genitalia</emphasis>
: Aedeagus without the prolonged section between subapical broadening and the apical and subapical teeth which is present in some species in the group. In ventral view the apex is asymmetrically curved. Base of subapical tooth distinctly curved basally (see Figs
<figureCitation id="C3652CF511DBE7BED4B5AC28777CE697" 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">8I</figureCitation>
,
<figureCitation id="72EE058384D6A3241F8284936FFE7639" 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">6G</figureCitation>
).
</paragraph>
<paragraph id="A379C49EF1CE4FB2705F4782CD73308E" pageId="0" pageNumber="45">
<emphasis id="C73E187DA77309B07F94B6513D0B8222" bold="true" pageId="0" pageNumber="45">Female</emphasis>
: Externally similar to males. Dorsal microreticulation slightly more impressed than in males.
</paragraph>
</subSubSection>
<subSubSection id="594790CE9A1D9FB63038662F0E706AFB" pageId="0" pageNumber="45" type="distribution">
<paragraph id="B4BA662BEB3B569723EA307CD3C3D1F8" pageId="0" pageNumber="45">Distribution.</paragraph>
<paragraph id="EC031BD8FFD7127AE58D16BED100CCA5" pageId="0" pageNumber="45">
Only known from the type locality, a lowland valley wetland at 31 m on the Agulhas Plain, Western Cape Province, Republic of South Africa (see Fig.
<figureCitation id="68DD5DA0AC078B94C24CFF930BDC3EA1" 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>
). The most southerly distributed
<taxonomicName id="ED5D79AA3E8760AA7BE5FD09798D4FE1" class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="genus">
<emphasis id="7C7E1CF62159D8266A2D276C3934CACD" italics="true" pageId="0" pageNumber="45">Agabus</emphasis>
</taxonomicName>
species in the world.
</paragraph>
</subSubSection>
<subSubSection id="619DBDA409755F747FE71BF395DC3B52" pageId="0" pageNumber="45" type="ecology">
<paragraph id="063D45BE035D3005160ACFF0781DAD00" pageId="0" pageNumber="45">Ecology.</paragraph>
<paragraph id="B9483A8EA29AB0E6E1976F40E2F9C906" pageId="0" pageNumber="45">Collected from the base of large tussocks in a valley wetland. Despite having largely lentic conditions, this is likely to experience some seepage flow, particularly following periods of high rainfall in winter and spring.</paragraph>
</subSubSection>
<subSubSection id="1CC45A13BAFB4A5BDEE48DA23864A0B6" pageId="0" pageNumber="45" type="etymology">
<paragraph id="910C5983688E228942B376881599F2EA" pageId="0" pageNumber="45">Etymology.</paragraph>
<paragraph id="494311F73E64C391475F040C566E5439" pageId="0" pageNumber="45">
Named after the Agulhas Plain, on which the type locality is situated. The Agulhas Plain is itself named in reference to nearby Cape Agulhas (Portuguese - Cabo das Agulhas = Cape of Needles), the most southerly point on the African continent. As with other members of the species group,
<taxonomicName id="C353994A5DA670AE80974B6DD89E97DB" lsidName="A. agulhas" pageId="0" pageNumber="45" rank="species" species="agulhas">
<emphasis id="EA7BE881523F2820334D7E728C5453D2" italics="true" pageId="0" pageNumber="45">A. agulhas</emphasis>
</taxonomicName>
sp. nov. has sharp, needle-like, teeth at the aedeagal apex.
</paragraph>
</subSubSection>
<subSubSection id="A2494BE5C3C256A327633771884F7AD8" pageId="0" pageNumber="45" type="comments">
<paragraph id="788395F6C45B98074273DBBA96EE12F4" pageId="0" pageNumber="45">Comments.</paragraph>
<paragraph id="4F124C41A38FA7A99059BCB506C84292" pageId="0" pageNumber="45">
COI sequence divergence between
<taxonomicName id="7418CE6B7E7C975A992084CEDCADA097" lsidName="A. agulhas" pageId="0" pageNumber="45" rank="species" species="agulhas">
<emphasis id="13C12F847F8E7FA0CC00AE7F329AF0FD" italics="true" pageId="0" pageNumber="45">A. agulhas</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName id="1125E8AE216B07FBEE9C9C121FFC104B" lsidName="A. austellus" pageId="0" pageNumber="45" rank="species" species="austellus">
<emphasis id="F4E99916123DE64D929EE993FD9AC320" italics="true" pageId="0" pageNumber="45">A. austellus</emphasis>
</taxonomicName>
sp. nov. ranges from 3.9 to 4.7%; that between
<taxonomicName id="FD4364CC1AC1DE2F17A28FF5D83F3BB1" lsidName="A. agulhas" pageId="0" pageNumber="45" rank="species" species="agulhas">
<emphasis id="A6BA044644ACCBD0E59D0EACE62FF38B" italics="true" pageId="0" pageNumber="45">A. agulhas</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName id="CCA1F605907699BFA591879C1AA5A5D5" lsidName="A. riberae" pageId="0" pageNumber="45" rank="species" species="riberae">
<emphasis id="5C340B9C2E15A691858DBD11D9E6BC6E" italics="true" pageId="0" pageNumber="45">A. riberae</emphasis>
</taxonomicName>
sp. nov. being 6.4% (I. Ribera, pers. comm.).
</paragraph>
<caption id="6A88A54CF9DC93BA7C3D1AD1EC3BFAD9" doi="10.3897/zookeys.963.53470.figure19" httpUri="https://binary.pensoft.net/fig/444416" pageId="0" pageNumber="45" start="Figure 19" startId="F19">
<paragraph id="EDE9653C1111BEBB13E1D7267CE593E5" pageId="0" pageNumber="45">
<emphasis id="09151F1E94295873252983C67A933114" bold="true" pageId="0" pageNumber="45">Figure 19.</emphasis>
Tra-Tra River at 485 m at Wupperthal, Cederberg range, Western Cape Province, South Africa. Locality for
<taxonomicName id="BB81B33CEDE3272A4A94BE6B4F5D8982" 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 id="C309FA089DC070A53D34159985472C48" italics="true" pageId="0" pageNumber="45">Agabus riberae</emphasis>
</taxonomicName>
sp. nov. September 28, 2018. Photo Stacey DeAmicis.
</paragraph>
</caption>
<caption id="763C8900A5D27354ACCD146E73FB30DA" doi="10.3897/zookeys.963.53470.figure20" httpUri="https://binary.pensoft.net/fig/444417" pageId="0" pageNumber="45" start="Figure 20" startId="F20">
<paragraph id="045477702EC9D37F4E66B8271D98C67F" pageId="0" pageNumber="45">
<emphasis id="5A328D0F8C21CB9883BAD2A48BA0AD41" bold="true" pageId="0" pageNumber="45">Figure 20.</emphasis>
Lake Ellis on Mount Kenya, Kenya. Type locality for
<taxonomicName id="8C8B00812624043C07904D2A4EB3155F" authorityName="Englund, Njoroge &amp; Bergsten" authorityYear="2020" class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus anguluverpus" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="species" species="anguluverpus">
<emphasis id="D7F04DEB64D72D053ECC1BC97F13E6B9" italics="true" pageId="0" pageNumber="45">Agabus anguluverpus</emphasis>
</taxonomicName>
sp. nov. September 17, 2015. Photo Wanyoike Wamiti.
</paragraph>
</caption>
<caption id="520163A31FA50181AA078227C41140E5" doi="10.3897/zookeys.963.53470.figure21" httpUri="https://binary.pensoft.net/fig/444418" pageId="0" pageNumber="45" start="Figure 21" startId="F21">
<paragraph id="A7AA250F44D208C2E9D906D548ED70E0" pageId="0" pageNumber="45">
<emphasis id="2E1963035C04F62A60FFFC327871D149" bold="true" pageId="0" pageNumber="45">Figure 21.</emphasis>
Part of Lake Ellis on Mount Kenya, Kenya. Type locality for
<taxonomicName id="618763011726823D44DC08E4E50E4030" authorityName="Englund, Njoroge &amp; Bergsten" authorityYear="2020" class="Insecta" family="Dytiscidae" genus="Agabus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Agabus anguluverpus" order="Coleoptera" pageId="0" pageNumber="45" phylum="Arthropoda" rank="species" species="anguluverpus">
<emphasis id="AC89A00041AD6B2AA6880E44757A84D0" italics="true" pageId="0" pageNumber="45">Agabus anguluverpus</emphasis>
</taxonomicName>
sp. nov. September 17, 2015. Photo Wanyoike Wamiti.
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>
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