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<article xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" article-type="research-article" xml:lang="en">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">KOEDOE</journal-id>
<journal-title-group>
<journal-title>KOEDOE - African Protected Area Conservation and Science</journal-title>
</journal-title-group>
<issn pub-type="ppub">0075-6458</issn>
<issn pub-type="epub">2071-0771</issn>
<publisher>
<publisher-name>AOSIS</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">KOEDOE-61-1531</article-id>
<article-id pub-id-type="doi">10.4102/koedoe.v61i1.1531</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Checklist</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>A checklist of the termites of Kruger National Park, South Africa</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-5306-9963</contrib-id>
<name>
<surname>Lind</surname>
<given-names>Brianna M.</given-names>
</name>
<xref ref-type="aff" rid="AF0001">1</xref>
</contrib>
<contrib contrib-type="author">
<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-0003-1435</contrib-id>
<name>
<surname>Davies</surname>
<given-names>Andrew B.</given-names>
</name>
<xref ref-type="aff" rid="AF0002">2</xref>
</contrib>
<aff id="AF0001"><label>1</label>Plant and Environmental Sciences Department, New Mexico State University, New Mexico, United States</aff>
<aff id="AF0002"><label>2</label>Department of Global Ecology, Carnegie Institution for Science, California, United States</aff>
</contrib-group>
<author-notes>
<corresp id="cor1"><bold>Corresponding author:</bold> Brianna M. Lind, <email xlink:href="brianna.m.lind@gmail.com">brianna.m.lind@gmail.com</email></corresp>
</author-notes>
<pub-date pub-type="epub"><day>09</day><month>04</month><year>2019</year></pub-date>
<pub-date pub-type="collection"><year>2019</year></pub-date>
<volume>61</volume>
<issue>1</issue>
<elocation-id>1531</elocation-id>
<history>
<date date-type="received"><day>14</day><month>03</month><year>2018</year></date>
<date date-type="accepted"><day>12</day><month>11</month><year>2018</year></date>
</history>
<permissions>
<copyright-statement>&#x00A9; 2019. The Authors</copyright-statement>
<copyright-year>2019</copyright-year>
<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by/4.0/">
<license-p>Licensee: AOSIS. This work is licensed under the Creative Commons Attribution License.</license-p>
</license>
</permissions>
<abstract>
<p>The protection of biodiversity is critical to ecosystem function and is a primary management goal for conservation areas globally. Maintaining a current inventory of known diversity is a central component of achieving this goal and serves as an essential starting point for future research endeavours. Since the first published survey of termites in South Africa&#x2019;s Kruger National Park (KNP) over 55 years ago, our understanding of termite diversity has expanded sufficiently to merit an update and formal checklist. Here we revise the inventory of termite diversity in KNP and summarise the taxonomic and functional diversity of termites in the park. A thorough review of recent termite research in KNP added 6 new genera and 13 species to what was found in Coaton&#x2019;s original survey, with one genus, <italic>Anenteotermes</italic>, recorded for the first time in southern Africa. Based on the updated species checklist, the majority of genera in the park belong to Feeding Group II (39&#x0025;) and the Termitidae family (75&#x0025;).</p>
<p><bold>Conservation implications</bold>: In savannas, termites play crucial roles in nutrient cycling, water redistribution and plant dynamics. Systematically cataloguing termite diversity and assemblage composition in the park provides an essential baseline for scientific research, aids biodiversity conservation efforts and encourages scientists and managers to consider termites in ecosystem functioning and management. Having more detailed descriptions of genera, species and feeding groups allows for more tangible, ecologically relevant attributions of termite influence, facilitates enhanced inquiry and allows for more realistic quantification of termite roles in key ecosystem processes.</p>
<p><bold>Keywords:</bold> termite; Kruger National Park; feeding groups; diversity; savanna.</p>
</abstract>
</article-meta>
</front>
<body>
<sec id="s0001">
<title>Introduction</title>
<p>A principal goal of most conservation areas is to conserve biodiversity in all its facets. To achieve this goal, scientists must first catalogue the known diversity of organisms to serve as a baseline for future conservation and research endeavours. This is particularly pertinent for groups that perform important ecological functions, such as termites. Termites are a prominent source of habitat heterogeneity in African savannas and have a substantial impact on ecosystem functioning (Jouquet et al. <xref ref-type="bibr" rid="CIT0022">2011</xref>; Sileshi et al. <xref ref-type="bibr" rid="CIT0032">2010</xref>). Termites are the dominant invertebrate decomposers in tropical forests and savannas (Collins <xref ref-type="bibr" rid="CIT0005">1981</xref>; Schuurman <xref ref-type="bibr" rid="CIT0031">2005</xref>), and, through their foraging activities, alter soil processes, redistribute nutrients and modify soil hydrology (Jouquet et al. <xref ref-type="bibr" rid="CIT0022">2011</xref>). These ecosystem alterations in turn affect vegetation distributions and assemblage composition (Davies, Baldeck &#x0026; Asner <xref ref-type="bibr" rid="CIT0006">2016a</xref>; Joseph et al. <xref ref-type="bibr" rid="CIT0021">2014</xref>; Sileshi et al. <xref ref-type="bibr" rid="CIT0032">2010</xref>), resulting in a patchy landscape that alters both mammalian and insect herbivore foraging behaviour (Davies et al. <xref ref-type="bibr" rid="CIT0011">2016b</xref>, <xref ref-type="bibr" rid="CIT0013">2016c</xref>; Grant &#x0026; Scholes <xref ref-type="bibr" rid="CIT0019">2006</xref>; Levick et al. <xref ref-type="bibr" rid="CIT0024">2010a</xref>).</p>
<p>Considering its latitude, southern Africa has a surprisingly high diversity of termites, with over 50 genera present (Eggleton <xref ref-type="bibr" rid="CIT0017">2000</xref>; Uys <xref ref-type="bibr" rid="CIT0034">2002</xref>). The termite fauna of the region is generally considered to be well documented as a result of the National Survey of Isoptera conducted by W.G.H. Coaton and co-workers between 1956 and 1979. During this survey, almost every quarter-degree grid cell of South Africa, Swaziland, Zimbabwe and Namibia was sampled for termites. Much of the information gained from this survey appeared in a series of publications in <italic>Cimbebasia</italic> between 1971 and 1980 (Uys <xref ref-type="bibr" rid="CIT0034">2002</xref>), and collected specimens are currently located in the South African National Collection of Insects (Uys <xref ref-type="bibr" rid="CIT0034">2002</xref>). Kruger National Park (KNP), South Africa, was also included in these surveys, with a checklist of the park&#x2019;s termite fauna published in 1962 (Coaton <xref ref-type="bibr" rid="CIT0003">1962</xref>). However, since this time, a considerable amount of termite research has been conducted in KNP, warranting an update of the checklist in order to have a comprehensive and current inventory of termite diversity in the park.</p>
<p>In his survey, Coaton collected 22 termite genera from four families within KNP (Coaton <xref ref-type="bibr" rid="CIT0003">1962</xref>). While the last few decades have seen a large increase in the number of studies investigating termite ecology in KNP, none have focused exclusively on termite taxonomy or sought to update the checklist, despite new genus and species records. Here we synthesise the termite occurrence records from published studies in KNP and compile an updated list of termite richness and functional diversity to facilitate better understanding of KNP&#x2019;s biodiversity, which will aid both management and future research endeavours. By characterising the park&#x2019;s termite assemblages, a fuller understanding of the fundamental roles termites play in critical ecosystem processes within Kruger&#x2019;s savannas can also be realised.</p>
</sec>
<sec id="s0002">
<title>Methods and materials</title>
<p>Kruger National Park is located in low-lying savanna in the northeast corner of South Africa between the foot slopes of the Drakensberg Escarpment and Mozambican coastal plains (Venter, Scholes &#x0026; Eckhardt et al. <xref ref-type="bibr" rid="CIT0035">2003</xref>). The region experiences high temperatures in the summer, mild winters and the majority of rainfall between October and April (ranging from ~300 mm/year in the north to ~700 mm/year in the south) (Venter et al. <xref ref-type="bibr" rid="CIT0035">2003</xref>). The park has two main underlying geologies, basalt in the east and granite in the west (Venter et al. <xref ref-type="bibr" rid="CIT0035">2003</xref>). Along with varying rainfall, these geologies contribute to a diverse range of soil types and influence vegetation patterns (Venter et al. <xref ref-type="bibr" rid="CIT0035">2003</xref>). Tree canopy cover ranges from 5&#x0025; to 60&#x0025;, and 75&#x0025; of the park is covered by broad-leaved savanna with the remaining 25&#x0025; composed of fine-leaved savanna (Venter et al. <xref ref-type="bibr" rid="CIT0035">2003</xref>).</p>
<p>We conducted a comprehensive literature review of research involving termites in KNP by searching Web of Science and Google Scholar and compiled a list of termite genera and species sampled from each study. Termites were listed by family, subfamily, genus and species and categorised into feeding groups (FG) following Donovan, Eggleton and Bignell (<xref ref-type="bibr" rid="CIT0015">2001</xref>) to characterise the assemblage. Feeding groups consist of four categories (I&#x2013;IV) based on the level of decay of food items, which ranged from live plants to decayed organic material. Donovan et al. (<xref ref-type="bibr" rid="CIT0015">2001</xref>) defines FGs to include a non-termitid group, FG-I, that feeds on dead wood and grass and three termitid groups that eat dead wood, litter and micro-epiphytes, FG-II, humus, FG-III, and soil minerals, FG-IV.</p>
</sec>
<sec id="s0003">
<title>Results</title>
<p>Our literature review of 20 termite-related studies conducted in KNP (see <xref ref-type="table" rid="T0001">Table 1</xref>) revealed that since Coaton&#x2019;s 1962 survey, six previously unlisted termite genera and 13 species have been sampled in the park (see <xref ref-type="table" rid="T0002">Table 2</xref>). In addition, one sampled genus, <italic>Anenteotermes</italic>, had not been previously recorded in southern Africa. Three studies in particular contributed to checklist updates: Meyer et al. (<xref ref-type="bibr" rid="CIT0026">1999</xref>); Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>); and Davies et al. (<xref ref-type="bibr" rid="CIT0010">2014a</xref>). While termites from all southern African families are present in the park, the majority (75&#x0025;) belong to the Termitidae family (see <xref ref-type="fig" rid="F0001">Figure 1a</xref>). In terms of FGs, FG-II makes up 40&#x0025; of termite genera in the park, FG-I and FG-III each make up 25&#x0025; and FG-IV, which is the least represented, makes up 10&#x0025; (see <xref ref-type="fig" rid="F0001">Figure 1b</xref>).</p>
<fig id="F0001">
<label>FIGURE 1</label>
<caption><p>Summary of the families (a) and feeding groups (b) of termites sampled in Kruger National Park, South Africa.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="KOEDOE-61-1531-g001.tif"/>
</fig>
<table-wrap id="T0001">
<label>TABLE 1</label>
<caption><p>History of published, termite-focused research conducted in the Kruger National Park, South Africa.</p></caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th valign="top" align="left">Year of Publication</th>
<th valign="top" align="left">Authors</th>
<th valign="top" align="left">Title</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">1962</td>
<td align="left">Coaton, W.G.H.</td>
<td align="left">Survey of the termites of Kruger National Park</td>
</tr>
<tr>
<td align="left">1995</td>
<td align="left">Braack, L.E.O.</td>
<td align="left">Seasonal activity of savanna termites during and after a severe drought</td>
</tr>
<tr>
<td align="left">1999</td>
<td align="left">Meyer, V.W. et al.</td>
<td align="left">Distribution and density of termite mounds in the northern Kruger National Park, with specific reference to those constructed by Macrotermes Holmgren (Isoptera: Termitidae)</td>
</tr>
<tr>
<td align="left">2000</td>
<td align="left">Meyer, V.W. et al.</td>
<td align="left">Distribution and density of <italic>Cubitermes</italic> Wasmann (Isoptera: Termitidae) mounds in the northern Kruger National Park</td>
</tr>
<tr>
<td align="left">2000</td>
<td align="left">Meyer, V.W. et al.</td>
<td align="left">Intracolonial demography of the mound-building termite <italic>Macrotermes natalensis</italic> (Haviland) (Isoptera, Termitidae) in the northern Kruger National Park, South Africa</td>
</tr>
<tr>
<td align="left">2001</td>
<td align="left">Meyer, V.W. et al.</td>
<td align="left">Biomass of <italic>Macrotermes natalensis</italic> in the northern Kruger National Park, South Africa &#x2013; the effects of land characteristics</td>
</tr>
<tr>
<td align="left">2003</td>
<td align="left">Meyer, V.W. et al.</td>
<td align="left">Estimates of food consumption by the fungus-growing termite <italic>Macrotermes natalensis</italic> in a South African savanna-woodland: news and views</td>
</tr>
<tr>
<td align="left">2005</td>
<td align="left">Sponheimer et al.</td>
<td align="left">Hominins, sedges, and termites: new carbon isotope data from the Sterkfontein valley and Kruger National Park</td>
</tr>
<tr>
<td align="left">2006</td>
<td align="left">Grant &#x0026; Scholes</td>
<td align="left">The importance of nutrient hot-spots in the conservation and management of large wild mammalian herbivores in semi-arid savannas</td>
</tr>
<tr>
<td align="left">2010</td>
<td align="left">Levick et al.</td>
<td align="left">Regional insight into savanna hydrogeomorphology from termite mounds</td>
</tr>
<tr>
<td align="left">2010</td>
<td align="left">Levick et al.</td>
<td align="left">The spatial extent of termite influences on herbivore browsing in an African savanna</td>
</tr>
<tr>
<td align="left">2012</td>
<td align="left">Coetsee et al.</td>
<td align="left">An overview of nitrogen cycling in a semi-arid savanna: some implications for management and conservation in a large African park</td>
</tr>
<tr>
<td align="left">2012</td>
<td align="left">Davies et al.</td>
<td align="left">The pyrodiversity-biodiversity hypothesis: a test with savanna termite assemblages</td>
</tr>
<tr>
<td align="left">2013</td>
<td align="left">Davies et al.</td>
<td align="left">Assessing the relative efficiency of termite sampling methods along a rainfall gradient in African savannas</td>
</tr>
<tr>
<td align="left">2014</td>
<td align="left">Davies et al.</td>
<td align="left">Variable effects of termite mounds on African savanna grass communities across a rainfall gradient</td>
</tr>
<tr>
<td align="left">2014</td>
<td align="left">Davies et al.</td>
<td align="left">Spatial variability and abiotic determinants of termite mounds throughout a savanna catchment</td>
</tr>
<tr>
<td align="left">2015</td>
<td align="left">Davies et al.</td>
<td align="left">Seasonal activity patterns of African savanna termites vary across a rainfall gradient</td>
</tr>
<tr>
<td align="left">2016</td>
<td align="left">Davies et al.</td>
<td align="left">Termite mounds alter the spatial distribution of African savanna tree species</td>
</tr>
<tr>
<td align="left">2016</td>
<td align="left">Davies et al.</td>
<td align="left">Termite mounds differ in their importance for herbivores across savanna types, seasons and spatial scales</td>
</tr>
<tr>
<td align="left">2016</td>
<td align="left">Joseph et al.</td>
<td align="left">Microclimates mitigate against hot temperatures in dryland ecosystems: termite mounds as an example</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T0002">
<label>TABLE 2</label>
<caption><p>Checklist of termites known to occur in the Kruger National Park, South Africa.</p></caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th valign="top" align="left">Family</th>
<th valign="top" align="left">Subfamily</th>
<th valign="top" align="left">Genus</th>
<th valign="top" align="left">Species</th>
<th valign="top" align="center">Original Coaton (<xref ref-type="bibr" rid="CIT0003">1962</xref>)</th>
<th valign="top" align="center">Added by Meyer (<xref ref-type="bibr" rid="CIT0026">1999</xref>)</th>
<th valign="top" align="center">Added by Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>)</th>
<th valign="top" align="center">FG</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Kalotermitidae</td>
<td align="left">-</td>
<td align="left"><italic>Cryptotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Epicalotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Bifiditermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Neotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left">Hodotermitidae</td>
<td align="left">-</td>
<td align="left"><italic>Hodotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>mossambicus</italic></td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left">Rhinotermitidae</td>
<td align="left">-</td>
<td align="left"><italic>Psammotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Schedorhinotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">I</td>
</tr>
<tr>
<td align="left">Termitidae</td>
<td align="left">Macrotermitinae</td>
<td align="left"><italic>Allodontermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>rhodesiensis</italic></td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Ancistrotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>latinotus</italic></td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Macrotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>falciger</italic></td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>michaelseni</italic></td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>natalensis</italic></td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>ukuzii</italic></td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Microtermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Odontotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>prob. Latericius</italic></td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left">Apicotermitinae</td>
<td align="left"><italic>Adaiphrotermes</italic></td>
<td align="left">spp.</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">III</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Alyscotermes</italic></td>
<td align="left">spp.</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">III</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Anenteotermes</italic></td>
<td align="left">spp.</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">III</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Apicotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">IV</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Astalotermes</italic></td>
<td align="left">spp.</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">III</td>
</tr>
<tr>
<td align="left"></td>
<td align="left">Termitinae</td>
<td align="left"><italic>Amitermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>hastatus</italic></td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>messinae</italic></td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Anguilitermes</italic></td>
<td align="left">spp.</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">III</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Anoplotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">III</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Cubitermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">IV</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Lepidotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">IV</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Microcerotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Promirotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">III</td>
</tr>
<tr>
<td align="left"></td>
<td align="left">Nausitermitinae</td>
<td align="left"><italic>Fulleritermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Coarctotermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Rhadinotermes</italic></td>
<td align="left">spp.</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>coarctus</italic></td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>Trinervitermes</italic></td>
<td align="left">spp.</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>dispar</italic></td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
<tr>
<td align="left"></td>
<td align="left"></td>
<td align="left"></td>
<td align="left"><italic>trinervoides</italic></td>
<td align="center">-</td>
<td align="center">x</td>
<td align="center">-</td>
<td align="center">II</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn><p>No. Genera, Original &#x2013; Coaton (<xref ref-type="bibr" rid="CIT0003">1962</xref>) = 22; Added by Meyer (<xref ref-type="bibr" rid="CIT0026">1999</xref>) = 0; Added by Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>) = 6.</p></fn>
<fn><p>No. Species, Original &#x2013; Coaton (<xref ref-type="bibr" rid="CIT0003">1962</xref>) = 0; Added by Meyer (<xref ref-type="bibr" rid="CIT0026">1999</xref>) = 5; Added by Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>) = 6.</p></fn>
<fn><p>FG, feeding group.</p></fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s0004">
<title>Discussion</title>
<p>Termite ecology in KNP, and in savannas globally, has advanced substantively since Coaton&#x2019;s 1962 survey. Our literature review revealed 6 previously unrecorded genera and 13 previously unrecorded species in KNP. Termite diversity in KNP appears to mirror the generalised diversity pattern of termites in southern Africa, with relatively few genera in the Hodotermitidae, Kalotermitidae and Rhinotermitidae families, with the majority of termite diversity being found within the Termitidae family (Uys <xref ref-type="bibr" rid="CIT0034">2002</xref>). As knowledge about how termites affect the structure and function of savanna ecosystems continues to accumulate, it will be increasingly important to understand how different species and genera interact with and affect the environment. Maintaining an accurate record of the termites that occur in KNP is a critical first step to achieving this understanding and provides an essential baseline for ongoing research.</p>
<p>There were two major research papers that contributed most significantly to updating the inventory of KNP&#x2019;s taxonomic diversity: Meyer et al. (<xref ref-type="bibr" rid="CIT0026">1999</xref>), and Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>). Meyer et al. (<xref ref-type="bibr" rid="CIT0026">1999</xref>) made substantial contributions to advancing our understanding of termite diversity in KNP through research that involved mapping the distribution and density of termite mounds in northern Kruger (Meyer et al. <xref ref-type="bibr" rid="CIT0026">1999</xref>, <xref ref-type="bibr" rid="CIT0025">2000a</xref>). Meyer et al. (<xref ref-type="bibr" rid="CIT0026">1999</xref>) documented <italic>Macrotermes natalensis</italic> as the most abundant mound-building termite species in this region, which led to several follow-up studies on the species, including an investigation of intracolonial demography (Meyer et al. <xref ref-type="bibr" rid="CIT0025">2000</xref>), biomass (Meyer et al. <xref ref-type="bibr" rid="CIT0029">2001</xref>) and food consumption (Meyer et al. <xref ref-type="bibr" rid="CIT0027">2003</xref>). While Meyer&#x2019;s research into <italic>M. natalensis</italic> provides an excellent baseline for the ecological influence of this termite species in northern KNP, future research should strive to account for the full breath of foraging power (i.e. litter removal) that all FG-II termites effect in KNP. When FG-II termites are evaluated collectively, we may find that litter removal by termites competes with burning and herbivory and could in turn subsequently influence fire regimes and patterns.</p>
<p>While Meyer et al. (<xref ref-type="bibr" rid="CIT0026">1999</xref>) increased our knowledge of mound-building termites, Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>) employed a broad range of sampling strategies that captured the full range of termite families in the park. Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>) sampled termites across varying burning regimes across KNP and found that termites are broadly resilient to fire. They found that FG-II termites are dominant across KNP&#x2019;s rainfall gradient with the caveat that the proportion of other FGs changed with rainfall, with the greatest FG diversity in the wettest region, Pretoriuskop. This overall finding is similar to our synthesis, where we also found a dominance of FG-II termites in KNP. Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>) also documented a first record of the soil-feeding genus <italic>Anenteotermes</italic> (Sands <xref ref-type="bibr" rid="CIT0030">1998</xref>) in KNP and in southern Africa. Although this genus is known to range from semi-arid savanna to rainforests (Sands <xref ref-type="bibr" rid="CIT0030">1998</xref>), Davies et al. (<xref ref-type="bibr" rid="CIT0007">2012</xref>) were the first to document the genus in southern Africa. <italic>Anenteotermes</italic> has previously been recorded from as far south as Malawi (Donovan et al. <xref ref-type="bibr" rid="CIT0016">2002</xref>) and is also well represented in central (e.g. Deblauwe et al. <xref ref-type="bibr" rid="CIT0014">2008</xref>; Eggleton et al. <xref ref-type="bibr" rid="CIT0018">1996</xref>) and east (e.g. Wanyonyi et al. <xref ref-type="bibr" rid="CIT0036">1984</xref>) Africa.</p>
<p>Although we have added numerous additions to Coaton&#x2019;s original 1962 checklist, many of the specimens he sampled still await identification. As such, our &#x2018;new&#x2019; findings compiled from recent research may be a consequence of a more comprehensive identification process rather than of sampling in a new environment. This highlights the lack of taxonomic resolution of termites (especially non-mound-building, soil-feeding termites) in Africa. Indeed, almost a third of termite genera in southern Africa are in need of taxonomic revision (Uys <xref ref-type="bibr" rid="CIT0034">2002</xref>). Because listing and categorising taxa is such an important component of conserving biodiversity and understanding ecosystems, a priority of future termite research should be to resolve these taxonomic difficulties (Wilson <xref ref-type="bibr" rid="CIT0037">2017</xref>). Deo-xyribonucleic acid metabarcoding, in conjunction with taxonomic and natural history data components, may facilitate enhanced taxonomic clarity and enable more species-level identifications.</p>
</sec>
<sec id="s0005">
<title>Conclusion</title>
<p>Collectively, termites as an insect group play considerably important roles in the functioning of dryland and subtropical environments. Maintaining a termite checklist in KNP allows us to begin the important work of attributing what we understand as &#x2018;termite influence&#x2019; to specific termite species and genera. It also provides an invaluable baseline for future research in the park and facilitates the implementation of biodiversity conservation, a key park management goal. The ecological significance and conservation value of insects in savannas has traditionally received little attention relative to vertebrates (Braack &#x0026; Kryger <xref ref-type="bibr" rid="CIT0002">2003</xref>). Knowledge of termite diversity and assemblage composition will hopefully help bridge this gap and facilitate the consideration of this important insect group in ecosystem functioning and management planning. Moving forward, assessing termite diversity in ecologically relevant FGs, that have targeted and discernible ecosystem impacts, will advance research endeavours and allow for more realistic quantification of their roles in key processes such as vegetation dynamics, nutrient cycling, litter removal and fuel-load management.</p>
</sec>
</body>
<back>
<ack>
<title>Acknowledgements</title>
<p>The authors would like to thank Dr Niall Hanan for insightful comments on an earlier version of the manuscript.</p>
<sec id="s20006" sec-type="COI-statement">
<title>Competing interests</title>
<p>The authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article.</p>
</sec>
<sec id="s20007">
<title>Authors&#x2019; contributions</title>
<p>B.M.L. and A.B.D. designed the study, analysed the data and wrote the article.</p>
</sec>
</ack>
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<fn><p><bold>How to cite this article:</bold> Lind, B.M. &#x0026; Davies, A.B., 2019, &#x2018;A checklist of the termites of Kruger National Park, South Africa&#x2019;, <italic>Koedoe</italic> 61(1), a1531. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.4102/koedoe.v61i1.1531">https://doi.org/10.4102/koedoe.v61i1.1531</ext-link></p></fn>
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