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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">SATNT</journal-id>
<journal-title-group>
<journal-title>Suid-Afrikaanse Tydskrif vir Natuurwetenskap en Tegnologie</journal-title>
</journal-title-group>
<issn pub-type="ppub">0254-3486</issn>
<issn pub-type="epub">2222-4173</issn>
<publisher>
<publisher-name>AOSIS OpenJournals</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">SATNT-34-1286</article-id>
<article-id pub-id-type="doi">10.4102/satnt.v34i1.1286</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Oorspronklike Navorsing</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Dopluise (Hemiptera: Coccoidea) geassosieer met die wipstertmier, <italic>Crematogaster peringueyi</italic> Emery (Hymenoptera: Formicidae)</article-title>
<alt-title alt-title-type="running-title">Dopluise (Hemiptera: Coccoidea) geassosieer met die wipstertmier, <italic>Crematogaster peringueyi</italic> Emery</alt-title>
<alt-title alt-title-type="running-authors">Johannes H. Giliomee</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Giliomee</surname>
<given-names>Johannes H.</given-names>
</name>
<xref ref-type="aff" rid="AF0001">1</xref>
</contrib>
<aff id="AF0001"><label>1</label>Centre for Invasion Biology, Department of Botany and Zoology, Stellenbosch University, South Africa</aff>
</contrib-group>
<author-notes>
<corresp id="cor1"><bold>Correspondence to</bold>: Johannes Giliomee  <bold>Email</bold>: <email xlink:href="jhg@sun.ac.za">jhg@sun.ac.za</email>, <bold>Postal address</bold>: Private Bag X1, Matieland 7602, South Africa</corresp>
<fn>
<p><bold>How to cite this article:</bold> Giliomee, J.H., 2015, &#x2018;Dopluise (Hemiptera: Coccoidea) geassosieer met die wipstertmier, <italic>Crematogaster peringueyi</italic> Emery (Hymenoptera: Formicidae)&#x2019;, <italic>Suid-Afrikaanse Tydskrif vir Natuurwetenskap en Tegnologie</italic> 34(1), Art. #1286, 4 pages. <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.4102/satnt.v34i1.1286">http://dx.doi.org/10.4102/satnt.v34i1.1286</ext-link></p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>30</day>
<month>04</month>
<year>2015</year>
</pub-date>
<pub-date pub-type="ppub">
<year>2015</year>
</pub-date>
<volume>34</volume>
<issue>1</issue>
<fpage>1</fpage>
<lpage>4</lpage>
<history>
<date date-type="received">
<day>29</day>
<month>10</month>
<year>2014</year>
</date>
<date date-type="accepted">
<day>26</day>
<month>01</month>
<year>2015</year>
</date>
</history>
<permissions>
<copyright-statement>&#x00A9; 2015. The Authors</copyright-statement>
<copyright-year>2015</copyright-year>
<license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/2.0/">
<license-p>AOSIS OpenJournals. This work is licensed under the Creative Commons Attribution License.</license-p>
</license>
</permissions>
<abstract>
<title>Abstrakte</title>
<p>Neste van die wipstertmier, <italic>Crematogaster peringueyi</italic> (Hymenoptera: Formicidae), is op verskeie plekke langs die kus van die Wes-Kaap versamel. Die doel was om vas te stel watter dopluisagtiges (Hemiptera: Coccoidea) in die neste in assosiasie met hierdie miere leef. Dopluise van drie families, naamlik die Pseudococcidae (witluise), Coccidae (sagtedopluise) en Kerriidae (lakdopluise) is in die neste gevind, almal bekend daarvoor dat hulle heuningdou afskei. Hierdie mutualistiese verhouding tussen die miere en dopluise, bekend as mirmekofilie, is fakultatief van aard. Die wipstertmier blyk ook nie spesifiek te wees wat betref die plant waarop hulle nes maak nie.</p>
</abstract>
<trans-abstract xml:lang="en">
<title>Abstract</title>
<p><bold>The scale insects (Hemiptera: Coccoidea) associated with the cocktail ant (Hymenoptera: Formicidae).</bold> Nests of the cocktail ant, <italic>Crematogaster peringueyi</italic> (Hymenoptera: Formicidae), were collected from various locations along the coast of the Western Cape. The purpose was to determine which scale insects (Hemiptera: Coccoidea) live in association with these ants. Scale insects of three families, that is, Pseudococcidae (mealy bugs), Coccidae (soft scales) and Kerriidae (lac scales) were found in the nests, all of them known for secreting honeydew. This mutualistic association between the ants and scale insects, known as myrmecophily, is facultative, as the cocktail ant and the scale insects were also both found separately. The cocktail ant does not appear to be selective in choosing a plant on which to build a nest.</p>
</trans-abstract>
</article-meta>
</front>
<body>
<sec id="s0001">
<title>Inleiding</title>
<p>Dit is welbekend dat baie mierspesies in mutualistiese verhoudings leef met insekte wat koolhidraatryke stowwe, algemeen bekend as heuningdou, uitskei (Buckley <xref ref-type="bibr" rid="CIT0002">1987a</xref>, <xref ref-type="bibr" rid="CIT0003">1987b</xref>; Gullan <xref ref-type="bibr" rid="CIT0006">1997</xref>). Die verskynsel staan bekend as mirmekofilie (Stadler &#x0026; Dixon <xref ref-type="bibr" rid="CIT0012">2008</xref>). Voorbeelde van sulke insekte is lede van die Coccoidea (dopluisagtiges soos witluise, sagtedopluise en lakdopluise), Membracidae (boomspringers) en Aphididae (plantluise). Die larwes (ruspes) van sekere lede van die skoenlapperfamilie Lycaenidae skei ook &#x2019;n stof af wat vir miere besonder aantreklik is (Clark &#x0026; Dickson <xref ref-type="bibr" rid="CIT0004">1971</xref>; Pierce <italic>et al.</italic> <xref ref-type="bibr" rid="CIT0011">2002</xref>).</p>
<p>Die verhouding tussen miere en hul gashere is gewoonlik fakultatief van aard, dit wil s&#x00EA;, die miere en hul gashere kan wel sonder mekaar klaarkom. Albei trek egter groot voordeel uit die verhouding: die miere kry &#x2019;n konstante en min of meer onbeweeglike bron van voedsel omdat die dop- en plantluise vasgesuig sit aan die plante waarop hulle voed, terwyl die miere hulle teen hul natuurlike vyande beskerm deur afwerende afskeidings en fisieke verwildering. Die verskaffer van voedsel in so &#x2019;n simbiotiese verhouding staan bekend as &#x2019;n trofobiont en die verhouding as trofobiose.</p>
<p>Een van die mierspesies wat simbioties met ruspes en dopluise lewe, is die wipstermier, <italic>Crematogaster peringueyi</italic> Emery (Hymenoptera: Formicidae). Dit kom redelik algemeen in die Wes-Kaap en aangrensende provinsies van Suid-Afrika voor. Die miere is veral volop langs die kus, soos by Vermont (eie waarnemings). Hulle maak kenmerkende groot, ronde neste wat gewoonlik bo die grond in struike aangetref word (<xref ref-type="fig" rid="F0001">Figuur 1</xref>). Die nes word gebou van plantmateriaal wat fyn gekou en gewoonlik kartonagtig van aard is met &#x2019;n doolhof van tonnels aan die binnekant. Benewens die groot nes is daar dikwels kleiner satellietneste op takke naby die moedernes. Hierdie neste kan ook kartonagtig wees (<xref ref-type="fig" rid="F0002">Figuur 2</xref> en <xref ref-type="fig" rid="F0003">3</xref>), maar is soms veselagtig (<xref ref-type="fig" rid="F0004">Figuur 4</xref>) of katoenagtig (<xref ref-type="fig" rid="F0005">Figuur 5</xref> en <xref ref-type="fig" rid="F0006">6</xref>), afhangende van die plantmateriaal wat beskikbaar is. Albei tipes neste kan dien as beskutting vir die ruspes en dopluisssoorte waarmee die miere simbioties verkeer.</p>
<fig id="F0001">
<label>FIGUUR 1</label>
<caption><p>Kartonnnes van <italic>Crematogaster peringueyi</italic> met satellietnes links bo.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="SATNT-34-1286-g001.tif"/>
</fig>
<fig id="F0002">
<label>FIGUUR 2</label>
<caption><p>Satellietnes van <italic>Crematogaster peringueyi</italic> (a) op <italic>Chrysanthemoides incana</italic> met onbedekte, swart <italic>Saissetia</italic>-spesies op linkerkantste takkie te sien en (b) waarin bruin wasdopluise (<italic>Ceroplastes tachardiaformis</italic>) gesien kan word.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="SATNT-34-1286-g002.tif"/>
</fig>
<fig id="F0003">
<label>FIGUUR 3</label>
<caption><p>(a) Satellietnes van <italic>Crematogaster peringueyi</italic> op <italic>Matalasia muricata</italic> en (b) volwasse lakdopluis (<italic>Tachardina minor</italic>) en eerste-instar kruipers (rooi spikkels) in die ontblote nes van <italic>Crematogaster peringueyi</italic>.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="SATNT-34-1286-g003.tif"/>
</fig>
<fig id="F0004">
<label>FIGUUR 4</label>
<caption><p>(a) Satellietnes van <italic>Crematogaster peringueyi</italic> op onbekende plantsoort en (b) onbekende witluissoort met werkers van <italic>Crematogaster peringueyi</italic> binne-in die oopgemaakte satellietnes.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="SATNT-34-1286-g004.tif"/>
</fig>
<fig id="F0005">
<label>FIGUUR 5</label>
<caption><p>(a) Klein satellietnes van <italic>Crematogaster peringueyi</italic>, gemaak van donsagtige materiaal, op <italic>Chrysanthemoides monilifera</italic> en (b) satellietnes van binne met &#x2019;n enkele sagtedopluis van die genus <italic>Saissetia.</italic></p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="SATNT-34-1286-g005.tif"/>
</fig>

<fig id="F0006">
<label>FIGUUR 6</label>
<caption><p>(a) Klein satellietnes van <italic>Crematogaster peringueyi</italic> op <italic>Crysanthemoides monilifera</italic> en (b) die nessie is bo-oor &#x2019;n enkele wasdopluis van die genus <italic>Ceroplastes</italic> gebou.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="SATNT-34-1286-g006.tif"/>
</fig>
<p>By <italic>Chrysoritus</italic>-spesies van die skoenlapperfamilie Lycaenidae, soos <italic>Chrysoritus thysbe</italic> Linnaeus (opaalkopervlerkie), word die ruspes as &#x2018;slawe&#x2019; in die nes aangehou en verlaat hulle net snags die nes om op blare van &#x2019;n gasheerplant soos <italic>Chrysanthemoides monilifera</italic> (bietou) te voed (Clark &#x0026; Dickson <xref ref-type="bibr" rid="CIT0004">1971</xref>; Henning <xref ref-type="bibr" rid="CIT0010">1987</xref>; Pierce <italic>et al.</italic> <xref ref-type="bibr" rid="CIT0011">2002</xref>; Terblanche &#x0026; Van Hamburg <xref ref-type="bibr" rid="CIT0014">2004</xref>; Woodhall <xref ref-type="bibr" rid="CIT0015">2005</xref>). In die geval van die skaars <italic>Chrysanthemoides dicksoni</italic> (Gabriel) (Strandveld-kopervlerkie) verlaat die ruspes nooit die miernes nie, maar word hulle deur die miere gevoed met materiaal wat laasgenoemde opbraak en wat waarskynlik bestaan uit heuningdou wat van dopluise daar naby afkomstig is (Heath &#x0026; Brinkman <xref ref-type="bibr" rid="CIT0009">1995</xref>; Heath <xref ref-type="bibr" rid="CIT0007">1998</xref>; Heath <xref ref-type="bibr" rid="CIT0008">2014</xref>). Die skoenlapperwyfie l&#x00EA; haar eiers juis op struike waar daar wipstertmiere aanwesig is en moontlik ook waar dopluise in die nabyheid is (Edge &#x0026; Terblanche <xref ref-type="bibr" rid="CIT0005">2010</xref>). Die proses van mond-tot-mond-voeding tussen lede van &#x2019;n gemeenskap staan bekend trofallaksis.</p>
<p>Dis nie net suigende insekte en skoenlapperruspes wat mutualisties met wipstertmiere saamlewe en so beskerming kry vir die verskaffing van voedsel nie. In Sentraal- en Oos-Afrika verskaf die sogenaamde &#x2018;whistling thorn&#x2019;-akasia, <italic>Vachellia</italic> (voorheen <italic>Acacia</italic>)<italic> drepanolobium</italic>, voedsel en skuiling aan verskeie soorte <italic>Crematogaster</italic>-miere in ruil vir beskerming teen die groot herbivore soos wildsbokke en selfs olifante (Stapley <xref ref-type="bibr" rid="CIT0013">1998</xref>).</p>
<p>In hierdie artikel word die resultate gerapporteer van &#x2019;n studie wat gedoen is om vas te stel watter dopluisagtiges (Hemiptera: Coccoidea) in die neste van die wipstertmier <italic>C. peringueyi</italic> in Suid-Afrika voorkom.</p>
</sec>
<sec id="s0002">
<title>Materiaal en metodes</title>
<p>Op verskillende plekke langs die kus van die Wes-Kaap is in&#x00A0;die natuurlike veld gesoek na struike met die groot, bolvormige kartonneste van <italic>C. peringueyi</italic>. Naby sulke neste is gewoonlik kleiner satellietneste op dieselfde plant aanwesig. Aangesien die groot neste nie <italic>in situ</italic> oopgesny kon word nie vanwe&#x00EB; die krioelende en bytende miere, en hulle verwoes sou word indien hulle weggeneem sou word, is op die satellietneste gekonsentreer. Hierdie kleiner neste is met &#x2019;n snoeisker afgesny en in dubbele plasieksakke geplaas om te verhoed dat die miere ontsnap. In die laboratorium is die neste vir &#x2019;n paar minute in die vrieshokkie van &#x2019;n yskas geplaas om die miere se bewegings te vertraag, of die miere is doodgemaak deur watte met &#x2019;n paar druppels etielasetaat in die plastieksakke met neste te plaas. Nadat die miere verlam of doodgemaak is, is die neste gefotografeer en dan oopgemaak om die takkies vir dopluise te deursoek. Soms is die takkies waarop die neste was deur middel van &#x2019;n stereomikroskoop ondersoek omdat die voorkoms van sommige dopluisagtiges baie kripties is. Nadat die nes oopgemaak is, is dit weer gefotografeer. Die dopluise wat gevind is, is in 75% etielalkohol gepreserveer en vir identifikasie gestuur na kundige taksonome, wat in bepaalde families spesialiseer (sien Erkenning). Die gasheerplante van die dopluise is vir identifikasie na planttaksonome by die Suid-Afrikaanse Nasionale Instituut vir Biodiversiteit (SANBI), Kirstenbosch Nasionale Botaniese Tuin, Nuweland gestuur. In enkele gevalle waar reproduktiewe strukture van plante nie aanwesig was toe die mierneste versamel is nie, kon die plante nie ge&#x00EF;dentifiseer word nie.</p>
</sec>
<sec id="s0003">
<title>Resultate</title>
<p>&#x2019;n Verskeidenheid van dopluisspesies uit drie families van die superfamilie Coccoidea, naamlik, die Pseudoco&#x00AD;ccidae (witluise), Coccidae (sagtedopluise) en Kerriidae (lakdo&#x00AD;pluise) is in die neste van <italic>C.</italic> <italic>peringueyi</italic> gevind (<xref ref-type="table" rid="T0001">Tabel 1</xref>, <xref ref-type="fig" rid="F0001">Figuur 1&#x2013;6</xref>). Die lede van al drie families is bekend daarvoor dat hulle heuningdou afskei waarop miere voed. Mierneste met dopluise het ook op &#x2019;n verskeidenheid van plantsoorte voorgekom.</p>
<table-wrap id="T0001">
<label>TABEL 1</label>
<caption><p>Dopluisspesies gevind in die neste van die wipstertmier, <italic>Crematogaster peringueyi</italic>, op verskillende plantspesies.</p></caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<td align="left">Spesies</td>
<td align="center">Datum</td>
<td align="center">Dopluisspesies</td>
<td align="center">Plek</td>
<td align="center">Plantspesies</td>
</tr>
</thead>
<tbody>
<tr>
<td align="left"><italic>Pseudococcidae</italic></td>
<td align="left">14.11.13</td>
<td align="left"><italic>Madeurycoccus bicolor</italic>De Lotto</td>
<td align="left">Vermont</td>
<td align="left"><italic>Euclea racemosa</italic></td>
</tr>
<tr>
<td align="left"/>
<td align="left">22.10 13</td>
<td align="left">Onbekende spesies</td>
<td align="left">Vermont</td>
<td align="left">Onbekend</td>
</tr>
<tr>
<td align="left"/>
<td align="left">1998/1999</td>
<td align="left"><italic>Planococcus ficus</italic> (Signoret)<xref ref-type="fn" rid="TFN0001">&#x2020;</xref></td>
<td align="left">Wydverspreid</td>
<td align="left"><italic>Vitis vinifera</italic></td>
</tr>
<tr>
<td align="left"><italic>Coccidae</italic></td>
<td align="left">31.10.12</td>
<td align="left"><italic>Saissetia</italic>-spesies</td>
<td align="left">Saldanha</td>
<td align="left"><italic>Chrysanthemoides incana</italic></td>
</tr>
<tr>
<td align="left"/>
<td align="left">17.1.13</td>
<td align="left"><italic>Saissetia</italic>-spesies</td>
<td align="left">Vermont</td>
<td align="left"><italic>Helichrysum pandurifolium</italic></td>
</tr>
<tr>
<td align="left"/>
<td align="left">31.10.12</td>
<td align="left"><italic>Ceroplastes tachardiaformis</italic> Brain</td>
<td align="left">Saldanha</td>
<td align="left"><italic>Chrysanthemoides incana</italic></td>
</tr>
<tr>
<td align="left"/>
<td align="left">29.7. 13</td>
<td align="left"><italic>Ceroplastes rusticus</italic> De Lotto</td>
<td align="left">Lambertsbaai</td>
<td align="left">Onbekend</td>
</tr>
<tr>
<td align="left"/>
<td align="left">22. 9.13</td>
<td align="left"><italic>Cryptinglisia</italic> sp. nr. <italic>zizyphi</italic></td>
<td align="left">Vermont</td>
<td align="left"><italic>Chrysanthemoides monilifera</italic></td>
</tr>
<tr>
<td align="left"/>
<td align="left">9.1.14</td>
<td align="left"><italic>Coccus longulus</italic> (Douglas)</td>
<td align="left">Hermanus</td>
<td align="left"><italic>Aspalathus-</italic>spesies</td>
</tr>
<tr>
<td align="left"/>
<td align="left">9.1.14</td>
<td align="left"><italic>Ceroplastes</italic> ?<italic>helichrysi</italic> Hall</td>
<td align="left">Hermanus</td>
<td align="left"><italic>Aspalathus-</italic>spesies</td>
</tr>
<tr>
<td align="left"><italic>Kerriidae</italic></td>
<td align="left">31.10.12</td>
<td align="left"><italic>Tachardina minor</italic> (Brain)</td>
<td align="left">Saldanha</td>
<td align="left"><italic>Chrysanthemoides incana</italic></td>
</tr>
<tr>
<td align="left"/>
<td align="left">17.1.13</td>
<td align="left"><italic>Tachardina minor</italic> (Brain)?<xref ref-type="fn" rid="TFN0001">&#x2021;</xref></td>
<td align="left">Vermont</td>
<td align="left"><italic>Metalasia muricata</italic></td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn id="TFN0001"><p>&#x2020;, Volgens Addison en Samways (<xref ref-type="bibr" rid="CIT0001">2000</xref>); &#x2021;, stem nie heeltemal ooreen met die sleutel nie.</p></fn>
</table-wrap-foot></table-wrap>
<p>In sommige gevalle was daar geen dopluise in die neste nie; trouens groot mierneste is waargeneem op dooie takke waarop dopluise natuurlik nie kan voed nie. Groterige neste is ook versamel vanaf lewende bietou en laventel (<italic>Lavandula</italic> <italic>angustifolia</italic>) waarin daar geen dopluise was nie. Dis moontlik &#x2019;n aanduiding dat die miere die dopluise nie in die neste in dra nie, maar nes maak waar hulle reeds aanwesig is. Die dopluisspesies wat gevind is, kom ook buite die mierneste voor.</p>
<p>In een geval (<xref ref-type="fig" rid="F0002">Figuur 2a, b</xref>) is dopluise van die familie Kerriidae in die miernes toegemaak gevind, terwyl ander van die familie Coccidae (wat ook al binne-in neste gevind is) daarnaas onbedek was.</p>
<p>In die gebied waar die ondersoek gedoen is, is geen skoenlapperlarwes in die mierneste gevind nie, alhoewel bietou daar voorkom en enkele volwassenes van <italic>C. thysbe</italic> by Vermont waargeneem is.</p>
</sec>
<sec id="s0004">
<title>Bespreking</title>
<p>Dit blyk dat <italic>C. peringyueyi</italic> nie baie spesifiek is oor die soort plant waarop hulle neste bou nie en in enkele gevalle is neste op die grond teenaan struike gevind. Selfs as die tak doodgaan waarop die nes gebou is, sal die nes bly voortbestaan. Aanvanklik is so &#x2019;n nes waarskynlik rondom dopluise gebou, maar as die tak van nature doodgaan, soos dikwels met die ouer takke van bietou gebeur (eie waarnemings), en die dopluise daarmee saam, sal die miere sonder die dopluise klaarkom. Laasgenoemde blyk ook uit die feit dat mierneste (grotes sowel as satellietneste) sonder dopluise soms op groeiende plante gevind word. Die miere is ook nie kieskeurig wat betref die dopluise waarmee hulle saamleef nie, solank hulle net heuningdou afskei. Die dopluise daarenteen geniet beskerming in die mierneste, maar is nie tot die neste beperk nie sodat die verhouding tussen wipstertmiere en die dopluisspesies vir beide partye fakultatief van aard is.</p>
<p>Die feit dat nuwe soorte dopluisagtiges telkens gevind is soos die ondersoek gevorder het, dui daarop dat baie ander spesies van die drie families waarskynlik nog in neste van <italic>C. peringueyi</italic> aangetref kan word. Verdere ondersoek van neste kan selfs lei tot die ontdekking van onbekende dopluisspesies, want miere is vernuftige soekers na kos en sal enige spesies wat heuningdou afskei, opspoor.</p>
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<title>Erkenning</title>
<p>Ian Millar en Chris Hodgson word bedank vir die identifikasie van die dopluisagtiges en onbekende taksonome van die Suid-Afrikaanse Nasionale Instituut vir Biodiversiteit vir die identifikasie van die plante.</p>
<sec id="s20006">
<title>Mededingende belange</title>
<p>Die outeur verklaar hiermee dat hy geen finansi&#x00EB;le of persoonlike verbintenis het met enige party wat hom nadelig of voordelig kon be&#x00EF;nvloed het in die skryf van hierdie artikel nie.</p>
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