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<article xmlns:tp="http://www.plazi.org/taxpub" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" 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">5</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:8D1BC1DD-8175-5933-B147-C839B202D5BA</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:C132FA7C-A22D-4474-AAAF-B7A6889D0FF9</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">NeoBiota</journal-title>
        <abbrev-journal-title xml:lang="en">NB</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1619-0033</issn>
      <issn pub-type="epub">1314-2488</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/neobiota.109.191014</article-id>
      <article-id pub-id-type="publisher-id">191014</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Arthropoda</subject>
          <subject>Hymenoptera</subject>
          <subject>Insecta</subject>
          <subject>Vespidae</subject>
          <subject>Vespoidea</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biological Invasions</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Cenozoic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Africa</subject>
          <subject>South Africa</subject>
          <subject> Lesotho and Swaziland</subject>
          <subject>Southern Africa</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Potential future economic impact of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> (Fabricius) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Hymenoptera">Hymenoptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Vespidae">Vespidae</tp:taxon-name-part></tp:taxon-name>) in South Africa</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Cook</surname>
            <given-names>David C.</given-names>
          </name>
          <email xlink:type="simple">david.cook2@dpird.wa.gov.au</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-2954-7697</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/visualization/">Visualization</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>McKirdy</surname>
            <given-names>Simon J.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-6578-5521</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Veldtman</surname>
            <given-names>Ruan</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-2258-6108</uri>
          <xref ref-type="aff" rid="A4">4</xref>
          <xref ref-type="aff" rid="A5">5</xref>
          <xref ref-type="aff" rid="A6">6</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/validation/">Validation</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Harry Butler Institute, Murdoch University, 90 South Street, Murdoch WA 6150, Australia</addr-line>
        <institution>Harry Butler Institute, Murdoch University</institution>
        <addr-line content-type="city">Murdoch</addr-line>
        <country>Australia</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">School of Agriculture and Environment, The University of Western Australia, 35 Stirling Highway, Perth WA 6009, Australia</addr-line>
        <institution>School of Agriculture and Environment, The University of Western Australia</institution>
        <addr-line content-type="city">Perth</addr-line>
        <country>Australia</country>
        <uri content-type="ror">https://ror.org/047272k79</uri>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Biosecurity and Emergency Response, Department of Primary Industries and Regional Development, PO Box 1231, Bunbury WA 6231, Australia</addr-line>
        <institution>Biosecurity and Emergency Response, Department of Primary Industries and Regional Development</institution>
        <addr-line content-type="city">Bunbury</addr-line>
        <country>Australia</country>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">South African National Biodiversity Institute, Kirstenbosch NBG, Rhodes Drive, Newlands, Cape Town, South Africa</addr-line>
        <institution>South African National Biodiversity Institute</institution>
        <addr-line content-type="city">Cape Town</addr-line>
        <country>South Africa</country>
      </aff>
      <aff id="A5">
        <label>5</label>
        <addr-line content-type="verbatim">Department of Conservation Ecology and Entomology, Stellenbosch University, Private Bag x1, Matieland 7602, Stellenbosch, South Africa</addr-line>
        <institution>Department of Conservation Ecology and Entomology, Stellenbosch University</institution>
        <addr-line content-type="city">Stellenbosch</addr-line>
        <country>South Africa</country>
        <uri content-type="ror">https://ror.org/05bk57929</uri>
      </aff>
      <aff id="A6">
        <label>6</label>
        <addr-line content-type="verbatim">National Institute for Theoretical and Computational Science, Private Bag x1, Matieland 7602, Stellenbosch, South Africa</addr-line>
        <institution>National Institute for Theoretical and Computational Science</institution>
        <addr-line content-type="city">Stellenbosch</addr-line>
        <country>South Africa</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: David C. Cook (<email xlink:type="simple">david.cook2@dpird.wa.gov.au</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Andrew (Sandy) Liebhold</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>16</day>
        <month>09</month>
        <year>2026</year>
      </pub-date>
      <volume>109</volume>
      <fpage>67</fpage>
      <lpage>84</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/3E571153-2F13-53D3-87B4-D19B70373A3A">3E571153-2F13-53D3-87B4-D19B70373A3A</uri>
      <history>
        <date date-type="received">
          <day>09</day>
          <month>03</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>16</day>
          <month>06</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>David C. Cook, Simon J. McKirdy, Ruan Veldtman</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <abstract>
        <label>Abstract</label>
        <p>The potential economic effects of the introduced German wasp, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> (Fabricius), on fruit and vegetable industries and households in South Africa were estimated over several decades if it continues to spread. The model described was used to simulate impacts on horticulture and wine grape production and household pest control services. The results suggest that, if unmanaged, the value of damage caused by this important pest could amount to US$160–300 million over the next 50 years. This implies that the investment ceiling for a prospective eradication program is approximately US$3–6 million per year. The results were highly sensitive to the rate of spread, indicating high potential returns on investments in spread-mitigation activities if eradication is not technically feasible.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>Benefit-cost analysis</kwd>
        <kwd>economic impact assessment</kwd>
        <kwd>European wasp</kwd>
        <kwd>German wasp</kwd>
        <kwd>
          <italic>
            <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name>
          </italic>
        </kwd>
        <kwd>yellowjacket</kwd>
      </kwd-group>
      <funding-group>
        <award-group>
          <funding-source>
            <named-content content-type="funder_name">Murdoch University</named-content>
            <named-content content-type="funder_identifier">501100001799</named-content>
            <named-content content-type="funder_ror">https://ror.org/00r4sry34</named-content>
            <named-content content-type="funder_doi">http://doi.org/10.13039/501100001799</named-content>
          </funding-source>
        </award-group>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p>The German wasp, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> (Fabricius, 1793) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Hymenoptera">Hymenoptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Vespidae">Vespidae</tp:taxon-name-part></tp:taxon-name>), is an agricultural, environmental, and social pest that has been introduced to the Southern Hemisphere on various occasions over the last century (<xref ref-type="bibr" rid="B37">Lester and Beggs 2019</xref>; <xref ref-type="bibr" rid="B76">Wilson Rankin 2021</xref>). The increased movement of people and cargo after World War II saw <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> spread from its native range in Europe and the Mediterranean region to North America, Argentina, Chile, Iceland, South Africa, New Zealand, and Australia, where it has become invasive (<xref ref-type="bibr" rid="B29">Gergonne et al. 2026</xref>). It was first reported in the Western Cape region of South Africa in 1972, although it could have been present before 1970 (<xref ref-type="bibr" rid="B74">Whitehead 1975</xref>; <xref ref-type="bibr" rid="B75">Whitehead and Prins 1975</xref>). It is currently listed as a category 1B invasive species in South Africa, which necessitates a management plan, but eradication is probably feasible given the relatively slow spread of the species until recently (<xref ref-type="bibr" rid="B68">van Zyl et al. 2018</xref>; <xref ref-type="bibr" rid="B71">Veldtman et al. 2021</xref>). This slow spread rate is unusual, since all reports of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in South Africa are surrounded by climatically optimal and highly suitable habitat (<xref ref-type="bibr" rid="B20">de Villiers et al. 2017</xref>).</p>
      <p>It is not known how <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> first arrived in the Western Cape region, but because only two haplotypes have been identified, denoted Hap1 and Hap2, it may be that two incursion events led to successful establishment (<xref ref-type="bibr" rid="B67">van Asch et al. 2022</xref>; <xref ref-type="bibr" rid="B29">Gergonne et al. 2026</xref>). Hap2 was most likely involved in the first incursion, since it is the dominant haplotype around the original invaded range in and around the Cape Peninsula, whereas Hap1 appears to have arrived in the Boland region northeast of Cape Town in 2000 and subsequently spread to the north, east, and south (<xref ref-type="bibr" rid="B19">Davies et al. 2020</xref>; <xref ref-type="bibr" rid="B71">Veldtman et al. 2021</xref>) (Fig. <xref ref-type="fig" rid="F1">1</xref>). It is therefore hypothesized that Hap1 is more climatically suited to the Western Cape region than Hap2 (<xref ref-type="bibr" rid="B67">van Asch et al. 2022</xref>).</p>
      <fig id="F1">
        <object-id content-type="arpha">956C4674-21DD-52D9-85F3-A442A7812B90</object-id>
        <label>Figure 1.</label>
        <caption>
          <p>Distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in South Africa. Currently, the wasp is only found in the Western Cape region. The location of nest sites recorded between 2012 and 2026 is shown in red.</p>
        </caption>
        <graphic xlink:href="neobiota-109-067_article-191014__-g001.jpg" id="oo_1773301.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1773301</uri>
        </graphic>
      </fig>
      <p>Given its accelerated spread in the past 20 years, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> could cause substantial economic costs to apiculture and fruit and vegetable industries. Bees from both wild and managed European honeybee (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Apis">Apis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mellifera">mellifera</tp:taxon-name-part></tp:taxon-name></italic> Linnaeus) colonies are attacked by the wasp, sometimes by direct raids on beehives but more frequently by attacks on individual adult bees at ground level (<xref ref-type="bibr" rid="B52">Pusceddu et al. 2017</xref>). The resultant decline in free and commercial pollination services is particularly serious in the case of South Africa, where there are currently insufficient managed hive rental services available to fruit and vegetable growers (<xref ref-type="bibr" rid="B5">Allsopp et al. 2008</xref>; <xref ref-type="bibr" rid="B70">Veldtman 2018</xref>), and only enough wild pollinators to satisfy 20% of horticultural pollination needs (<xref ref-type="bibr" rid="B4">Allsopp 2020</xref>). It is therefore reasonable to assume that the demand for pollination services is highly inelastic. Any reduction in commercial hive capacity caused by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> will produce a proportional increase in the price of hive rentals for fruit and vegetable growers, increasing their variable costs of production.</p>
      <p>In this paper, a model is described that was used to predict the likely economic cost of a spreading <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> population in South Africa. If the population has entered an accelerated growth phase, it is important to highlight the potential costs that could occur as a result, thereby communicating to policymakers the benefits of taking management actions now before the scale of the outbreak becomes unmanageable. This can happen rapidly, as overwintering nests can reach large sizes by the following summer and produce thousands of new queens (<xref ref-type="bibr" rid="B63">Tennant et al. 2011</xref>), and as new colonies take hold in an area, their foraging behavior will begin to affect the surrounding environment. Pollination services to horticulture will be affected as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> attacks managed and wild honeybee hives; potential feeding by wasps on fruit will affect wine grape growers; and household costs will rise due to the need to remove nests and avoid disruption to recreational activities. These effects were simulated, and their financial costs were summed over 50 years to highlight the economic significance of this pest and the potential return on investment in actions that slow or stop its spread. All costs are stated in United States dollars (<abbrev xlink:title="United States dollars">US$</abbrev>).</p>
    </sec>
    <sec sec-type="methods" id="sec2">
      <title>Methods</title>
      <sec sec-type="Sectors" id="sec3">
        <title>Sectors</title>
        <p>To predict the economic impacts from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> over time, the direct and indirect impacts on horticulture and direct impacts on wine grapes and households were considered.</p>
      </sec>
      <sec sec-type="Horticulture" id="sec4">
        <title>Horticulture</title>
        <p>Fruit and vegetable growers are highly dependent on managed honeybees for pollination services due to the intensity of production and the large seasonal demand for crop pollination (<xref ref-type="bibr" rid="B41">Melin et al. 2014</xref>; <xref ref-type="bibr" rid="B70">Veldtman 2018</xref>). South Africa is home to wild honeybee populations of the subspecies <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Apis">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mellifera">mellifera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="scutellata">scutellata</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Apis">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mellifera">mellifera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="capensis">capensis</tp:taxon-name-part></tp:taxon-name></italic>, which are also used in the honeybee industry (<xref ref-type="bibr" rid="B48">Pirk et al. 2016</xref>). It has been estimated that these wild populations and those of other native pollinators supply approximately one fifth of the pollination services required by fruit and vegetable growers (<xref ref-type="bibr" rid="B4">Allsopp 2020</xref>), while the remainder is supplied by a large commercial beekeeping industry. This often involves transporting hives over large distances for crop pollination (<xref ref-type="bibr" rid="B41">Melin et al. 2014</xref>), and periodic supply shortages can result at times of peak demand (<xref ref-type="bibr" rid="B70">Veldtman 2018</xref>).</p>
        <p>Currently, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> has a limited and localized impact on commercial honeybee hives where it occurs in South Africa. Typically, colonies are not destroyed, but the strength of the hive can be reduced to such an extent that it becomes unusable for crop pollination. If <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> becomes widespread in the region, it is expected to exacerbate supply shortages by attacking bees from commercial hives and reducing the number of colonies available for crop pollination (<xref ref-type="bibr" rid="B12">Clapperton et al. 1989</xref>), placing upward pressure on hive hire costs and raising variable production costs for fruit and vegetable growers. The horticultural industries used to simulate the effects of pollination disruption appear in Table <xref ref-type="table" rid="T2">2</xref>. Increased variable production costs were approximated using the number of hives required to pollinate each crop multiplied by the change in hive hire costs attributable to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> honeybee predation, and horticulture yield losses were approximated by the percentage change in commercial bee numbers (Table <xref ref-type="table" rid="T1">1</xref>).</p>
        <table-wrap id="T1" position="float" orientation="portrait">
          <label>Table 1.</label>
          <caption>
            <p>Fruit and vegetable industries affected by reductions in honeybee pollination services in South Africa.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="1" colspan="1">
                  <bold>Industry</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Area<sup>†</sup> (ha)</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Volume<sup>†</sup> (T)</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Gross Value<sup>†</sup> (<abbrev xlink:title="United States dollars">US$</abbrev>’000,000)</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Pollinator reliance<sup>‡</sup> (%)</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Beehives per ha<sup>§</sup> (#)</bold>
                </th>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Apples</td>
                <td rowspan="1" colspan="1">28,130</td>
                <td rowspan="1" colspan="1">1,198,900</td>
                <td rowspan="1" colspan="1">570.7</td>
                <td rowspan="1" colspan="1">0.50</td>
                <td rowspan="1" colspan="1">2</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Apricots</td>
                <td rowspan="1" colspan="1">3,010</td>
                <td rowspan="1" colspan="1">24,890</td>
                <td rowspan="1" colspan="1">30.1</td>
                <td rowspan="1" colspan="1">0.70</td>
                <td rowspan="1" colspan="1">1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Avocados</td>
                <td rowspan="1" colspan="1">22,050</td>
                <td rowspan="1" colspan="1">108,880</td>
                <td rowspan="1" colspan="1">81.9</td>
                <td rowspan="1" colspan="1">1.00</td>
                <td rowspan="1" colspan="1">4</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Cantaloupes</td>
                <td rowspan="1" colspan="1">1,480</td>
                <td rowspan="1" colspan="1">13,680</td>
                <td rowspan="1" colspan="1">4.1</td>
                <td rowspan="1" colspan="1">1.00</td>
                <td rowspan="1" colspan="1">1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Cherries</td>
                <td rowspan="1" colspan="1">1,060</td>
                <td rowspan="1" colspan="1">1,800</td>
                <td rowspan="1" colspan="1">7.8</td>
                <td rowspan="1" colspan="1">0.70</td>
                <td rowspan="1" colspan="1">4</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Cucumbers</td>
                <td rowspan="1" colspan="1">1,620</td>
                <td rowspan="1" colspan="1">28,960</td>
                <td rowspan="1" colspan="1">18.6</td>
                <td rowspan="1" colspan="1">1.00</td>
                <td rowspan="1" colspan="1">1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Mangoes</td>
                <td rowspan="1" colspan="1">4,990</td>
                <td rowspan="1" colspan="1">92,190</td>
                <td rowspan="1" colspan="1">84.5</td>
                <td rowspan="1" colspan="1">0.50</td>
                <td rowspan="1" colspan="1">2</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Macadamia nuts</td>
                <td rowspan="1" colspan="1">81,300</td>
                <td rowspan="1" colspan="1">87,230</td>
                <td rowspan="1" colspan="1">159.9</td>
                <td rowspan="1" colspan="1">0.95</td>
                <td rowspan="1" colspan="1">1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Peaches and nectarines</td>
                <td rowspan="1" colspan="1">10,100</td>
                <td rowspan="1" colspan="1">188,050</td>
                <td rowspan="1" colspan="1">181.4</td>
                <td rowspan="1" colspan="1">0.70</td>
                <td rowspan="1" colspan="1">1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Pears</td>
                <td rowspan="1" colspan="1">12,610</td>
                <td rowspan="1" colspan="1">481,360</td>
                <td rowspan="1" colspan="1">232.1</td>
                <td rowspan="1" colspan="1">0.50</td>
                <td rowspan="1" colspan="1">4</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Plums</td>
                <td rowspan="1" colspan="1">11,800</td>
                <td rowspan="1" colspan="1">94,290</td>
                <td rowspan="1" colspan="1">56.6</td>
                <td rowspan="1" colspan="1">0.70</td>
                <td rowspan="1" colspan="1">6</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Pumpkins</td>
                <td rowspan="1" colspan="1">37,190</td>
                <td rowspan="1" colspan="1">282,930</td>
                <td rowspan="1" colspan="1">117.7</td>
                <td rowspan="1" colspan="1">0.90</td>
                <td rowspan="1" colspan="1">1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Strawberries</td>
                <td rowspan="1" colspan="1">1,050</td>
                <td rowspan="1" colspan="1">13,270</td>
                <td rowspan="1" colspan="1">35.3</td>
                <td rowspan="1" colspan="1">0.40</td>
                <td rowspan="1" colspan="1">4</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Watermelons</td>
                <td rowspan="1" colspan="1">3,880</td>
                <td rowspan="1" colspan="1">63,690</td>
                <td rowspan="1" colspan="1">31.5</td>
                <td rowspan="1" colspan="1">1.00</td>
                <td rowspan="1" colspan="1">1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">TOTAL</td>
                <td rowspan="1" colspan="1">220,270</td>
                <td rowspan="1" colspan="1">2,680,120</td>
                <td rowspan="1" colspan="1">1,612.2</td>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1"/>
              </tr>
            </tbody>
          </table>
          <table-wrap-foot>
            <fn>
              <p><sup>†</sup><xref ref-type="bibr" rid="B26">FAO (2025)</xref>, except for macadamias taken from <xref ref-type="bibr" rid="B44">Moobi (2025)</xref> and <xref ref-type="bibr" rid="B66">UN Commtrade Database (2024)</xref>. <sup>‡</sup><xref ref-type="bibr" rid="B17">Cunningham et al. (2002)</xref>. <sup>§</sup><xref ref-type="bibr" rid="B5">Allsopp et al. (2008)</xref>; <xref ref-type="bibr" rid="B15">Cook et al. (2007)</xref>.</p>
            </fn>
          </table-wrap-foot>
        </table-wrap>
        <table-wrap id="T2" position="float" orientation="portrait">
          <label>Table 2.</label>
          <caption>
            <p>Parameters used to simulate the impact of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> on South African apiculture, horticulture, wine grapes, and households.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="2" colspan="1">
                  <bold>Parameter</bold>
                </th>
                <th rowspan="1" colspan="4">
                  <bold>Sector</bold>
                </th>
              </tr>
              <tr>
                <th rowspan="1" colspan="1">
                  <bold>Pollination</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Horticulture</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Winegrapes</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Households</bold>
                </th>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Maximum units affected, <mml:math id="M20"><mml:msubsup><mml:mi>S</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup></mml:math> (#)<sup>†</sup></td>
                <td rowspan="1" colspan="1">450 000</td>
                <td rowspan="1" colspan="1">see Table <xref ref-type="table" rid="T2">2</xref></td>
                <td rowspan="1" colspan="1">87 850</td>
                <td rowspan="1" colspan="1">185 000</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Maximum proportion affected, <mml:math id="M21"><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup></mml:math> (%)<sup>‡</sup></td>
                <td rowspan="1" colspan="1">Uniform(5, 10)</td>
                <td rowspan="1" colspan="1">Uniform(60, 70)</td>
                <td rowspan="1" colspan="1">Uniform(20, 30)</td>
                <td rowspan="1" colspan="1">Uniform(80, 90)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Minimum proportion affected, <mml:math id="M22"><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">min</mml:mo></mml:mrow></mml:msubsup></mml:math> (%)<sup>§</sup></td>
                <td rowspan="1" colspan="1">0.01</td>
                <td rowspan="1" colspan="1">0.01</td>
                <td rowspan="1" colspan="1">0.01</td>
                <td rowspan="1" colspan="1">0.01</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Proportion affected at <mml:math id="M23"><mml:msup><mml:mi>t</mml:mi><mml:mrow><mml:msub><mml:mi>θ</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:msup></mml:math>, <italic>θ<sub>i</sub></italic> (%)<sup>§</sup></td>
                <td rowspan="1" colspan="1">Uniform(4, 5)</td>
                <td rowspan="1" colspan="1">Uniform(50, 60)</td>
                <td rowspan="1" colspan="1">Uniform(10, 20)</td>
                <td rowspan="1" colspan="1">Uniform(70, 80)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Time taken for <italic>θ<sub>i</sub></italic> to be affected (yr)<sup>§</sup></td>
                <td rowspan="1" colspan="1">Uniform(30, 50)</td>
                <td rowspan="1" colspan="1">Uniform(30, 50)</td>
                <td rowspan="1" colspan="1">Uniform(30, 50)</td>
                <td rowspan="1" colspan="1">Uniform(30, 50)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Yield loss despite control, <italic>Y<sub>it</sub></italic> (%)<sup>|</sup></td>
                <td rowspan="1" colspan="1">5–10</td>
                <td rowspan="1" colspan="1">see Fig. <xref ref-type="fig" rid="F3">3</xref></td>
                <td rowspan="1" colspan="1">0</td>
                <td rowspan="1" colspan="1">na</td>
              </tr>
              <tr>
                <td rowspan="14" colspan="1">Price of per unit, <italic>P<sub>it</sub></italic> ($US/T)<sup>¶</sup></td>
                <td rowspan="14" colspan="1">75</td>
                <td rowspan="1" colspan="1">Apples 476</td>
                <td rowspan="14" colspan="1">332</td>
                <td rowspan="14" colspan="1">na</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Apricots 1209</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Avocados 752</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Cantaloupes 301</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Cherries 4334</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Cucumbers 640</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Mangoes 916</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Macadamia nuts 1834</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Peaches and nectarines 964</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Pears 482</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Plums 600</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Pumpkins 416</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Strawberries 2658</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Watermelons 493</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Increased variable cost, <italic>V<sub>it</sub></italic> (<abbrev xlink:title="United States dollars">US$</abbrev>)<sup>#</sup></td>
                <td rowspan="1" colspan="1">na</td>
                <td rowspan="1" colspan="1">see Fig. <xref ref-type="fig" rid="F3">3</xref></td>
                <td rowspan="1" colspan="1">Uniform(115, 150)</td>
                <td rowspan="1" colspan="1">Uniform(200, 250)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Elasticity of demand for pollination services, <italic>e</italic> (unitless)<sup>§</sup></td>
                <td rowspan="1" colspan="1">Uniform(−0.5, −0.2)</td>
                <td rowspan="1" colspan="1">na</td>
                <td rowspan="1" colspan="1">na</td>
                <td rowspan="1" colspan="1">na</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Discount rate, <italic>v</italic> (%)<sup>††</sup></td>
                <td rowspan="1" colspan="1">na</td>
                <td rowspan="1" colspan="1">Uniform(5, 7)</td>
                <td rowspan="1" colspan="1">Uniform(5, 7)</td>
                <td rowspan="1" colspan="1">Uniform(5, 7)</td>
              </tr>
            </tbody>
          </table>
          <table-wrap-foot>
            <fn>
              <p><sup>†</sup> Pollination – no. hives (R. van Zyl, South African National Department of Agriculture, pers. comm.). Horticulture – no. ha, <xref ref-type="bibr" rid="B26">FAO (2025)</xref>. Winegrapes – no. ha, <xref ref-type="bibr" rid="B27">Floris-Samuels (2024)</xref>. Households – no. middle-class households × 5%, <xref ref-type="bibr" rid="B35">Lemanski (2017)</xref>; <xref ref-type="bibr" rid="B47">Pest Control South Africa (2025)</xref>; <xref ref-type="bibr" rid="B62">Statistics South Africa (2023)</xref>. <sup>‡</sup> Pollination – <xref ref-type="bibr" rid="B14">Cook (2019)</xref>. Horticulture – <xref ref-type="bibr" rid="B4">Allsopp (2020)</xref>. Winegrapes – <xref ref-type="bibr" rid="B14">Cook (2019)</xref>; <xref ref-type="bibr" rid="B31">Goodall and Smith (2001)</xref>; <xref ref-type="bibr" rid="B64">Thomas (1993)</xref>. Households – <xref ref-type="bibr" rid="B14">Cook (2019)</xref>. <sup>§</sup> Plausible estimate. <sup>|</sup> Pollination – see Fig. <xref ref-type="fig" rid="F3">3</xref>. Horticulture – <xref ref-type="bibr" rid="B31">Goodall and Smith (2001)</xref>; <xref ref-type="bibr" rid="B64">Thomas (1993)</xref>. Winegrapes – <xref ref-type="bibr" rid="B14">Cook (2019)</xref>. <sup>¶</sup> Pollination – <xref ref-type="bibr" rid="B73">Western Cape Bee Industry Association (2024)</xref>. Horticulture – <xref ref-type="bibr" rid="B26">FAO (2025)</xref>. Winegrapes – <xref ref-type="bibr" rid="B27">Floris-Samuels (2024)</xref>. <sup>#</sup> Horticulture – <xref ref-type="bibr" rid="B14">Cook (2019)</xref>. Winegrapes – <xref ref-type="bibr" rid="B14">Cook (2019)</xref>. Households – <xref ref-type="bibr" rid="B30">Get Pest Control (2024)</xref>; <xref ref-type="bibr" rid="B47">Pest Control South Africa (2025)</xref>; <xref ref-type="bibr" rid="B51">Procompare (2025)</xref>. <sup>††</sup><xref ref-type="bibr" rid="B13">Commonwealth of Australia (2006)</xref>.</p>
            </fn>
          </table-wrap-foot>
        </table-wrap>
      </sec>
      <sec sec-type="Winegrapes" id="sec5">
        <title>Winegrapes</title>
        <p>The feeding behavior of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> can damage wine grapes on the vine. The wasps hollow out wine grapes, causing yield losses and introducing foreign yeasts that interfere with fermentation (<xref ref-type="bibr" rid="B7">Bashford 2001</xref>). In some grape-growing areas where they have not been managed, yield losses of 10–15% have been observed (<xref ref-type="bibr" rid="B31">Goodall and Smith 2001</xref>; <xref ref-type="bibr" rid="B64">Thomas 1993</xref>). This places an onus on vineyard managers to bait to control wasp numbers and reduce damage to fruit and vegetables, which adds to their costs of production. Bait stations deployed at a density of approximately five stations per hectare per year, estimated to cost approximately US$120–150 per hectare (<xref ref-type="bibr" rid="B77">Wine Tasmania 2018</xref>), were assumed to reduce yield losses to negligible levels.</p>
        <p>South African wine grape production is most prominent in the Western Cape province, with smaller-scale production occurring in the Northern Cape and KwaZulu-Natal provinces. The Cape Winelands region is the most significant in terms of wine production, encompassing the Stellenbosch, Paarl, Franschhoek, and Constantia regions. Other notable areas include the Breede River Valley, Olifants River Valley, and the Cape South Coast. Production in 2022/23 took place over an area of 87,850 hectares, producing approximately 1.2 million tonnes of wine grapes valued at approximately US$393.2 million (<xref ref-type="bibr" rid="B27">Floris-Samuels 2024</xref>).</p>
      </sec>
      <sec sec-type="Households" id="sec6">
        <title>Households</title>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> can become a serious pest in urban areas if uncontrolled, especially in warmer climates where breeding and nest construction can occur continuously, resulting in large summer colonies containing many thousands of individuals (<xref ref-type="bibr" rid="B63">Tennant et al. 2011</xref>). Health statistics related specifically to wasp stings are unavailable. A total of 109 deaths were recorded in South Africa from contact with wasps and honeybees for the period 2001–2011 (<xref ref-type="bibr" rid="B23">du Toit-Prinsloo et al. 2016</xref>). However, the nuisance value associated with large colonies near dwellings as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> spreads into urban areas is assumed to be large enough to motivate some householders to invest in private pest management services in the absence of government-provided services.</p>
        <p>In 2022, South Africa had a population of approximately 61.4 million living in 18.5 million dwellings (<xref ref-type="bibr" rid="B62">Statistics South Africa 2023</xref>). It is difficult to estimate how many households would engage pest management services to exterminate wasp nests, but in the absence of specific information, a proportion of upper- and middle-class households was used as a proxy. These are households with a level of disposable income high enough to warrant expenditure on professional pest insect removal services. What constitutes a middle-class household is debatable, but it was assumed that 20% of households fall into this category (<xref ref-type="bibr" rid="B35">Lemanski 2017</xref>). It was further assumed that 5% of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> colonies located in urban areas pose sufficient risk to residents’ health and recreation activities to justify expenditure on professional pest control services. The cost of such services is assumed to be US$120–300 per nest (<xref ref-type="bibr" rid="B30">Get Pest Control 2024</xref>; <xref ref-type="bibr" rid="B47">Pest Control South Africa 2025</xref>; <xref ref-type="bibr" rid="B51">Procompare 2025</xref>).</p>
      </sec>
      <sec sec-type="Simulation model" id="sec7">
        <title>Simulation model</title>
        <p>Following <xref ref-type="bibr" rid="B14">Cook (2019)</xref>, future impacts of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> were estimated using a Monte Carlo simulation model. Uncertain parameters were specified as distributions, and a Latin hypercube sampling algorithm was used to sample from each using the MonteCarlo package (<xref ref-type="bibr" rid="B36">Leschinski 2019</xref>) for the R programming language (<xref ref-type="bibr" rid="B53">R Core Team 2023</xref>). Parameter distributions used in the model are uniform or rectangular distributions bounded by minimum and maximum values.</p>
        <p>Different sectors affected by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in the model (i.e., horticulture, viticulture, and households) are denoted <italic>i</italic>, and the simulation was divided into yearly time steps, denoted <italic>t</italic>. To predict the proportion of each sector affected from one time step to the next, the model incorporated a <xref ref-type="bibr" rid="B59">Schaefer (1957)</xref> logistic growth equation, whereby the length of time taken for horticulture, wine grapes, and households to be affected was specified using plausible distributions. This model was combined with a measure of the marginal cost to simulate economic costs over time.</p>
        <p>The proportion of sector <italic>i</italic> affected in time step <italic>t</italic>(<italic>S<sub>it</sub></italic>) was estimated as:</p>
        <p><mml:math id="M1"><mml:msub><mml:mi>S</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:msubsup><mml:mi>S</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup><mml:mfrac><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup><mml:mrow><mml:mn>1</mml:mn><mml:mo>+</mml:mo><mml:mrow><mml:mo>(</mml:mo><mml:mfrac><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">min</mml:mo></mml:mrow></mml:msubsup></mml:mfrac><mml:mo>−</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow><mml:msup><mml:mi>e</mml:mi><mml:mrow><mml:mo>−</mml:mo><mml:msub><mml:mi>ω</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mi>t</mml:mi></mml:mrow></mml:msup></mml:mrow></mml:mfrac></mml:math>, (1)</p>
        <p>where:</p>
        <p><mml:math id="M2"><mml:msubsup><mml:mi>S</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup></mml:math> is the total number of units of <italic>i</italic> potentially affected (i.e., ha × pollinator reliance for horticulture, ha winegrapes, and the number of dwellings for households);</p>
        <p><mml:math id="M3"><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup></mml:math> is the maximum proportion of sector <italic>i</italic> affected;</p>
        <p><mml:math id="M4"><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">min</mml:mo></mml:mrow></mml:msubsup></mml:math> is the proportion of sector <italic>i</italic> affected in timestep 0;</p>
        <p><italic>ω<sub>i</sub></italic> is the rate of spread or transition from <mml:math id="M5"><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">min</mml:mo></mml:mrow></mml:msubsup></mml:math> to <mml:math id="M6"><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup></mml:math>.</p>
        <p>A hypothetical value of <italic>ω<sub>i</sub></italic> was estimated by specifying a proportion of each sector close to <mml:math id="M7"><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup></mml:math> and a probable number of time steps taken before <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> affects this proportion, such that:</p>
        <p><mml:math id="M8"><mml:msub><mml:mi>ω</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mo>−</mml:mo><mml:msup><mml:mi>t</mml:mi><mml:mrow><mml:msub><mml:mi>θ</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:msup><mml:mi>ln</mml:mi><mml:mo>⁡</mml:mo><mml:mrow><mml:mo>[</mml:mo><mml:mfrac><mml:mrow><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup><mml:mo>−</mml:mo><mml:msub><mml:mi>θ</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>θ</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mrow><mml:mo>(</mml:mo><mml:mfrac><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup><mml:msubsup><mml:mi>I</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">min</mml:mo></mml:mrow></mml:msubsup></mml:mfrac><mml:mo>−</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:mrow></mml:mfrac><mml:mo>]</mml:mo></mml:mrow></mml:math>, (2)</p>
        <p>where:</p>
        <p><italic>θ<sub>i</sub></italic> is a specified proportion of <mml:math id="M9"><mml:msubsup><mml:mi>S</mml:mi><mml:mi>i</mml:mi><mml:mrow><mml:mo movablelimits="true">max</mml:mo></mml:mrow></mml:msubsup></mml:math> affected;</p>
        <p><mml:math id="M10"><mml:msup><mml:mi>t</mml:mi><mml:mrow><mml:msub><mml:mi>θ</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:msup></mml:math> is the number of time steps taken for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> to affect <italic>θ<sub>i</sub></italic>.</p>
        <p>The value of damage to <italic>n</italic> sectors caused by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in time step <italic>t</italic> if no coordinated response actions are taken to mitigate spread (<italic>d<sub>t</sub></italic>) was estimated as:</p>
        <p><mml:math id="M11"><mml:msub><mml:mi>d</mml:mi><mml:mi>t</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:msubsup><mml:mrow><mml:mo>∑</mml:mo></mml:mrow><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>n</mml:mi></mml:msubsup><mml:msub><mml:mi>S</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:msub><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>Y</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:msub><mml:msub><mml:mi>P</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:msub><mml:mo>+</mml:mo><mml:msub><mml:mi>V</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math>, (3)</p>
        <p>where:</p>
        <p><italic>Y<sub>it</sub></italic> is the mean change in yield in sector <italic>i</italic> attributable to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in time step <italic>t</italic>;</p>
        <p><italic>P<sub>it</sub></italic> is the price per unit in sector <italic>i</italic> (i.e., ha, dwellings) in time step <italic>t</italic>;</p>
        <p><italic>V<sub>it</sub></italic> is the increase in variable cost per unit in sector <italic>i</italic> caused by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in time step <italic>t</italic>.</p>
        <p>For horticulture and wine grapes, an estimate of price, <italic>P<sub>it</sub></italic>, was specified for the first time step of the model (i.e., <italic>P<sub>it=0</sub></italic>, corresponding to the year 2024). In the case of wine grapes, this was assumed to remain constant in real terms throughout the simulation period since the change in supply conditions induced by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> is relatively minor. However, in the case of horticulture, the price was assumed to change as the supply of managed pollination services contracts due to wasp predation, in turn raising the price for beehive hire.</p>
        <p>If any change in price for pollination services is directly passed on to horticulture growers, the increase in variable production costs for fruits and vegetables could be estimated by simulating price effects in pollination services as their supply contracts in response to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> predation. Further assuming an isoelastic demand schedule for pollination services, the change in the price of commercial hives for pollination was estimated as:</p>
        <p><mml:math id="M12"><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>P</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup><mml:mo>=</mml:mo><mml:mfrac><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>Q</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup></mml:mrow><mml:mi>e</mml:mi></mml:mfrac></mml:math>, (4)</p>
        <p>where:</p>
        <p><mml:math id="M13"><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>P</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup></mml:math> is the proportionate change in the price of commercial beehives from one time step to the next induced by a contraction in the supply of pollination services attributable to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> predation (i.e., <mml:math id="M14"><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>P</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup><mml:mo>=</mml:mo><mml:mfrac><mml:mrow><mml:msubsup><mml:mi>P</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mo>−</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>H</mml:mi></mml:msubsup><mml:mo>−</mml:mo><mml:msubsup><mml:mi>P</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup></mml:mrow><mml:msubsup><mml:mi>P</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mo>−</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>H</mml:mi></mml:msubsup></mml:mfrac></mml:math>);</p>
        <p><mml:math id="M15"><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>Q</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup></mml:math> is the proportionate change in the quantity of beehives available for pollination (i.e., <mml:math id="M16"><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>Q</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup><mml:mo>=</mml:mo><mml:mfrac><mml:mrow><mml:msubsup><mml:mi>Q</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mo>−</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>H</mml:mi></mml:msubsup><mml:mo>−</mml:mo><mml:msubsup><mml:mi>Q</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup></mml:mrow><mml:msubsup><mml:mi>Q</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mo>−</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>H</mml:mi></mml:msubsup></mml:mfrac></mml:math>);</p>
        <p><italic>e</italic> is the elasticity of demand for pollination services.</p>
        <p>Since the change in the price of pollination services was assumed to be passed on to horticulturalists, the increase in the variable cost of producing horticultural product <italic>j</italic> in time step <italic>t</italic>, <italic>V<sub>jt</sub></italic>, was estimated as <mml:math id="M17"><mml:msub><mml:mi>V</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:msub><mml:mi>V</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>t</mml:mi><mml:mo>−</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:msub><mml:mo>+</mml:mo><mml:mrow><mml:mo>(</mml:mo><mml:msubsup><mml:mi>P</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mo>−</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>H</mml:mi></mml:msubsup><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>P</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup><mml:mo>)</mml:mo></mml:mrow></mml:math>. The change in yield in horticulture production was also assumed to be directly proportional to the pollination dependence of the product concerned. Therefore, given a pollinator reliance of horticultural product <italic>j</italic> of <italic>r<sub>j</sub></italic> (Table <xref ref-type="table" rid="T2">2</xref>), the change in yield loss experienced by growers in time step <italic>t</italic>, <italic>Y<sub>jt</sub></italic>, was estimated as <mml:math id="M18"><mml:msub><mml:mi>Y</mml:mi><mml:mrow><mml:mi>j</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mi mathvariant="normal">Δ</mml:mi><mml:msubsup><mml:mi>Q</mml:mi><mml:mi>t</mml:mi><mml:mi>H</mml:mi></mml:msubsup><mml:msub><mml:mi>r</mml:mi><mml:mi>j</mml:mi></mml:msub></mml:math>.</p>
        <p>The value of damages, <italic>d<sub>t</sub></italic>, accrued over time and was therefore subject to discounting. Discounting has an erosive effect on monetary values that increases with time, which means the present value of one unit of damage caused early in the simulation was worth more in real terms than the same amount of damage caused later in the simulation. The present value of economic damage from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> over <italic>T</italic> time steps (<italic>PVd<sub>T</sub></italic>) was calculated as:</p>
        <p><mml:math id="M19"><mml:mi>P</mml:mi><mml:mi>V</mml:mi><mml:msub><mml:mi>d</mml:mi><mml:mi>T</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:msubsup><mml:mrow><mml:mo>∑</mml:mo></mml:mrow><mml:mrow><mml:mi>t</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>T</mml:mi></mml:msubsup><mml:mrow><mml:mo>[</mml:mo><mml:mfrac><mml:msub><mml:mi>d</mml:mi><mml:mi>t</mml:mi></mml:msub><mml:mrow><mml:mo stretchy="false">(</mml:mo><mml:mn>1</mml:mn><mml:mo>+</mml:mo><mml:mi>v</mml:mi><mml:msup><mml:mo stretchy="false">)</mml:mo><mml:mi>t</mml:mi></mml:msup></mml:mrow></mml:mfrac><mml:mo>]</mml:mo></mml:mrow></mml:math>, (5)</p>
        <p>where:</p>
        <p><italic>v</italic> is the discount rate.</p>
        <p>The values assigned to each parameter in relation to each sector are provided in Table <xref ref-type="table" rid="T2">2</xref>. When the specific values of parameters were uncertain, they were specified as uniform distributions bookended by minimum and maximum values. Using the Monte Carlo method, one value was randomly sampled from each distribution and used to calculate spread and impact over time in each of the 5,000 iterations of the model.</p>
      </sec>
    </sec>
    <sec sec-type="Results" id="sec8">
      <title>Results</title>
      <p>The extent of losses experienced in each sector is expressed in appropriate units in Fig. <xref ref-type="fig" rid="F2">2</xref>, which includes the effects of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> on commercial beehives. Financial losses to the pollination industry were not quantified in the assessment but were assumed to affect horticulture producers indirectly, as they were passed on as increased hive rental costs and yield losses from wasps attacking bees in beekeepers’ apiaries. The number of commercial beehives affected over time is therefore shown in panel A, while the area of horticulture affected is shown in panel B. The area of wine grapes affected is shown in panel C, and the number of households likely to be affected is shown in panel D. The box-whisker plots show the 5<sup>th</sup>, 25<sup>th</sup>, median, 75<sup>th</sup>, and 95<sup>th</sup> percentile values.</p>
      <fig id="F2">
        <object-id content-type="arpha">0935B275-DDEB-5523-8AFD-552AEA8DFE09</object-id>
        <label>Figure 2.</label>
        <caption>
          <p>Cumulative losses likely to be experienced in different sectors because of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in South Africa over 50 years. The plots show the 5<sup>th</sup>, 25<sup>th</sup>, median, 75<sup>th</sup>, and 95<sup>th</sup> percentiles of simulation model outputs. Panels (<bold>A</bold>–<bold>D</bold>) show the number of commercial beehives affected, area of horticulture affected, area of wine grapes affected, and number of households affected, respectively. After 50 years, it is estimated that 6,500–20,400 commercial beehives could experience losses from predation (panel <bold>A</bold>), 127,000–136,000 ha of horticulture crops could experience yield loss and rising production costs (panel <bold>B</bold>), 10,000–19,000 ha of wine grapes could experience rising production costs (panel <bold>C</bold>), and 109,000–120,000 households could experience increased pest management expenses (panel <bold>D</bold>).</p>
        </caption>
        <graphic xlink:href="neobiota-109-067_article-191014__-g002.jpg" id="oo_1773302.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1773302</uri>
        </graphic>
      </fig>
      <p>With the set of assumptions used in the model, it was estimated that 6,500–20,400 beehives would be affected by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> attacking foraging bees after 50 years (panel A). This range corresponds to the 25<sup>th</sup> and 75<sup>th</sup> percentiles, indicating that 50% of model iterations produced results between these values. Likewise, the results showed that 127,000–136,000 ha of horticulture crops would be affected by yield losses and rising production costs (panel B), 10,000–19,000 ha of wine grapes would be affected by higher production costs because of wasp control expenses (panel C), and 109,000–120,000 households would experience increased pest removal expenses (panel D) by the end of the simulation period.</p>
      <p>The predicted indirect effects on horticulture from induced changes in the commercial pollination industry are shown in Fig. <xref ref-type="fig" rid="F3">3</xref>. Panel A shows the change in the price per hive paid for pollination services in real or present-value terms. As foraging bees are attacked by spreading <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> colonies, the resulting contraction in supply places upward pressure on the hire price. This effect had a negligible impact on hive hire prices for the first 25–30 years but increased rapidly thereafter. After 50 years, the estimated price per hive increased from US$75.00 to US$78.20–84.50. Panel B shows the estimated yield lost in horticulture industries due to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> attacking bees. This had the same lag before losses accelerated, with yield losses increasing by 4–13% by the end of the 50-year simulation.</p>
      <fig id="F3">
        <object-id content-type="arpha">3ADBB44F-8BAC-5336-BD47-8C49597A3861</object-id>
        <label>Figure 3.</label>
        <caption>
          <p>Increased beehive hire costs and horticulture yield losses resulting from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> spread in South Africa over 50 years. The plots show the 5<sup>th</sup>, 25<sup>th</sup>, median, 75<sup>th</sup>, and 95<sup>th</sup> percentiles of simulation model outputs. Panel (<bold>A</bold>) shows the price charged to horticulturalists to hire beehives for pollination services, and panel (<bold>B</bold>) shows yield losses in horticulture crops caused by bee predation. After 50 years, it is estimated that the price of managed hives for pollination could increase from US$75.00/hive to US$78.20–84.50/hive and that horticultural yield losses could increase by 4–13%.</p>
        </caption>
        <graphic xlink:href="neobiota-109-067_article-191014__-g003.jpg" id="oo_1773303.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1773303</uri>
        </graphic>
      </fig>
      <p>The present value of damages predicted in the model simulations (i.e., <italic>PVd<sub>T</sub></italic>) is shown in Fig. <xref ref-type="fig" rid="F4">4</xref>. Panels A, B, and C show predicted <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> impacts on growers of pollinator-dependent horticulture crops, wine grape growers, and householders, respectively, while panel D shows the summed value of damages across all three sectors.</p>
      <fig id="F4">
        <object-id content-type="arpha">E7C606B1-2D12-5E87-A18C-E05FD791CC8F</object-id>
        <label>Figure 4.</label>
        <caption>
          <p>Cumulative value of damages predicted from the simulation of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> spread in South Africa over 50 years. The plots show the 5<sup>th</sup>, 25<sup>th</sup>, median, 75<sup>th</sup>, and 95<sup>th</sup> percentiles. Panels (<bold>A</bold>–<bold>C</bold>) show the value of cumulative damage to horticulture industries, wine grape production, and households, respectively. Panel (<bold>D</bold>) shows the combined value of damages across all three sectors. After 50 years, combined damages were estimated at US$160–300 million. Households accounted for 82% of these losses, horticulture for 17%, and wine grapes for 1%.</p>
        </caption>
        <graphic xlink:href="neobiota-109-067_article-191014__-g004.jpg" id="oo_1773304.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1773304</uri>
        </graphic>
      </fig>
      <p>Disruption to honeybee pollination services is likely to cost horticulture industries US$30–55 million over 50 years. Costs related to wine grape production over the same period were estimated to be US$1–3 million, while costs incurred by households were estimated to be US$110–260 million. The summed value of damages from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> across all sectors was US$160–300 million after 50 years, with a median value of US$225 million (Table <xref ref-type="table" rid="T3">3</xref>).</p>
      <table-wrap id="T3" position="float" orientation="portrait">
        <label>Table 3.</label>
        <caption>
          <p>Results.</p>
        </caption>
        <table>
          <tbody>
            <tr>
              <th rowspan="2" colspan="1">
                <bold>Sector</bold>
              </th>
              <th rowspan="1" colspan="3">
                <bold>Value of damages (<abbrev xlink:title="United States dollars">US$</abbrev> million)</bold>
              </th>
            </tr>
            <tr>
              <th rowspan="1" colspan="1">
                <bold>25%</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>50%</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>75%</bold>
              </th>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Horticulture</td>
              <td rowspan="1" colspan="1">30</td>
              <td rowspan="1" colspan="1">40</td>
              <td rowspan="1" colspan="1">55</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Winegrapes</td>
              <td rowspan="1" colspan="1">1</td>
              <td rowspan="1" colspan="1">2</td>
              <td rowspan="1" colspan="1">3</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Households</td>
              <td rowspan="1" colspan="1">110</td>
              <td rowspan="1" colspan="1">180</td>
              <td rowspan="1" colspan="1">260</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Combined</td>
              <td rowspan="1" colspan="1">160</td>
              <td rowspan="1" colspan="1">225</td>
              <td rowspan="1" colspan="1">300</td>
            </tr>
          </tbody>
        </table>
      </table-wrap>
      <p>Given the uncertainties in many model parameters, a sensitivity analysis was conducted to determine which exerted the greatest influence on the results. A simple approach was used, involving each parameter being set to the minimum and maximum values of its range while all other parameters were held constant. The effect that the minimum and maximum values for each parameter had on the present value of combined costs is reported in Fig. <xref ref-type="fig" rid="F5">5</xref>. Here, the influence of each parameter is indicated by the length of the corresponding bars, and the parameters are ranked from top to bottom in order of the change they produced in the present value of cumulative costs after 50 years.</p>
      <fig id="F5">
        <object-id content-type="arpha">8D00A6CA-563D-52EE-B2B6-D46E3961AA9F</object-id>
        <label>Figure 5.</label>
        <caption>
          <p>Sensitivity of total costs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> spread in South Africa to changes in model parameters. Parameters are ranked from top to bottom according to the sensitivity of the present value of cumulative damages over 50 years to changes in each parameter, with the length of the corresponding bars reflecting the changes in results produced. Each parameter specified as a distribution was changed to the minimum and maximum values of its range while all other parameters remained unchanged, and the resultant cost estimate was recorded. Results were most sensitive to changes in the rate of spread and the discount rate.</p>
        </caption>
        <graphic xlink:href="neobiota-109-067_article-191014__-g005.jpg" id="oo_1773305.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1773305</uri>
        </graphic>
      </fig>
      <p>The results were most sensitive to changes in the rate at which <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> was assumed to spread. The estimation technique used to shape the logistic spread function (Eq. 2) specified a proportion of each sector, denoted <italic>θ</italic>, close to the asymptote to indicate where the function began to plateau. The number of time steps taken before <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> affected <italic>θ</italic>, <italic>t<sup>θ</sup></italic>, determined the rate of spread. For all sectors, <italic>t<sup>θ</sup></italic> was assumed to be 30–50 years (Table <xref ref-type="table" rid="T1">1</xref>). Increasing <italic>t<sup>θ</sup></italic> to 50 years lowered the median value of damages from US$225 million to US$115 million (i.e., –50%), and lowering <italic>t<sup>θ</sup></italic> to 30 years increased the median value of damages to US$345 million (+55%).</p>
      <p>Large changes in the results were also observed when varying the discount rate. The simulation period was relatively long at 50 years, and cost accumulation was skewed to later time steps, so the erosive effects of discounting were pronounced. Lowering the discount rate to 5% from its median value of 6% (i.e., –17%) produced an increase in the median value of damages to US$335 million (+49%). Likewise, increasing the discount rate to 7% (+17%) lowered the present value of costs to US$160 million (−28%). The discount rates used were suggested in the <xref ref-type="bibr" rid="B13">Commonwealth of Australia (2006)</xref>, but it was acknowledged that choosing an appropriate value for this parameter in a case study with cost streams that extend so far into the future is difficult (<xref ref-type="bibr" rid="B1">Abelson and Dalton 2018</xref>).</p>
      <p>Results were also relatively sensitive to changes in the parameter <italic>θ</italic>, which was used to determine the asymptotic phase of the spread function. This was assumed to be 50–60% for horticulture, 10–20% for wine grapes, and 70–80% for households. Increasing <italic>θ</italic> to the maximum of its range for each sector increased the median value of damages to US$280 million (i.e., +25%), and lowering <italic>θ</italic> to the minimum value of its range for each sector lowered the median value of damages to US$190 million (−15%).</p>
    </sec>
    <sec sec-type="Discussion" id="sec9">
      <title>Discussion</title>
      <p>The model outlined in this paper is a first attempt to estimate the potential financial impacts <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> could have in South Africa if it continues to spread unabated over the next 50 years. Although relatively simple, it captures important spread characteristics that make this a difficult species to deal with, namely its propensity to establish and maintain a relatively low economic cost for several decades before population growth and damage accelerate. The model aggregates economic impacts over time, indicating the financial implications of the spread to communities and agricultural industries. To this end, it provides information that could be used in future decisions concerning national response policies.</p>
      <p>There is a growing literature highlighting increasing economic costs associated with invasive species, but this often relies on aggregated impacts from multiple species (<xref ref-type="bibr" rid="B25">Early et al. 2016</xref>; <xref ref-type="bibr" rid="B45">Paini et al. 2016</xref>; <xref ref-type="bibr" rid="B18">Cuthbert et al. 2022</xref>; <xref ref-type="bibr" rid="B50">Polce et al. 2023</xref>; <xref ref-type="bibr" rid="B28">Gallardo et al. 2024</xref>; <xref ref-type="bibr" rid="B61">Soto et al. 2025</xref>). These are frequently cited in the introductory sections of many research papers and articles to highlight the scale of the issue, but beyond this, their use is questionable. Policymakers are generally aware that the invasive species problem is dire and the costs are enormous, but they are seldom informed by predictive analyses of what the effects of response policies enacted in the present might be (<xref ref-type="bibr" rid="B55">Regan et al. 2002</xref>; <xref ref-type="bibr" rid="B72">Waage et al. 2005</xref>).</p>
      <p>The model provides policymakers with prospective policy analysis. Although this can be achieved using simple partial budgeting techniques or deterministic models (<xref ref-type="bibr" rid="B21">de Wit et al. 2001</xref>; <xref ref-type="bibr" rid="B40">MacLeod et al. 2004</xref>; <xref ref-type="bibr" rid="B8">Bolda et al. 2010</xref>), more complex approaches such as the one set out in this paper rely on stochastic simulation to specifically incorporate policy risks (<xref ref-type="bibr" rid="B54">Rafoss 2003</xref>; <xref ref-type="bibr" rid="B79">Yemshanov et al. 2009</xref>; <xref ref-type="bibr" rid="B14">Cook 2019</xref>). Other approaches that integrate ecological models with economic management frameworks (<xref ref-type="bibr" rid="B3">Alexei and Andrew 1998</xref>; <xref ref-type="bibr" rid="B60">Sharov and Liebhold 1998</xref>; <xref ref-type="bibr" rid="B6">Barbier 2001</xref>; <xref ref-type="bibr" rid="B10">Cacho et al. 2008</xref>; <xref ref-type="bibr" rid="B34">Hyder et al. 2008</xref>; <xref ref-type="bibr" rid="B11">Carrasco et al. 2010</xref> ) or use metapopulation models to forecast future impacts (<xref ref-type="bibr" rid="B2">Albers et al. 2010</xref>; <xref ref-type="bibr" rid="B58">Sanchirico et al. 2010</xref>; <xref ref-type="bibr" rid="B46">Pepper et al. 2019</xref>) can also highlight complex interactions between spreading populations and heterogeneous landscapes to policy audiences. However, these models can be expensive to develop and require technical skills to use, and their complexity can make results difficult to communicate to policymakers.</p>
      <p>In the current paper, it was assumed that no coordinated eradication or spread-mitigation response will be mounted as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> spreads in South Africa over the next 50 years, but the model can be used to evaluate future response policies. For instance, the potential human health consequences of this pest as it spreads could create a political imperative for authorities to respond, particularly around the 20-year mark when the simulations suggest spread will reach an inflection point. A benefit-cost analysis of any prospective response policy could simply be carried out using these results as a counterfactual scenario and comparing them with the potential impacts of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> in a policy scenario.</p>
      <p>A simple example drawing on the results of the sensitivity analysis can be used to explain how a benefit-cost analysis could be carried out. Sensitivity test results (Fig. <xref ref-type="fig" rid="F5">5</xref>) suggest that if a spread-mitigation policy can increase the parameter <italic>t</italic>θ to 50 years, the value of damages will be reduced by US$110 million over 30 years, or approximately US$3.7 million per year. This represents the threshold or break-even investment cost for the policy, which might take the form of surveillance activities to detect and destroy wasp nests (<xref ref-type="bibr" rid="B24">Duthie and Lester 2013</xref>; <xref ref-type="bibr" rid="B14">Cook 2019</xref>), subsidies for pest control services (<xref ref-type="bibr" rid="B57">Saga et al. 2025</xref>), bounties for nest reporting (<xref ref-type="bibr" rid="B32">Hester and Cacho 2012</xref>), or potentially biological control programs (<xref ref-type="bibr" rid="B56">Rose et al. 1999</xref>; <xref ref-type="bibr" rid="B43">Merino et al. 2013</xref>; <xref ref-type="bibr" rid="B9">Cacho and Hester 2022</xref>; <xref ref-type="bibr" rid="B33">Hester et al. 2024</xref>; <xref ref-type="bibr" rid="B69">van Zyl et al. 2024</xref>). If the policy costs less than this amount, it will produce a net benefit for society over time, but if not, a net social welfare loss is likely to result. A benefit-cost analysis of this nature would be an important tool for policymakers, as it condenses a great deal of complex information into monetary terms, making it relatively simple to communicate to policymakers and stakeholders (<xref ref-type="bibr" rid="B72">Waage et al. 2005</xref>). However, this must be accompanied by a technical feasibility study that assesses logistical constraints involved in applying sustained pressure to wasp populations given their high mobility, high reproductive rate, and ability to exploit a range of food resources (<xref ref-type="bibr" rid="B68">van Zyl et al. 2018</xref>).</p>
      <p>The simplifying assumption was used in the model that the supply of managed hives remains constant over time despite a rising demand for pollination services. This implies that no new commercial beekeepers will enter the market, even as hive hire prices rise, to increase supply and place downward pressure on prices. As a result, the model predicts the average cost per honeybee hive supplied for pollination to rise from approximately US$75.00/hive to US$78.20–84.50/hive over the next several decades if <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> continues to spread (Fig. <xref ref-type="fig" rid="F3">3</xref>). The assumptions reflect the barriers new suppliers face when entering the market. South Africa has banned the importation of live honeybees since the early 1980s (<xref ref-type="bibr" rid="B22">Dietemann et al. 2009</xref>), and the availability of wild bee stock to build or replenish commercial stock is limited (<xref ref-type="bibr" rid="B70">Veldtman 2018</xref>; <xref ref-type="bibr" rid="B4">Allsopp 2020</xref>). Access to year-round forage resources in South Africa’s dry climate is also limited and subject to seasonal variations that could affect the ability of the commercial pollination industry to expand (<xref ref-type="bibr" rid="B70">Veldtman 2018</xref>). Concerted conservation efforts specifically oriented toward plant species that benefit honeybees are needed to provide sufficient forage for wild and managed colonies (<xref ref-type="bibr" rid="B42">Mensah et al. 2017</xref>).</p>
      <p>If the set of assumptions used to characterize <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> spread is correct, or even a reasonable approximation, impacted sectors are likely to begin experiencing substantial costs after 20 years due to the logistic growth assumed in the model. This implies a significant lag period given that the pest has been present in the Western Cape province for many decades. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">Vespula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> has taken 50 years to spread from the southern slopes of Table Mountain after crossing a major geographical barrier, the Cape Flats (a large, low-lying region to the southeast of Cape Town), through human-mediated spread 25 years ago. However, since then, spread has been somewhat sporadic. While it quickly proliferated in the Boland region, spread appeared to slow after 2005 before increasing again from 2014, again due to human-mediated spread. This suggests that the increase in area occupied might not be occurring as uniformly as the model predicts, at least in the short term, although the total area affected remains relatively small. The complicated spread pattern observed indicates that a spatial time-step analysis would be beneficial, which the model does not currently include. This would be particularly useful when predicting regional and subregional spread and the economic implications for local agricultural industries and communities.</p>
      <p>The lack of a geospatial dimension in the model could be addressed in future work using modeling techniques that delineate landscapes into spatial units, such as species dispersal models or cellular automata models. Dispersal models incorporating the processes regulating species survival, reproduction, and movement have been used effectively to explain the spread of invasive species over large spatial and temporal scales (<xref ref-type="bibr" rid="B49">Pitt et al. 2009</xref>; <xref ref-type="bibr" rid="B78">Wittmann et al. 2016</xref>; <xref ref-type="bibr" rid="B39">Lustig et al. 2017</xref>). However, these have yet to be combined with economic models to inform policy and can be problematic when used for prediction (<xref ref-type="bibr" rid="B38">Liu et al. 2020</xref>). Cellular automata models have been effective policy support tools for invasive species preparedness and response efforts when combined with measures of economic impact (<xref ref-type="bibr" rid="B16">Cook et al. 2025</xref>) but may be unsuited to diverse landscapes where movement probability varies according to exogenous factors such as wind or human-mediated dispersal (<xref ref-type="bibr" rid="B65">Thompson et al. 2021</xref>). Given problems with more complex approaches, the aspatial model presented in this analysis is an important first step in informing policymakers of the potential economic threat posed by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic>.</p>
    </sec>
    <sec sec-type="Conclusion" id="sec10">
      <title>Conclusion</title>
      <p>An estimate of the damage likely to be caused by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> if it is allowed to spread in South Africa over the next 50 years has been provided in this paper. By applying a spread model to multiple sectors, it was shown how costs could increase at an increasing rate over time. The effects of wasps preying on foraging honeybees could raise variable production costs and yield losses for horticulture industries, and the value of damages was estimated to be US$30–55 million over 50 years. Cost increases to the wine grape industry from wasp control measures to prevent feeding on grapes were shown to be relatively minor, with costs estimated at US$1–3 million over 50 years. The financial impacts on households were shown to be high, with the predicted increase in pest extermination services in urban areas likely to cost US$110–260 million. These results suggest that the combined costs to these three sectors could be US$160–300 million after 50 years, implying annual costs of US$3–6 million. This implies that if a response was mounted by the South African government to mitigate the spread of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic> at a cost of up to US$3 million per year, this would most likely produce a net benefit for society. Further research is needed to expand on the simulation model introduced in this paper. More scientific investigation into the spread of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vespula">V.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="germanica">germanica</tp:taxon-name-part></tp:taxon-name></italic>, particularly Hap1, would be extremely helpful to improve the model, given that results were highly sensitive to parameters that affected the spread rate. Introducing a spatial dimension to explore interactions between different habitats and spreading wasp populations may help in the design of future responses at a regional and subregional scale.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>The authors thank Marc Widmer (Western Australian Department of Primary Industries and Regional Development) and Dr. Francois (Gulu) Bekker (Stellenbosch University) for assistance generously provided. The authors have no support to report.</p>
    </ack>
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    <sec sec-type="Additional information" id="sec11">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec12">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec13">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec14">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.</p>
        <p>No AI tools were used in the preparation of this manuscript.</p>
      </sec>
      <sec sec-type="Funding" id="sec15">
        <title>Funding</title>
        <p>The authors have no funding to report. Publication charges were generously provided by the Harry Butler Institute, Murdoch University.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec16">
        <title>Author contributions</title>
        <p>David C. Cook: conceptualization, methodology, formal analysis, investigation, writing – original draft, writing – review and editing, visualization. Simon J. McKirdy: conceptualization, methodology, writing – original draft, writing – review and editing, funding acquisition. Ruan Veldtman: conceptualization, methodology, data curation, validation, writing – original draft, writing – review and editing, funding acquisition.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec17">
        <title>Author ORCIDs</title>
        <p>D.C. Cook <ext-link xlink:href="https://orcid.org/0000-0002-2954-7697" ext-link-type="uri">https://orcid.org/0000-0002-2954-7697</ext-link></p>
        <p>S.J. McKirdy <ext-link xlink:href="https://orcid.org/0000-0001-6578-5521" ext-link-type="uri">https://orcid.org/0000-0001-6578-5521</ext-link></p>
        <p>R. Veldtman <ext-link xlink:href="https://orcid.org/0000-0002-2258-6108" ext-link-type="uri">https://orcid.org/0000-0002-2258-6108</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec18">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text.</p>
      </sec>
    </sec>
  </back>
</article>
