<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3-mathml3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" article-type="research-article" dtd-version="1.3" xml:lang="EN">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2026.1746978</article-id>
<article-version article-version-type="Version of Record" vocab="NISO-RP-8-2008"/>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Original Research</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>The invasive bark beetle, <italic>Pagiocerus frontalis</italic> (Fabricius): as an emerging maize storage pest in Tanzania</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Chidege</surname><given-names>Maneno Y.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>*</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/1398445/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="visualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/visualization/">Visualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="validation" vocab-term-identifier="https://credit.niso.org/contributor-roles/validation/">Validation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="software" vocab-term-identifier="https://credit.niso.org/contributor-roles/software/">Software</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="conceptualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
</contrib>
<contrib contrib-type="author">
<name><surname>Venkataramana</surname><given-names>Pavithravani B.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="conceptualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="visualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/visualization/">Visualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="validation" vocab-term-identifier="https://credit.niso.org/contributor-roles/validation/">Validation</role>
</contrib>
<contrib contrib-type="author">
<name><surname>Ndakidemi</surname><given-names>Patrick A.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/615624/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="visualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/visualization/">Visualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="validation" vocab-term-identifier="https://credit.niso.org/contributor-roles/validation/">Validation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="conceptualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
</contrib>
</contrib-group>
<aff id="aff1"><label>1</label><institution>Departiment of Life Sciences and Bioengeneering, The Nelson Mandela African Institution of Science and Technology (NM-AIST)</institution>, <city>Arusha</city>,&#xa0;<country country="tz">Tanzania</country></aff>
<aff id="aff2"><label>2</label><institution>Depertiment of Plant Health and Biosecurity, Tanzania Plant Health and Pesticides Authority (TPHPA)</institution>, <city>Arusha</city>,&#xa0;<country country="tz">Tanzania</country></aff>
<author-notes>
<corresp id="c001"><label>*</label>Correspondence: Maneno Y. Chidege, <email xlink:href="mailto:chidegem@nm-aist.ac.tz">chidegem@nm-aist.ac.tz</email>; <email xlink:href="mailto:mchidege@yahoo.com">mchidege@yahoo.com</email></corresp>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2026-02-09">
<day>09</day>
<month>02</month>
<year>2026</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2026</year>
</pub-date>
<volume>17</volume>
<elocation-id>1746978</elocation-id>
<history>
<date date-type="received">
<day>17</day>
<month>11</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>08</day>
<month>01</month>
<year>2026</year>
</date>
<date date-type="rev-recd">
<day>29</day>
<month>12</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2026 Chidege, Venkataramana and Ndakidemi.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Chidege, Venkataramana and Ndakidemi</copyright-holder>
<license>
<ali:license_ref start_date="2026-02-09">https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
<license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License (CC BY)</ext-link>. The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</license-p>
</license>
</permissions>
<abstract>
<p>In Tanzania, smallholder farmers often sell maize immediately after harvest to avoid post-harvest losses caused by storage pests, a practice that exacerbates food insecurity. The invasive bark beetle <italic>Pagiocerus frontalis</italic> (Fabricius, 1801) (Coleoptera: Curculionidae), which infests maize and avocado seeds, was first detected in Tanzania in December 2018 in stored provitamin A yellow maize (CP 201). Host plant resistance represents a safe and sustainable strategy for managing storage insect pests. In this study, we evaluated the susceptibility of 27 maize varieties commonly cultivated in Tanzania. The varieties were assessed for grain damage, weight loss, progeny production, time to progeny emergence, and adult insect mortality. Significant differences were observed in grain damage and adult mortality, whereas no significant differences were detected in grain weight loss, progeny number, or time to progeny emergence. These findings demonstrate that <italic>P. frontalis</italic> can inflict substantial damage across all major maize varieties in Tanzania. This study provides the first evidence of varietal susceptibility to this invasive pest and establishes a foundation for developing integrated pest management strategies aimed at safeguarding maize production and enhancing food security in Tanzania and across Africa.</p>
</abstract>
<kwd-group>
<kwd>food insecurity</kwd>
<kwd>invasive</kwd>
<kwd>maize varieties</kwd>
<kwd>new storage insect pest</kwd>
<kwd>provitamin-A</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declared that financial support was not received for this work and/or its publication.</funding-statement>
</funding-group>
<counts>
<fig-count count="11"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="46"/>
<page-count count="13"/>
<word-count count="4780"/>
</counts>
<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Plant Pathogen Interactions</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Maize (<italic>Zea mays</italic> L.) is Tanzania&#x2019;s most important staple crop, underpinning household food security, nutrition, and rural livelihoods (<xref ref-type="bibr" rid="B34">Mutungi et al., 2019</xref>; <xref ref-type="bibr" rid="B35">Mutungi et al., 2022</xref>). It accounts for over 70% of national cereal production and serves as a primary source of calories for millions of Tanzanians (<xref ref-type="bibr" rid="B32">Mmasa and Mathur, 2020</xref>). Beyond consumption, maize supports income generation and national grain markets, and it plays a key role in agricultural development strategies, food security frameworks, and climate resilience agendas. Ensuring the availability and quality of maize after harvest is therefore both a production challenge and a national policy priority, closely aligned with poverty reduction, nutrition security, and the sustainable development goals.</p>
<p>Despite increased maize productivity, post-harvest losses remain a major constraint, particularly in smallholder systems where conventional storage methods, such as polypropylene bags, cribs, and unimproved granaries dominate (<xref ref-type="bibr" rid="B2">Abass et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B26">Kumar and Kalita, 2017</xref>). Storage insect pests are a leading cause of these losses (<xref ref-type="bibr" rid="B33">Mutambuki and Likhayo, 2021</xref>; <xref ref-type="bibr" rid="B39">Nwosu, 2018</xref>). In Tanzania, the maize weevil <italic>Sitophilus zeamais</italic> (Motschulsky) (Coleoptera: Curculionidae) can reduce grain quantity by 20&#x2013;30% within three to six months of storage, while the larger grain borer <italic>Prostephanus truncatus</italic>&#x2002;(horn) (Coleoptera:&#x2002;Bostrichidae), an invasive species introduced into East Africa in the late 1970s, often causes losses exceeding 30&#x2013;40% (<xref ref-type="bibr" rid="B13">Dunstan and Magazini, 1981</xref>; <xref ref-type="bibr" rid="B21">Hodges et&#xa0;al., 1983</xref>; <xref ref-type="bibr" rid="B20">Hodges, 1986</xref>; <xref ref-type="bibr" rid="B17">Haines, 1991</xref>; <xref ref-type="bibr" rid="B38">Njoroge et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B3">Addo, 2002</xref>). Such damage also compromises grain quality, seed viability, and market value (<xref ref-type="bibr" rid="B31">Mesterha&#x301;&#x17a;y et al., 2020</xref>).</p>
<p>The accidental introduction of <italic>P. truncatus</italic> illustrated how invasive storage pests can rapidly spread and overwhelm local management systems (<xref ref-type="bibr" rid="B29">Markham et al., 1994</xref>). In a similar concern, the invasive bark beetle <italic>P. frontalis</italic>, native to Central and South America, has been associated primarily with avocado (<italic>Persea americana</italic> Mill.) seeds since its first records in the 1930s (<xref ref-type="bibr" rid="B24">Kirkendall, 2018</xref>). The insect bores into partially or fully exposed seeds on the ground but does not attack fruits on the tree (<xref ref-type="bibr" rid="B10">De Dios &#xc1;vila et&#xa0;al., 2021</xref>). It has also been recorded in coffee berries in Ecuador and can be laboratory-reared on cassava chips as well as maize grains (<xref ref-type="bibr" rid="B16">Gianoli et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B46">Yust, 1957</xref>; <xref ref-type="bibr" rid="B40">Okello et&#xa0;al., 1996</xref>). <italic>Pagiocerus frontalis</italic> can cause up to 44% yield loss in maize (<xref ref-type="bibr" rid="B10">De Dios &#xc1;vila et&#xa0;al., 2021</xref>), infesting cobs in the field before harvest and continuing to feed during storage (<xref ref-type="bibr" rid="B40">Okello et&#xa0;al., 1996</xref>).</p>
<p>The bark beetle <italic>Pagiocerus frontalis</italic> was first detected in Tanzania in December 2018, infesting stored provitamin A maize (CP 201) in Usariver, Arusha (latitude 3&#xb0;21&#x2032;45&#x2033; S, longitude 36&#xb0;52&#x2032;20&#x2033; E; altitude 1, 300&#x2013;1, 500 meters above sea level). Preliminary laboratory rearing confirmed it feeds on both maize and avocado seeds and can perforate polypropylene storage bags, causing damage comparable to <italic>P. truncatus</italic> (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>). These observations indicate a high risk of rapid spread through household storage and informal grain trade networks.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Adult <italic>P. frontalis</italic><bold>(A)</bold>, bored holes in woven polypropylene bags and spilled maize flour on the floor (between red dots) <bold>(B)</bold> .Severe damage caused by <italic>P. frontalis</italic>: maize grains with exhausted endosperm and visible maize flour <bold>(C-E)</bold>; avocado seeds with extensive perforation <bold>(F, G)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g001.tif">
<alt-text content-type="machine-generated">Photographs showing insect infestation and damage. Panel A: adult beetle. Panel B: infested sack with powdery residue. Panel C: food substrate with dark spots and residue. Panel D: damaged grains with perforations, larvae, eggs, and pupae. Panel E: damaged yellow maize kernels with black specks. Panel F: damaged avocado seeds with surface discoloration. Panel G: severely damaged avocado seed with multiple holes.</alt-text>
</graphic></fig>
<p>Current storage pest management in smallholder systems relies heavily on synthetic insecticides. While these can suppress pest populations, their effectiveness is limited by high costs, improper application, restricted access, and emerging resistance. Furthermore, pesticide use raises serious concerns about food safety, human health, and environmental contamination (<xref ref-type="bibr" rid="B2">Abass et&#xa0;al., 2014</xref>). Hermetic storage technologies, particularly Purdue Improved Crop Storage (PICS) bags, have demonstrated effectiveness against several storage pests by restricting oxygen and suppressing insect development (<xref ref-type="bibr" rid="B7">Baoua et&#xa0;al., 2014</xref>). However, adoption is uneven, and such technologies may not fully prevent damage by highly aggressive tunneling species, highlighting the need for complementary strategies such as host plant resistance.</p>
<p>The emergence of <italic>P. frontalis</italic> has policy and regional implications. Increased post-harvest losses threaten food availability, exacerbate price volatility, and may heighten dependence on emergency grain imports. Its regional spread could also complicate cross-border grain trade and phytosanitary regulation within East Africa. Effective early-stage management is therefore critical to prevent <italic>P. frontalis</italic> from becoming a region-wide storage pest with significant economic and food security consequences.</p>
<p>Host plant resistance is a cornerstone of sustainable integrated pest management (IPM) and aligns with policy goals related to climate-smart agriculture, environmental protection, and smallholder resilience. Resistant maize varieties offer a low-cost, scalable, and farmer-accessible option for reducing storage losses without additional chemical inputs (<xref ref-type="bibr" rid="B27">L&#xf3;pez-Castillo et&#xa0;al., 2018</xref>). While substantial research has addressed resistance to established storage pests such as <italic>S. zeamais</italic> and <italic>P. truncatus</italic>, resistance to <italic>P. frontalis</italic> remains poorly understood, particularly within Tanzanian widely cultivated maize varieties.</p>
<p>This study presents the first assessment of susceptibility among widely cultivated maize varieties in Tanzania to the newly detected storage pest <italic>P. frontalis</italic>. By generating baseline data to inform varietal selection, breeding programs, and national post-harvest loss reduction strategies, the findings contribute to early-stage management of this emerging pest. Beyond Tanzania, the results have broader regional relevance for invasive storage pest preparedness in East Africa and support global efforts to promote sustainable, pesticide-reducing approaches to post-harvest pest management.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<p>The invasive bark beetle <italic>P. frontalis</italic> was obtained from stored provitamin A maize (CP 201) grains harvested in December 2018 in Usa River, Arusha Region, Tanzania (latitude 3&#xb0;21&#x2032;45&#x2033; S, longitude 36&#xb0;52&#x2032;20&#x2033; E; altitude 1, 300&#x2013;1, 500 m above sea level). A colony was established and maintained at the Postharvest Entomology Laboratory of the Tanzania Plant Health and Pesticide Authority (TPHPA). Mixed-sex insects were reared in 2-kg glass jars containing provitamin A maize for 60 days under controlled conditions (23 &#xb1; 5&#xb0;C, 65 &#xb1; 5% RH) and continuous darkness (<xref ref-type="bibr" rid="B44">Sakka et&#xa0;al., 2022</xref>) to produce sufficient numbers for experimentation. Adults emerging insects less than seven days old were collected for experimental assays to ensure physiological uniformity and consistent reproductive potential (<xref ref-type="bibr" rid="B38">Njoroge et&#xa0;al., 2019</xref>).</p>
<p>Clean, undamaged grains of 27 maize varieties were sourced from seed companies, farmers, and the Tanzania Agricultural Research Institute (TARI) (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>) and were disinfected by fumigation with aluminium phosphide tablets (56% a.i.) at a rate of 3 g m<sup>&#x2212;3</sup> in airtight plastic drums for 7 days, followed by aeration for 72 h to remove residual phosphine. Grain moisture content was adjusted to 12% (wet basis) and verified using the oven-dry method at 105 &#xb1; 2&#xb0;C for 24 h, in accordance with <xref ref-type="bibr" rid="B22">ISTA (2020)</xref>. To minimize within-variety variation, grains were standardized by counting and weighing, and only samples within &#xb1;5&#x2013;10% of the mean grain weight for each variety sample were used in the experiments; grains smaller or larger than this range were discarded (<xref ref-type="bibr" rid="B19">Harden and Wood, 2017</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Grains variety name and sources.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">S.N</th>
<th valign="middle" align="left">Variety name</th>
<th valign="middle" align="left">Source</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">1</td>
<td valign="middle" align="left">SC419</td>
<td valign="middle" align="left">Seed Co. Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">2</td>
<td valign="middle" align="left">SC403</td>
<td valign="middle" align="left">Seed Co. Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">3</td>
<td valign="middle" align="left">UH628</td>
<td valign="middle" align="left">Farmers-Mbeya</td>
</tr>
<tr>
<td valign="middle" align="left">4</td>
<td valign="middle" align="left">H625</td>
<td valign="middle" align="left">Seed Co. Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">5</td>
<td valign="middle" align="left">SC719</td>
<td valign="middle" align="left">Seed Co. Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">6</td>
<td valign="middle" align="left">Njano-CP201</td>
<td valign="middle" align="left">Farmers-Morogoro</td>
</tr>
<tr>
<td valign="middle" align="left">7</td>
<td valign="middle" align="left">SY6444</td>
<td valign="middle" align="left">Syngenta Tanzania ltd</td>
</tr>
<tr>
<td valign="middle" align="left">8</td>
<td valign="middle" align="left">DK777</td>
<td valign="middle" align="left">Monsanto tz ltd</td>
</tr>
<tr>
<td valign="middle" align="left">9</td>
<td valign="middle" align="left">PAN691</td>
<td valign="middle" align="left">Pannar Seeds Co. Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">10</td>
<td valign="middle" align="left">SC627</td>
<td valign="middle" align="left">Seed Co</td>
</tr>
<tr>
<td valign="middle" align="left">11</td>
<td valign="middle" align="left">DK8031</td>
<td valign="middle" align="left">Monsanto tz ltd</td>
</tr>
<tr>
<td valign="middle" align="left">12</td>
<td valign="middle" align="left">ZMS 606</td>
<td valign="middle" align="left">Bajuta International (T) Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">13</td>
<td valign="middle" align="left">Kitale628</td>
<td valign="middle" align="left">Farmers-Arusha</td>
</tr>
<tr>
<td valign="middle" align="left">14</td>
<td valign="middle" align="left">Situka-M1</td>
<td valign="middle" align="left">TARI</td>
</tr>
<tr>
<td valign="middle" align="left">15</td>
<td valign="middle" align="left">Pionner3253</td>
<td valign="middle" align="left">Farmers-Arusha</td>
</tr>
<tr>
<td valign="middle" align="left">16</td>
<td valign="middle" align="left">Staha</td>
<td valign="middle" align="left">Farmers-Morogoro</td>
</tr>
<tr>
<td valign="middle" align="left">17</td>
<td valign="middle" align="left">Aminika505</td>
<td valign="middle" align="left">Farmers-Arusha</td>
</tr>
<tr>
<td valign="middle" align="left">18</td>
<td valign="middle" align="left">Kaspidi</td>
<td valign="middle" align="left">IFFA Seed Company Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">19</td>
<td valign="middle" align="left">Kilima</td>
<td valign="middle" align="left">Farmers-Shinyanga</td>
</tr>
<tr>
<td valign="middle" align="left">20</td>
<td valign="middle" align="left">MeruHB515</td>
<td valign="middle" align="left">Meru Agro Tours &amp; Consultant Co. Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">21</td>
<td valign="middle" align="left">Lubango</td>
<td valign="middle" align="left">IFFA Seed Company Ltd</td>
</tr>
<tr>
<td valign="middle" align="left">22</td>
<td valign="middle" align="left">UH615</td>
<td valign="middle" align="left">Farmers-Mbeya</td>
</tr>
<tr>
<td valign="middle" align="left">23</td>
<td valign="middle" align="left">UHS5350</td>
<td valign="middle" align="left">Farmers-Mbeya</td>
</tr>
<tr>
<td valign="middle" align="left">24</td>
<td valign="middle" align="left">UH630</td>
<td valign="middle" align="left">Farmers-Mbeya</td>
</tr>
<tr>
<td valign="middle" align="left">25</td>
<td valign="middle" align="left">TMV1</td>
<td valign="middle" align="left">Farmers-Morogoro</td>
</tr>
<tr>
<td valign="middle" align="left">26</td>
<td valign="middle" align="left">TMV2</td>
<td valign="middle" align="left">Farmers-Arusha</td>
</tr>
<tr>
<td valign="middle" align="left">27</td>
<td valign="middle" align="left">Hybrid513</td>
<td valign="middle" align="left">Farmers-Arusha</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>The experiment was conducted using a completely randomized design with 27 maize varieties and four replicates, giving a total of 108 petri dishes. Each replicate consisted of 30 grains placed in 90 &#xd7; 15 mm transparent plastic Petri dishes, to which 20 mixed-sex insects (&lt;7 days old) were added using a fine brush after collection by sieving (1-mm mesh), assuming an approximate 1:1 sex ratio (<xref ref-type="bibr" rid="B42">Quellhorst et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B41">Panagiotakis et&#xa0;al., 2023</xref>). The dishes were randomly positioned on laboratory shelves at 23 &#xb1; 5&#xb0;C and 65 &#xb1; 5 % RH under continuous darkness (<xref ref-type="bibr" rid="B44">Sakka et&#xa0;al., 2022</xref>). Ten days after infestation, parental insects were removed, and the number of dead specimens as well as grain damage in each replicate were recorded (Mwololo, 2012; <xref ref-type="bibr" rid="B5">Altun&#xe7; et&#xa0;al., 2023</xref>). Beginning 14 days after infestation, the dishes were monitored daily for the emergence of F<sub>1</sub> progeny. Emerging F<sub>1</sub> adults were not removed during the incubation period in order to quantify cumulative grain weight loss (Mwololo, 2012). Grains were incubated for a total of 65 days, after which progeny production and grain weight loss were assessed (<xref ref-type="bibr" rid="B12">Dobie, 1974</xref>; <xref ref-type="bibr" rid="B45">Tefera et&#xa0;al., 2011</xref>).</p>
<sec id="s2_1">
<label>2.1</label>
<title>Data collection</title>
<p>Data were recorded on the following parameters: grain damage, grain weight loss, number of emerged progeny, days to first F<sub>1</sub> progeny emergence, and adult mortality. Grain damage and adult mortality were assessed 10 days after insect introduction, while the number of emerged progeny and grain weight loss were recorded 65 days after insect introduction (<xref ref-type="bibr" rid="B15">Gerken and Campbell, 2020</xref>; <xref ref-type="bibr" rid="B23">Kasozi et al., 2016</xref>). These parameters were recorded in accordance with the descriptions below.</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Grain damage</title>
<p>Grain damage was assessed by manually inspecting all grains in each experimental unit. Grains were visually classified into two categories: damaged grains, which are grains showing visible insect feeding holes, tunnels, or structural collapse caused by insect activity; and undamaged grains, which are grains with no visible signs of feeding or structural damage. Counting was performed using a standardized tally method to ensure consistency and reproducibility. Each grain was examined individually, and a mark was recorded on a tally sheet for every damaged or undamaged grain. Observations were conducted under consistent lighting conditions, and all counts were carried out by the same person to minimize observer bias. The percentage of damaged grains was then calculated using the formula described by <xref ref-type="bibr" rid="B37">Njoroge et&#xa0;al. (2014)</xref>:</p>
<p>Grain damage (%) = (Nd/(Nd + Nu)) &#xd7; 100, where Nd=number of damaged grains, Nu is the number of undamaged grains (<xref ref-type="bibr" rid="B37">Njoroge et&#xa0;al., 2014</xref>).</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Grain weight loss</title>
<p>Grains were separated into damaged and undamaged categories using a fine-mesh sieve to remove loose frass and debris. The frass and debris were collected in pre-weighed containers to avoid overestimation of grain weight. Each fraction (undamaged grains, damaged grains, and frass) was weighed using an analytical balance. Grains were then gently shaken through a 2-mm mesh sieve to ensure complete removal of frass without causing additional grain damage, and weights were recorded immediately after sieving. Grain weight loss was calculated as follows:</p>
<p>Grain weight loss (%) = ((Wu - Wi)/Wu) &#xd7; 100, where, Wu is the weight of uninfested grains; Wi is the weight of infested grains (<xref ref-type="bibr" rid="B14">Gad et al., 2024</xref>).</p>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Adult mortality</title>
<p>Adult mortality of <italic>P. frontalis</italic> was determined by carefully examining individual insects. Adults that showed no movement or response to gentle probing with a fine brush were classified as dead (Di), while all others were recorded as alive. To minimize error, mortality assessments were performed at a consistent time across treatments. Adult mortality (%) was then calculated using the formula:</p>
<p>Adult mortality (%) = (Di/Ti) &#xd7; 100, where Di is the number of dead adults and Ti is the total number of adults initially introduced (<xref ref-type="bibr" rid="B6">Athanassiou and Arthur, 2020</xref>).</p>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Progeny production</title>
<p>Progeny production was quantified by counting adult insects present in each experimental unit after 65 days. This measure reflects the reproductive performance and population growth potential of the species under the tested varieties. Progeny production was calculated as follows;</p>
<p>Progeny production=Total number of new adults at 65 days (<xref ref-type="bibr" rid="B5">Altun&#xe7; et&#xa0;al., 2023</xref>)</p>
</sec>
<sec id="s2_6">
<label>2.6</label>
<title>Days to progeny production</title>
<p>The minimum time required for progeny production was measured by recording the number of days from insect inoculation until the first emergence of new <italic>P. frontalis</italic> adults. To obtain this data, containers were inspected daily from 14 days at approximately the same time to ensure consistent observations. Any petridish with a newly emerged adults were carefully noted. For each replicate, the day on which the first new adult appeared was recorded, and the mean value across replicates was used to determine days to progeny production, defined as the average number of days to first progeny emergence (<xref ref-type="bibr" rid="B4">Ahmad et&#xa0;al., 2018</xref>).</p>
<p>Days to progeny production = Mean days to first emergence of new adults.</p>
</sec>
<sec id="s2_7">
<label>2.7</label>
<title>Data analysis</title>
<p>Data collected were tested for normality using the Shapiro&#x2013;Francia test (<xref ref-type="bibr" rid="B30">Mbah and Paothong, 2015</xref>), and homogeneity of variances was assessed using Levene&#x2019;s test (<xref ref-type="bibr" rid="B9">D&#x2019;Isita et&#xa0;al., 2024</xref>). After confirming that all data were normally distributed and that variances were homogeneous, a one-way analysis of variance (ANOVA) was performed. Mean values for parameters showing significant differences were separated using Tukey&#x2019;s honestly significant difference (HSD) <italic>post hoc</italic> test at the 5% significance level, using Statistica version 10 software.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Grain damage</title>
<p>A preliminary assessment under farmer storage conditions indicated that <italic>Pagiocerus frontalis</italic> caused a mean grain damage (GD) of 9.95% and grain weight loss (GWL) of 17.65% within two months after harvest. Laboratory observations showed that <italic>P. frontalis</italic> infested maize grains by boring into the grain, producing exit holes, and generating fine powdery frass. Infested grains exhibited partial to complete endosperm consumption.</p>
<p>Laboratory screening of maize varieties revealed a significant effect of variety on grain damage (F(26, 81) = 9.65, <italic>P</italic> &lt; 0.001) (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>). Although all 27 varieties sustained damage, the magnitude varied significantly among them. TMV1 and Hybrid 513 recorded the lowest mean grain damage at 42.5% and 45.0%, respectively, whereas Staha and Pioneer 3253 showed the highest damage levels at 90.83% and 92.5%, respectively. In varieties with higher damage, most grains exhibited extensive perforation and endosperm loss. Grain damage was detectable as early as 10 days after infestation. No variety remained undamaged under the experimental conditions (<xref ref-type="fig" rid="f2"><bold>Figures&#xa0;2</bold></xref>, <xref ref-type="fig" rid="f3"><bold>3</bold></xref>).</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>One-way ANOVA showing the effect of maize variety on grain damage, insect performance, and grain weight loss caused by <italic>Pagiocerus frontalis</italic>.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Variable</th>
<th valign="middle" align="center">df (treatment, error)</th>
<th valign="middle" align="center">F</th>
<th valign="middle" align="center">P</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Grain damage (%)</td>
<td valign="middle" align="left">26, 81</td>
<td valign="middle" align="right">9.65</td>
<td valign="middle" align="right">&lt;0.001</td>
</tr>
<tr>
<td valign="middle" align="left">Adult mortality (%)</td>
<td valign="middle" align="left">26, 81</td>
<td valign="middle" align="right">3.39</td>
<td valign="middle" align="right">&lt;0.001</td>
</tr>
<tr>
<td valign="middle" align="left">Days to first F<sub>1</sub> emergence</td>
<td valign="middle" align="left">26, 81</td>
<td valign="middle" align="right">1.17</td>
<td valign="middle" align="right">0.294</td>
</tr>
<tr>
<td valign="middle" align="left">Number of progenies</td>
<td valign="middle" align="left">26, 81</td>
<td valign="middle" align="right">1.04</td>
<td valign="middle" align="right">0.428</td>
</tr>
<tr>
<td valign="middle" align="left">Grain weight loss (%)</td>
<td valign="middle" align="left">26, 81</td>
<td valign="middle" align="right">0.62</td>
<td valign="middle" align="right">0.915</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>One-way analysis of variance (ANOVA) was used to test for varietal effects. Significant differences were declared at <italic>P</italic> &lt; 0.05.</p></fn>
</table-wrap-foot>
</table-wrap>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Mean grain damage (%) caused by <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (first set of varieties). Bars represent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varieties sharing the same letter are not significantly different. Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g002.tif">
<alt-text content-type="machine-generated">Bar chart showing mean percent grain damage among maize varieties, with Tembo-SC419 showing the lowest damage and Pioneer 3253 the highest. Error bars indicate variability, and letters denote significant differences.</alt-text>
</graphic></fig>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Mean grain damage (%) caused by <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (second set of maize varieties). Barsrepresent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varieties sharing the same letter are not significantly different. Sample size was n = 4. for readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g003.tif">
<alt-text content-type="machine-generated">Bar chart of mean percent grain damage across maize varieties. Staha and UH615 show the highest damage, while TMV1 and UH630 show the lowest. Error bars and letters indicate significant differences. </alt-text>
</graphic></fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Adult mortality</title>
<p>Adult mortality differed significantly among maize varieties (F(26, 81) = 3.39; P &lt; 0.001) (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>), although overall mortality levels were low. The lowest mortality rates were observed in DK8031 (13.75%), Staha (21.25%), and Hybrid 513 (29.5%), while other varieties exhibited intermediate mortality levels (<xref ref-type="fig" rid="f4"><bold>Figures&#xa0;4</bold></xref>, <xref ref-type="fig" rid="f5"><bold>5</bold></xref>). Differences in adult mortality did not consistently correspond with patterns of grain damage.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Mean mortality (%) of <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (first set of varieties). Bars represent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varieties sharing the same letter are not significantly different. Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g004.tif">
<alt-text content-type="machine-generated">Bar chart showing mean percent mortality (&#xb1; standard error) for maize varieties, with values ranging from approximately 18% to 28%.</alt-text>
</graphic></fig>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Mean mortality (%) of <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (second set of maize varieties). Bars represent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varieties sharing the sameletter are not significantly different. Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g005.tif">
<alt-text content-type="machine-generated">Bar chart showing mean percent mortality (&#xb1; standard error) for maize varieties. Hybrid 513 shows the highest mortality and DK8031 the lowest. Letters indicate significant differences.</alt-text>
</graphic></fig>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Progeny production</title>
<p>No significant differences were detected among maize varieties in mean time to first F<sub>1</sub> adult emergence (F(26, 81) = 1.17; P = 0.294) or in the mean number of F<sub>1</sub> progeny produced (F(26, 81) = 1.04; P = 0.428) (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>). Time to first adult emergence ranged from 20 to 23 days across varieties (<xref ref-type="fig" rid="f6"><bold>Figures&#xa0;6</bold></xref>, <xref ref-type="fig" rid="f7"><bold>7</bold></xref>), while progeny production varied from 6.75 to 10.25 adults per sample (<xref ref-type="fig" rid="f8"><bold>Figures&#xa0;8</bold></xref>, <xref ref-type="fig" rid="f9"><bold>9</bold></xref>).</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Mean number of days to first F1 progeny of <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (first set of varieties). Barsrepresent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varietiessharing the same letter are not significantly different. Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g006.tif">
<alt-text content-type="machine-generated">Bar chart showing mean number of days to F1 emergence for maize varieties, with error bars. Values range from approximately 21 to 23 days.</alt-text>
</graphic></fig>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>Mean number of days to first F1 progeny of <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (second set of maize varieties). Bars represent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varieties sharing the same letter are not significantly different. Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g007.tif">
<alt-text content-type="machine-generated">Bar chart showing mean number of days to F1 emergence for maize varieties. TMV2 shows the shortest duration, and UH630 the longest.</alt-text>
</graphic></fig>
<fig id="f8" position="float">
<label>Figure&#xa0;8</label>
<caption>
<p>Mean number of progeny of <italic>Pagiocerus frontalis</italic> emerging on 27 maize varieties under laboratory conditions (first set of varieties). Bars represent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varieties sharing the same letter are not significantly different. Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g008.tif">
<alt-text content-type="machine-generated">Bar chart showing mean number of progeny for maize varieties, with error bars. UH615 and UHS5350 show the lowest means, while UH630 and SY6444 show the highest. </alt-text>
</graphic></fig>
<fig id="f9" position="float">
<label>Figure&#xa0;9</label>
<caption>
<p>Mean number of progeny of <italic>Pagiocerus frontalis</italic> emerging on 27 maize varieties under laboratory conditions (second set of maize varieties). Bars represent means &#xb1; SE. Different letters above bars indicate significant differences among varieties according to Tukey&#x2019;s HSD test (P &#x2264; 0.05); varieties sharing the same letter are not significantly different. Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g009.tif">
<alt-text content-type="machine-generated">Bar chart showing mean number of progeny for maize varieties, with error bars. Hybrid 513, Lubango, and Njano-CP201 show the highest means. </alt-text>
</graphic></fig>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Grain weight loss</title>
<p>Grain weight loss did not differ significantly among maize varieties (F(26, 81) = 0.62; P = 0.915) (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>). Mean weight loss ranged from 16.69% in TMV1 to 24.43% in SY6444. Although some varieties exhibited lower grain damage or higher adult mortality, these differences did not translate into significant variation in cumulative grain weight loss (<xref ref-type="fig" rid="f10"><bold>Figures&#xa0;10</bold></xref>, <xref ref-type="fig" rid="f11"><bold>11</bold></xref>).</p>
<fig id="f10" position="float">
<label>Figure&#xa0;10</label>
<caption>
<p>Mean grain weight loss (%) caused by <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (first set of varieties). Bars represent means &#xb1; SE. No significant differences were observed among varieties (P &#x2264; 0.05). Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g010.tif">
<alt-text content-type="machine-generated">Bar chart showing mean percent grain weight loss for maize varieties, with error bars. SY6444 shows the highest loss, while Tumbili-SC403, SC719, Zamseed 606, and Sutka show lower losses.</alt-text>
</graphic></fig>
<fig id="f11" position="float">
<label>Figure&#xa0;11</label>
<caption>
<p>Mean grain weight loss (%) caused by <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (second set of maize varieties). Bars represent means &#xb1; SE. No significant differences were observed among varieties (P &#x2264; 0.05). Sample size was n = 4. Mean grain weight loss (%) caused by <italic>Pagiocerus frontalis</italic> on 27 maize varieties under laboratory conditions (second set of maize varieties). Bars represent means &#xb1; SE. No significant differences were observed among varieties (P &#x2264; 0.05). Sample size was n = 4. For readability, a complete table of means is provided in <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Table&#xa0;3</bold></xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1746978-g011.tif">
<alt-text content-type="machine-generated">Bar chart showing mean percent grain weight loss for maize varieties, with error bars. TMV1 and UH630 show the highest losses, while UH615 and MeruHB515 show the lowest.</alt-text>
</graphic></fig>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>This study showed that <italic>Pagiocerus frontalis</italic> can cause substantial post-harvest damage across widely cultivated maize varieties in Tanzania, highlighting its potential as an emerging storage pest. Variation in grain damage among varieties suggests differences in susceptibility, consistent with previous studies showing that traits such as grain hardness, pericarp thickness, endosperm composition, amylose content, and secondary metabolite profiles can influence pest performance (<xref ref-type="bibr" rid="B36">Ngom et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B9">D&#x2019;Isita et&#xa0;al., 2024</xref>).</p>
<p>TMV1 and Hybrid 513 exhibited lower grain damage than Staha and Pioneer 3253, indicating relative differences in susceptibility among varieties. Similar patterns have been observed for other storage pests, such as <italic>Sitophilus zeamais</italic>, where certain maize genotypes consistently show reduced progeny emergence, grain damage, and weight loss (<xref ref-type="bibr" rid="B28">Marcos et&#xa0;al., 2025</xref>). These differences have been attributed to grain physical and chemical characteristics. Although <italic>P. frontalis</italic> and <italic>S. zeamais</italic> differ biologically, the results support the general principle that maize genotype can influence pest performance in controlled assays.</p>
<p>Adult mortality also varied among varieties, though overall levels were low and did not consistently correspond with patterns of grain damage. While the mechanisms were not directly investigated in this study, previous research suggests that varietal traits, such as secondary metabolites, may affect adult feeding efficiency or survival (<xref ref-type="bibr" rid="B8">Carneiro et&#xa0;al., 2025</xref>). Further mechanistic studies, including biochemical analyses or choice tests, are required to confirm these effects.</p>
<p>No significant differences were detected among maize varieties in progeny production or development time, indicating that once oviposition occurred, larval and pupal development proceeded similarly across varieties. Similar findings have been reported for other storage pests, such as <italic>Sitophilus zeamais</italic>, where varietal differences in progeny emergence and development were often minimal once eggs were laid, suggesting that post-oviposition development may be less influenced by maize genotype (<xref ref-type="bibr" rid="B12">Dobie, 1974</xref>; <xref ref-type="bibr" rid="B18">Haines, 1990</xref>). These results suggest that observed differences in overall susceptibility are more likely driven by factors affecting adult feeding and oviposition rather than larval performance.</p>
<p>Despite significant differences in grain damage among maize varieties, cumulative grain weight loss did not differ significantly. Similar trends have been reported in studies on stored-maize pests such as <italic>Sitophilus zeamais</italic>, where differences in kernel damage among varieties were not always accompanied by proportional differences in total weight loss over time (<xref ref-type="bibr" rid="B25">Kossou et&#xa0;al., 1993</xref>; <xref ref-type="bibr" rid="B45">Tefera et&#xa0;al., 2011</xref>). This pattern has been attributed to continued feeding and reproduction by insects that successfully establish on the grain, which can compensate for initial reductions in feeding on less susceptible varieties. Factors such as the duration of adult exposure, the ability of <italic>Pagiocerus frontalis</italic> to sustain feeding after establishment, and the generally limited physical or biochemical defenses of stored grain likely contributed to this outcome. These findings suggest that although some maize varieties may exhibit reduced susceptibility under controlled conditions, host-plant traits alone may not be sufficient to substantially limit overall quantitative losses during storage, highlighting the importance of integrated pest management strategies.</p>
<p>The close similarity between <italic>P. frontalis</italic> and <italic>P. truncatus</italic> in damage patterns, characterized by extensive endosperm consumption, multiple exit holes, and perforation of storage bags raises concerns for maize storage systems in Africa. The historical spread of <italic>P. truncatus</italic> following its introduction in the late 1970s and the severe losses that followed (Hodges et&#xa0;al., 1996; <xref ref-type="bibr" rid="B43">Quellhorst et&#xa0;al., 2024</xref>) highlight the urgency of proactive management of <italic>P. frontalis</italic>.</p>
<p>From an integrated pest management (IPM) perspective, these findings indicate that host-plant traits alone are unlikely to provide adequate control of <italic>Pagiocerus frontalis</italic>. Similar conclusions have been reported for other major stored-grain pests, including <italic>Sitophilus zeamais</italic> and <italic>Prostephanus truncatus</italic>, where varietal resistance reduced early damage but did not consistently prevent population buildup during storage (<xref ref-type="bibr" rid="B25">Kossou et&#xa0;al., 1993</xref>; <xref ref-type="bibr" rid="B45">Tefera et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B1">Abass et&#xa0;al., 2018</xref>). Varieties showing preliminary signs of lower susceptibility, such as TMV1, may contribute to reduced initial damage; however, effective management typically relies on integrating multiple control measures. Studies have demonstrated that improved storage hygiene, hermetic storage technologies, and complementary biological or selective chemical controls can substantially reduce pest populations and grain losses when used together (<xref ref-type="bibr" rid="B11">De Groote et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B7">Baoua et&#xa0;al., 2014</xref>). Such integrated approaches are therefore likely to be necessary for managing <italic>P. frontalis</italic> under smallholder storage conditions.</p>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusion</title>
<p>This study provides the first comprehensive evaluation of Tanzanian maize varieties in response to infestation by the invasive bark beetle <italic>Pagiocerus frontalis</italic>. Although significant varietal differences were detected in grain damage and adult mortality, all tested varieties were susceptible, and none expressed resistance strong enough to disrupt progeny development or reduce cumulative grain weight loss.</p>
<p>The observed patterns indicate that resistance in the tested germplasm is partial and likely mediated through physical or biochemical grain traits that reduce feeding efficiency rather than directly suppress survival or reproduction. However, these effects were insufficient to limit population buildup or economic losses, confirming that host-plant resistance alone cannot provide effective control of <italic>P. frontalis</italic> under tropical storage conditions.</p>
<p>Effective management of <italic>P. frontalis</italic> will therefore require integrated pest management approaches that combine moderately tolerant varieties with improved storage infrastructure, particularly hermetic storage systems, alongside complementary biological or selective chemical controls. Strengthening early detection, containment, and surveillance efforts is essential to prevent widespread establishment. Future breeding programs should prioritize resistance-associated traits such as increased grain hardness and elevated phenolic content to support the development of durable resistance and safeguard maize-based food security in Tanzania and across sub-Saharan Africa.</p>
</sec>
<sec id="s6">
<label>6</label>
<title>Recommendations</title>
<p>Future research should expand resistance screening and breeding efforts by evaluating both local and improved maize germplasm for traits associated with reduced susceptibility to <italic>Pagiocerus frontalis</italic>, including grain hardness, pericarp thickness,&#xa0;and potential biochemical defenses such as phenolic compounds. Further studies should also investigate integrated management strategies under controlled and on-farm conditions, combining maize varieties showing relatively lower damage with improved storage technologies (e.g., hermetic systems) and complementary control options to better understand their combined effects on pest survival, reproduction, and grain loss. In addition, long-term ecological studies are needed to monitor the distribution, population dynamics, and competitive interactions of <italic>P. frontalis</italic> with other major storage pests. Research should also assess effective extension approaches for early detection and post-harvest handling practices that limit infestation and reinfestation. Finally, broader studies examining the role of <italic>P. frontalis</italic> within national post-harvest systems would help inform evidence-based strategies for strengthening maize storage resilience and food security.</p>
</sec>
</body>
<back>
<sec id="s7" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Material</bold></xref>. Further inquiries can be directed to the corresponding author.</p></sec>
<sec id="s8" sec-type="ethics-statement">
<title>Ethics statement</title>
<p>The manuscript presents research on animals that do not require ethical approval for their study.</p></sec>
<sec id="s9" sec-type="author-contributions">
<title>Author contributions</title>
<p>MC: Formal Analysis, Writing &#x2013; original draft, Data curation, Methodology, Visualization, Validation, Software, Investigation, Writing &#x2013; review &amp; editing, Conceptualization. PV: Methodology, Writing &#x2013; review &amp; editing, Investigation, Conceptualization, Writing &#x2013; original draft, Visualization, Validation. PN: Formal Analysis, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Visualization, Validation, Methodology, Investigation, Conceptualization.</p></sec>
<ack>
<title>Acknowledgments</title>
<p>The authors wish to thank the farmers, seed companies, Tanzania agricultural research institute (TARI) for providing maize grains for this study.</p>
</ack>
<sec id="s11" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p></sec>
<sec id="s12" sec-type="disclaimer">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p></sec>
<sec id="s13" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p></sec>
<sec id="s14" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fpls.2026.1746978/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fpls.2026.1746978/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Table1.docx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document"/></sec>
<ref-list>
<title>References</title>
<ref id="B1">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Abass</surname> <given-names>A. B.</given-names></name>
<name><surname>Fischler</surname> <given-names>M.</given-names></name>
<name><surname>Schneider</surname> <given-names>K.</given-names></name>
<name><surname>Daudi</surname> <given-names>S.</given-names></name>
<name><surname>Gaspar</surname> <given-names>A.</given-names></name>
<name><surname>R&#xfc;st</surname> <given-names>J.</given-names></name>
<etal/>
</person-group>. (<year>2018</year>). 
<article-title>On-farm comparison of different postharvest storage technologies in a maize farming system of Tanzania</article-title>. <source>J. Stored Products Res.</source> <volume>77</volume>, <fpage>55</fpage>&#x2013;<lpage>65</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2018.03.002</pub-id>
</mixed-citation>
</ref>
<ref id="B2">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Abass</surname> <given-names>A. B.</given-names></name>
<name><surname>Mgonja</surname> <given-names>M. A.</given-names></name>
<name><surname>Amri</surname> <given-names>E.</given-names></name>
<name><surname>Mvumi</surname> <given-names>B. M.</given-names></name>
</person-group> (<year>2014</year>). 
<article-title>Postharvest losses of maize in Tanzania: Causes and management strategies</article-title>. <source>J. Stored Products Res.</source> <volume>58</volume>, <fpage>18</fpage>&#x2013;<lpage>26</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2014.03.002</pub-id>
</mixed-citation>
</ref>
<ref id="B3">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Addo</surname> <given-names>S.</given-names></name>
<name><surname>Birkinshaw</surname> <given-names>L. A.</given-names></name>
<name><surname>Hodges</surname> <given-names>R. J.</given-names></name>
</person-group> (<year>2002</year>). 
<article-title>Ten years after the arrival in ghana of larger grain borer: farmers&#x2019; responses and adoption of ipm strategies</article-title>. <source>International J. Pest Management.</source> <volume>48</volume> (<issue>2</issue>), <fpage>315</fpage>&#x2013;<lpage>325</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1080/09670870210151670</pub-id>
</mixed-citation>
</ref>
<ref id="B4">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Ahmad</surname> <given-names>F.</given-names></name>
<name><surname>Srebrnik</surname> <given-names>L.</given-names></name>
<name><surname>Gr&#x10d;man</surname> <given-names>D.</given-names></name>
</person-group> (<year>2018</year>). 
<article-title>The effects of population densities and diet on <italic>Tribolium castaneum</italic> (Herbst) life parameters</article-title>. <source>J. Stored Products Res.</source> <volume>79</volume>, <fpage>60</fpage>&#x2013;<lpage>65</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2018.09.002</pub-id>
</mixed-citation>
</ref>
<ref id="B5">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Altun&#xe7;</surname> <given-names>Y. E.</given-names></name>
<name><surname>Agrafioti</surname> <given-names>P.</given-names></name>
<name><surname>Lampiri</surname> <given-names>E.</given-names></name>
<name><surname>G&#xfc;ncan</surname> <given-names>A.</given-names></name>
<name><surname>Tsialtas</surname> <given-names>I. T.</given-names></name>
<name><surname>Athanassiou</surname> <given-names>C. G.</given-names></name>
</person-group> (<year>2023</year>). 
<article-title>Population growth of <italic>Prostephanus truncatus</italic> and <italic>Sitophilus zeamais</italic> and infestation patterns in three maize hybrids</article-title>. <source>J. Stored Products Res.</source> <volume>101</volume>, <fpage>102091</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2023.102091</pub-id>
</mixed-citation>
</ref>
<ref id="B6">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Athanassiou</surname> <given-names>C. G.</given-names></name>
<name><surname>Arthur</surname> <given-names>F. H.</given-names></name>
</person-group> (<year>2020</year>). 
<article-title>Cool down&#x2013;warm up: Differential responses of stored product insects after gradual temperature changes</article-title>. <source>Insects</source> <volume>11</volume>, <elocation-id>158</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/insects11030158</pub-id>, PMID: <pub-id pub-id-type="pmid">32121521</pub-id>
</mixed-citation>
</ref>
<ref id="B7">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Baoua</surname> <given-names>I. B.</given-names></name>
<name><surname>Amadou</surname> <given-names>L.</given-names></name>
<name><surname>Nielsen</surname> <given-names>J.</given-names></name>
<name><surname>Murdock</surname> <given-names>L. L.</given-names></name>
</person-group> (<year>2014</year>). 
<article-title>Triple bagging maize to reduce postharvest losses caused by insects in Niger</article-title>. <source>J. Stored Products Res.</source> <volume>56</volume>, <fpage>13</fpage>&#x2013;<lpage>19</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2013.10.003</pub-id>
</mixed-citation>
</ref>
<ref id="B8">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Carneiro</surname> <given-names>Z. F.</given-names></name>
<name><surname>Serpa-Neto</surname> <given-names>E. L.</given-names></name>
<name><surname>Pereira</surname> <given-names>H. C.</given-names></name>
<name><surname>Andrade</surname> <given-names>G. S.</given-names></name>
<name><surname>Zanuncio</surname> <given-names>J. C.</given-names></name>
<name><surname>Contreiras-Rodrigues</surname> <given-names>A. P. D.</given-names></name>
<etal/>
</person-group>. (<year>2025</year>). 
<article-title>Corn physicochemical properties affect <italic>Sitophilus zeamais</italic> losses</article-title>. <source>Braz. J. Biol.</source> <volume>85</volume>, <fpage>e291868</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1590/1519-6984.291868</pub-id>, PMID: <pub-id pub-id-type="pmid">40834174</pub-id>
</mixed-citation>
</ref>
<ref id="B9">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>D&#x2019;Isita</surname> <given-names>I.</given-names></name>
<name><surname>Pistillo</surname> <given-names>O. M.</given-names></name>
<name><surname>Di Palma</surname> <given-names>A. M.</given-names></name>
<name><surname>De Vita</surname> <given-names>P.</given-names></name>
<name><surname>Germinara</surname> <given-names>G. S.</given-names></name>
</person-group> (<year>2024</year>). 
<article-title>Susceptibility of old and modern wheat genotypes to <italic>Sitophilus granarius</italic> (L.) and <italic>Rhyzopertha Dominica</italic> (F.)</article-title>. <source>J. Stored Products Res.</source> <volume>106</volume>, <fpage>102265</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2024.102265</pub-id>
</mixed-citation>
</ref>
<ref id="B10">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>De Dios &#xc1;vila</surname> <given-names>N.</given-names></name>
<name><surname>Velasco</surname> <given-names>C. R.</given-names></name>
<name><surname>Campos</surname> <given-names>J. C.</given-names></name>
<name><surname>Solorio</surname> <given-names>A. B.</given-names></name>
<name><surname>Miranda</surname> <given-names>J. L. D.</given-names></name>
<name><surname>Virgen</surname> <given-names>M. O. E.</given-names></name>
</person-group> (<year>2021</year>). 
<article-title><italic>Pagiocerus frontalis</italic> (Fabricius) (Coleoptera: Curculionidae: Scolytinae) associated with avocado <italic>Persea Americana</italic> Miller (Lauraceae) fruit in Nayarit, Mexico</article-title>. <source>Florida Entomologist</source> <volume>104</volume>, <fpage>320</fpage>&#x2013;<lpage>321</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1653/024.104.0409</pub-id>
</mixed-citation>
</ref>
<ref id="B11">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>De Groote</surname> <given-names>H.</given-names></name>
<name><surname>Kimenju</surname> <given-names>S. C.</given-names></name>
<name><surname>Likhayo</surname> <given-names>P.</given-names></name>
<name><surname>Kanampiu</surname> <given-names>F.</given-names></name>
<name><surname>Tefera</surname> <given-names>T.</given-names></name>
<name><surname>Hellin</surname> <given-names>J.</given-names></name>
</person-group> (<year>2013</year>). 
<article-title>Effectiveness of hermetic systems in controlling maize storage pests in Kenya</article-title>. <source>J. Stored Products Res.</source> <volume>53</volume>, <fpage>27</fpage>&#x2013;<lpage>36</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2013.01.005</pub-id>
</mixed-citation>
</ref>
<ref id="B12">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Dobie</surname> <given-names>P.</given-names></name>
</person-group> (<year>1974</year>). 
<article-title>The laboratory assessment of the inherent susceptibility of maize varieties to post-harvest infestation by <italic>Sitophilus zeamais</italic> (Coleoptera: Curculionidae)</article-title>. <source>J. Stored Products Res.</source> <volume>10</volume>, <fpage>183</fpage>&#x2013;<lpage>197</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/0022-474X(74)90016-0</pub-id>
</mixed-citation>
</ref>
<ref id="B13">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Dunstan</surname> <given-names>W.</given-names></name>
<name><surname>Magazini</surname> <given-names>I.</given-names></name>
</person-group> (<year>1981</year>). 
<article-title>Outbreaks and new records. Tanzania. The larger grain borer on stored products</article-title>. <source>FAO Plant Prot. Bull.</source> <volume>29</volume>, <fpage>81</fpage>.
</mixed-citation>
</ref>
<ref id="B14">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Gad</surname> <given-names>L.</given-names></name>
<name><surname>Mwololo</surname> <given-names>J.</given-names></name>
<name><surname>Quellhorst</surname> <given-names>R.</given-names></name>
</person-group> (<year>2024</year>). 
<article-title>Evaluation of grain damage and weight loss formulas for storage pests</article-title>. <source>J. Stored Products Res.</source> <volume>104</volume>, <elocation-id>102153</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2024.102153</pub-id>
</mixed-citation>
</ref>
<ref id="B15">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Gerken</surname> <given-names>A. R.</given-names></name>
<name><surname>Campbell</surname> <given-names>J. F.</given-names></name>
</person-group> (<year>2020</year>). 
<article-title>Oviposition and development of <italic>Tribolium castaneum</italic> Herbst (Coleoptera: Tenebrionidae) on different types of flour</article-title>. <source>Agronomy</source> <volume>10</volume>, <fpage>1593</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/agronomy10101593</pub-id>
</mixed-citation>
</ref>
<ref id="B16">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Gianoli</surname> <given-names>E.</given-names></name>
<name><surname>Salazar</surname> <given-names>M.</given-names></name>
<name><surname>Araya</surname> <given-names>J.</given-names></name>
</person-group> (<year>2006</year>). 
<article-title>Occurrence of <italic>Pagiocerus frontalis</italic> in coffee berries in Ecuador</article-title>. <source>J. Trop. Agric.</source> <volume>44</volume>, <fpage>101</fpage>&#x2013;<lpage>105</lpage>.
</mixed-citation>
</ref>
<ref id="B17">
<mixed-citation publication-type="book">
<person-group person-group-type="author">
<name><surname>Haines</surname> <given-names>C. P.</given-names></name>
</person-group> (Ed.) (<year>1991</year>). <source>Insects and arachnids of tropical stored products: Their biology and identification</source> (
<publisher-name>Natural Resources Institute. Food and Agriculture Organization of the United Nations</publisher-name>). Available online at: <uri xlink:href="http://www.fao.org/3/T1838E/T1838E1f.htm">http://www.fao.org/3/T1838E/T1838E1f.htm</uri> (Accessed <date-in-citation content-type="access-date">June 30, 2022</date-in-citation>).
</mixed-citation>
</ref>
<ref id="B18">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Haines</surname> <given-names>C. P.</given-names></name>
<name><surname>Rees</surname> <given-names>D. P.</given-names></name>
<name><surname>Ryder</surname> <given-names>K.</given-names></name>
<name><surname>Sistyanto</surname> <given-names>S.</given-names></name>
<name><surname>Cahyana</surname> <given-names>Y.</given-names></name>
</person-group> (<year>1990</year>). 
<article-title>Brown-rice bait-bags for monitoring insect pest populations in bag stacks of milled rice as an aid to pest control decision-making</article-title>. In <source>Proceedings of the 5th International Working Conference on Stored-Product Protection</source> (<publisher-loc>Bordeaux, France</publisher-loc>). pp. <page-range>1351&#x2013;1358</page-range>.
</mixed-citation>
</ref>
<ref id="B19">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Harden</surname> <given-names>S.</given-names></name>
<name><surname>Wood</surname> <given-names>J. A.</given-names></name>
</person-group> (<year>2017</year>). 
<article-title>A single parameter for within-sample uniformity of seed size in grain, with an emphasis on pulses</article-title>. <source>Cereal Chem.</source> <volume>94</volume>, <fpage>430</fpage>&#x2013;<lpage>436</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1094/CCHEM-04-16-0121-R</pub-id>
</mixed-citation>
</ref>
<ref id="B20">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Hodges</surname> <given-names>R.</given-names></name>
</person-group> (<year>1986</year>). 
<article-title>The biology and control of <italic>Prostephanus truncatus</italic> (Horn) (Coleoptera: Bostrichidae)&#x2014;A destructive storage pest with an increasing range</article-title>. <source>J.&#xa0;Stored Products Res.</source> <volume>22</volume>, <fpage>21</fpage>&#x2013;<lpage>14</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/0022-474X(86)90040-8</pub-id>
</mixed-citation>
</ref>
<ref id="B21">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Hodges</surname> <given-names>R. J.</given-names></name>
<name><surname>Bell</surname> <given-names>C. H.</given-names></name>
<name><surname>Throne</surname> <given-names>J. E.</given-names></name>
</person-group> (<year>1983</year>). 
<article-title>The biology and control of the larger grain borer (<italic>Prostephanus truncatus</italic>)</article-title>. <source>Annu. Rev. Entomology</source> <volume>28</volume>, <fpage>1</fpage>&#x2013;<lpage>24</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1146/annurev.en.28.010183.000245</pub-id>
</mixed-citation>
</ref>
<ref id="B22">
<mixed-citation publication-type="book">
<person-group person-group-type="author"><collab>International Seed Testing Association</collab>
</person-group> (<year>2020</year>). <source>International rules for seed testing (Rule 9: Moisture content determination)</source> (<publisher-loc>Wallisellen, Switzerland</publisher-loc>: 
<publisher-name>ISTA</publisher-name>). doi:&#xa0;<pub-id pub-id-type="doi">10.15258/istarules.2020</pub-id>
</mixed-citation>
</ref>
<ref id="B23">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Kasozi</surname> <given-names>L.</given-names></name>
<name><surname>Derera</surname> <given-names>J.</given-names></name>
<name><surname>Tongoona</surname> <given-names>P.</given-names></name>
<name><surname>Tukamuhabwa</surname> <given-names>P.</given-names></name>
<name><surname>Muwonge</surname> <given-names>A.</given-names></name>
<name><surname>Asea</surname> <given-names>G.</given-names></name>
</person-group> (<year>2016</year>). 
<article-title>Genotypic variation for maize weevil resistance in eastern and southern Africa maize inbred lines</article-title>. <source>Uganda J. Agric. Sci.</source> <volume>17</volume>, <fpage>83</fpage>&#x2013;<lpage>97</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.4314/ujas.v17i1.8</pub-id>
</mixed-citation>
</ref>
<ref id="B24">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Kirkendall</surname> <given-names>L. R.</given-names></name>
</person-group> (<year>2018</year>). 
<article-title>Invasive bark beetles (Coleoptera, Curculionidae, Scolytinae) in Chile and Argentina, including two species new for South America, and the correct identity of the Orthotomicus species in Chile and Argentina</article-title>. <source>Diversity</source> <volume>10</volume>, <fpage>40</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/d10020040</pub-id>
</mixed-citation>
</ref>
<ref id="B25">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Kossou</surname> <given-names>D. K.</given-names></name>
<name><surname>Bosque-P&#xe9;rez</surname> <given-names>N. A.</given-names></name>
<name><surname>Mareck</surname> <given-names>J. H.</given-names></name>
</person-group> (<year>1993</year>). 
<article-title>Comparison of improved and local maize varieties in the Republic of Benin with emphasis on susceptibility to <italic>Sitophilus zeamais</italic> Motschulsky</article-title>. <source>J. Stored Products Res.</source> <volume>29</volume>, <fpage>333</fpage>&#x2013;<lpage>343</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/0022-474X(93)90031-H</pub-id>
</mixed-citation>
</ref>
<ref id="B26">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Kumar</surname> <given-names>D.</given-names></name>
<name><surname>Kalita</surname> <given-names>P.</given-names></name>
</person-group> (<year>2017</year>). 
<article-title>Reducing postharvest losses during storage of grain crops to strengthen food security in developing countries</article-title>. <source>Foods</source> <volume>6</volume>, <fpage>8</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/foods6010008</pub-id>, PMID: <pub-id pub-id-type="pmid">28231087</pub-id>
</mixed-citation>
</ref>
<ref id="B27">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>L&#xf3;pez-Castillo</surname> <given-names>L. M.</given-names></name>
<name><surname>Silva-Fern&#xe1;ndez</surname> <given-names>S. E.</given-names></name>
<name><surname>Winkler</surname> <given-names>R.</given-names></name>
<name><surname>Bergvinson</surname> <given-names>D. J.</given-names></name>
<name><surname>Arnason</surname> <given-names>J. T.</given-names></name>
<name><surname>Garc&#xed;a-Lara</surname> <given-names>S.</given-names></name>
</person-group> (<year>2018</year>). 
<article-title>Postharvest insect resistance in maize</article-title>. <source>J. Stored Products Res.</source> <volume>77</volume>, <fpage>66</fpage>&#x2013;<lpage>76</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2018.03.004</pub-id>
</mixed-citation>
</ref>
<ref id="B28">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Marcos</surname> <given-names>R. A.</given-names></name>
<name><surname>Holtz</surname> <given-names>F. G.</given-names></name>
<name><surname>de Oliveira Ramos</surname> <given-names>M. M.</given-names></name>
<name><surname>da Silva</surname> <given-names>J. A.</given-names></name>
<name><surname>Ferreira Simeone</surname> <given-names>M. L.</given-names></name>
<name><surname>Zago</surname> <given-names>H. B.</given-names></name>
<etal/>
</person-group>. (<year>2025</year>). 
<article-title>Antixenosis of different maize genotypes in storage affects feeding preferences and oviposition of <italic>Sitophilus zeamais</italic> Motschulsky</article-title>. <source>Scientia Agropecuaria</source> <volume>16</volume>, <fpage>409</fpage>&#x2013;<lpage>416</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.17268/sci.agropecu.2025.031</pub-id>
</mixed-citation>
</ref>
<ref id="B29">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Markham</surname> <given-names>R.</given-names></name>
<name><surname>Bosque-Perez</surname> <given-names>N.</given-names></name>
<name><surname>Borgemeister</surname> <given-names>C.</given-names></name>
<name><surname>Meikle</surname> <given-names>W.</given-names></name>
</person-group> (<year>1994</year>). 
<article-title>Developing pest management strategies for <italic>Sitophilus zeamais</italic> and <italic>Prostephanus truncatus</italic> in the tropics</article-title>. <source>FAO Plant Protection Bulletin</source>, <volume>42</volume> (<issue>3</issue>), <page-range>97&#x2013;116</page-range>.
</mixed-citation>
</ref>
<ref id="B30">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Mbah</surname> <given-names>A. K.</given-names></name>
<name><surname>Paothong</surname> <given-names>A.</given-names></name>
</person-group> (<year>2015</year>). 
<article-title>Shapiro&#x2013;Francia test compared to other normality tests using expected p-value</article-title>. <source>J. Stat. Comput. Simulation</source> <volume>85</volume>, <fpage>3002</fpage>&#x2013;<lpage>3016</lpage>.
</mixed-citation>
</ref>
<ref id="B31">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Mesterh&#xe1;zy</surname> <given-names>&#xc1;.</given-names></name>
<name><surname>Ol&#xe1;h</surname> <given-names>J.</given-names></name>
<name><surname>Popp</surname> <given-names>J.</given-names></name>
</person-group> (<year>2020</year>). 
<article-title>Losses in the grain supply chain: Causes and solutions</article-title>. <source>Sustainability</source> <volume>12</volume>, <fpage>2342</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/su12062342</pub-id>
</mixed-citation>
</ref>
<ref id="B32">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Mmasa</surname> <given-names>Z.</given-names></name>
<name><surname>Mathur</surname> <given-names>S.</given-names></name>
</person-group> (<year>2020</year>). 
<article-title>Maize production and food security in Tanzania</article-title>. <source>Afr. J. Agric. Res.</source> <volume>15</volume>, <fpage>1052</fpage>&#x2013;<lpage>1061</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.5897/AJAR2020.15050</pub-id>
</mixed-citation>
</ref>
<ref id="B33">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Mutambuki</surname> <given-names>K.</given-names></name>
<name><surname>Likhayo</surname> <given-names>P.</given-names></name>
</person-group> (<year>2021</year>). 
<article-title>Efficacy of different hermetic bag storage technologies against insect pests and aflatoxin incidence in stored maize grain</article-title>. <source>Bull. Entomological Res.</source> <volume>111</volume>, <fpage>499</fpage>&#x2013;<lpage>510</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1017/S0007485321000213</pub-id>, PMID: <pub-id pub-id-type="pmid">33766166</pub-id>
</mixed-citation>
</ref>
<ref id="B34">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Mutungi</surname> <given-names>C.</given-names></name>
<name><surname>Muthoni</surname> <given-names>F.</given-names></name>
<name><surname>Bekunda</surname> <given-names>M.</given-names></name>
<name><surname>Gaspar</surname> <given-names>A.</given-names></name>
<name><surname>Kabula</surname> <given-names>E.</given-names></name>
<name><surname>Abass</surname> <given-names>A.</given-names></name>
</person-group> (<year>2019</year>). 
<article-title>Physical quality of maize grain harvested and stored by smallholder farmers in the Northern highlands of Tanzania: Effects of harvesting and pre-storage handling practices</article-title>. <source>J. Stored Products Res.</source> <volume>84</volume>, <fpage>101517</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2019.101517</pub-id>
</mixed-citation>
</ref>
<ref id="B35">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Mutungi</surname> <given-names>C.</given-names></name>
<name><surname>Tungu</surname> <given-names>J.</given-names></name>
<name><surname>Amri</surname> <given-names>J.</given-names></name>
<name><surname>Gaspar</surname> <given-names>A.</given-names></name>
<name><surname>Abass</surname> <given-names>A.</given-names></name>
</person-group> (<year>2022</year>). 
<article-title>Nutritional benefits of improved post-harvest handling practices for maize and common beans in Northern Tanzania: A quantitative farm-level assessment</article-title>. <source>J. Stored Products Res.</source> <volume>95</volume>, <fpage>101918</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2021.101918</pub-id>
</mixed-citation>
</ref>
<ref id="B36">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Ngom</surname> <given-names>D.</given-names></name>
<name><surname>Fauconnier</surname> <given-names>M.-L.</given-names></name>
<name><surname>Malumba</surname> <given-names>P.</given-names></name>
<name><surname>Dia</surname> <given-names>C. A. K. M.</given-names></name>
<name><surname>Thiaw</surname> <given-names>C.</given-names></name>
<name><surname>Semb&#xe8;ne</surname> <given-names>M.</given-names></name>
</person-group> (<year>2020</year>). 
<article-title>Varietal susceptibility of maize to larger grain borer, <italic>Prostephanus truncatus</italic> (Horn) (Coleoptera; Bostrichidae), based on grain physicochemical parameters</article-title>. <source>PloS One</source> <volume>15</volume>, <elocation-id>e0232164</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1371/journal.pone.0232164</pub-id>, PMID: <pub-id pub-id-type="pmid">32330189</pub-id>
</mixed-citation>
</ref>
<ref id="B37">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Njoroge</surname> <given-names>A.</given-names></name>
<name><surname>Affognon</surname> <given-names>H.</given-names></name>
<name><surname>Mutungi</surname> <given-names>C.</given-names></name>
<name><surname>Manono</surname> <given-names>J.</given-names></name>
<name><surname>Lamuka</surname> <given-names>P.</given-names></name>
<name><surname>Murdock</surname> <given-names>L.</given-names></name>
</person-group> (<year>2014</year>). 
<article-title>Triple bag hermetic storage delivers a lethal punch to Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae) in stored maize</article-title>. <source>J. Stored Products Res.</source> <volume>58</volume>, <fpage>12</fpage>&#x2013;<lpage>19</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2014.02.005</pub-id>
</mixed-citation>
</ref>
<ref id="B38">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Njoroge</surname> <given-names>A. W.</given-names></name>
<name><surname>Mugo</surname> <given-names>S.</given-names></name>
<name><surname>Wanjohi</surname> <given-names>B.</given-names></name>
</person-group> (<year>2019</year>). 
<article-title>Post-harvest losses of maize due to storage insects in East Africa</article-title>. <source>Int. J. Agric. Res.</source> <volume>14</volume>, <fpage>123</fpage>&#x2013;<lpage>134</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3923/ijar.2019.123.134</pub-id>
</mixed-citation>
</ref>
<ref id="B39">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Nwosu</surname> <given-names>L. C.</given-names></name>
</person-group> (<year>2018</year>). 
<article-title>Maize and the maize weevil: Advances and innovations in postharvest control of the pest</article-title>. <source>Food Qual. Saf.</source> <volume>2</volume>, <fpage>145</fpage>&#x2013;<lpage>152</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/fqsafe/fyy011</pub-id>
</mixed-citation>
</ref>
<ref id="B40">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Okello</surname> <given-names>S.</given-names></name>
<name><surname>Reichmuth</surname> <given-names>F.</given-names></name>
<name><surname>Schulz</surname> <given-names>F.</given-names></name>
</person-group> (<year>1996</year>). 
<article-title>Laboratory investigations on the developmental rate at low relative humidity and the developmental temperature limit of <italic>Pagiocerus frontalis</italic> (Fab.) (Col., Scolytidae) at high and low temperatures</article-title>. <source>Anzeiger f&#xfc;r Sch&#xe4;dlingskunde Pflanzenschutz Umweltschutz</source> <volume>69</volume>, <fpage>180</fpage>&#x2013;<lpage>182</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/BF01908441</pub-id>
</mixed-citation>
</ref>
<ref id="B41">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Panagiotakis</surname> <given-names>A.</given-names></name>
<name><surname>Baliota</surname> <given-names>G. V.</given-names></name>
<name><surname>Rumbos</surname> <given-names>C. I.</given-names></name>
<name><surname>Athanassiou</surname> <given-names>C. G.</given-names></name>
</person-group> (<year>2023</year>). 
<article-title>Efficacy of contact insecticides for the control of the larger grain borer, <italic>prostephanus truncatus</italic> (Horn), on stored maize</article-title>. <source>Agriculture</source> <volume>13</volume>, <elocation-id>1502</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/agriculture13081502</pub-id>
</mixed-citation>
</ref>
<ref id="B42">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Quellhorst</surname> <given-names>H.</given-names></name>
<name><surname>Athanassiou</surname> <given-names>C. G.</given-names></name>
<name><surname>Bruce</surname> <given-names>A.</given-names></name>
<name><surname>Scully</surname> <given-names>E. D.</given-names></name>
<name><surname>Morrison</surname> <given-names>W. R.</given-names> <suffix>III</suffix></name>
</person-group> (<year>2020</year>). 
<article-title>Temperature-mediated competition between the invasive larger grain borer (Coleoptera: Bostrichidae) and the cosmopolitan maize weevil (Coleoptera: Curculionidae)</article-title>. <source>Environ. Entomology</source> <volume>49</volume>, <fpage>255</fpage>&#x2013;<lpage>264</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/ee/nvz151</pub-id>, PMID: <pub-id pub-id-type="pmid">31848593</pub-id>
</mixed-citation>
</ref>
<ref id="B43">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Quellhorst</surname> <given-names>H. E.</given-names></name>
<name><surname>Sakka</surname> <given-names>M. K.</given-names></name>
<name><surname>Morrison</surname> <given-names>W. R.</given-names> <suffix>III</suffix></name>
<name><surname>Zhu</surname> <given-names>K. Y.</given-names></name>
<name><surname>Athanassiou</surname> <given-names>C. G.</given-names></name>
</person-group> (<year>2024</year>). 
<article-title>Population growth of two strains of the larger grain borer, <italic>Prostephanus truncatus</italic> on three maize hybrids grown in Europe</article-title>. <source>Crop Prot.</source> <volume>183</volume>, <fpage>106752</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.cropro.2024.106752</pub-id>
</mixed-citation>
</ref>
<ref id="B44">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Sakka</surname> <given-names>M. K.</given-names></name>
<name><surname>Gatzali</surname> <given-names>F.</given-names></name>
<name><surname>Karathanos</surname> <given-names>V.</given-names></name>
<name><surname>Athanassiou</surname> <given-names>C. G.</given-names></name>
</person-group> (<year>2022</year>). 
<article-title>Efficacy of low oxygen against <italic>Trogoderma granarium</italic> Everts, <italic>Tribolium castaneum</italic> (Herbst), and <italic>Callosobruchus maculatus</italic> (F.) in commercial applications</article-title>. <source>J. Stored Products Res.</source> <volume>97</volume>, <fpage>101968</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2022.101968</pub-id>
</mixed-citation>
</ref>
<ref id="B45">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Tefera</surname> <given-names>T.</given-names></name>
<name><surname>Mugo</surname> <given-names>S.</given-names></name>
<name><surname>Likhayo</surname> <given-names>P.</given-names></name>
</person-group> (<year>2011</year>). 
<article-title>Effects of insect population density and storage time on grain damage and weight loss due to the maize weevil, <italic>Sitophilus zeamais</italic>, in maize</article-title>. <source>J. Stored Products Res.</source> <volume>47</volume>, <fpage>20</fpage>&#x2013;<lpage>25</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jspr.2010.08.002</pub-id>
</mixed-citation>
</ref>
<ref id="B46">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Yust</surname> <given-names>C.</given-names></name>
</person-group> (<year>1957</year>). 
<article-title>Observations on the biology of <italic>Pagiocerus frontalis</italic></article-title>. <source>J. Economic Entomology</source> <volume>50</volume>, <fpage>405</fpage>&#x2013;<lpage>408</lpage>.
</mixed-citation>
</ref>
</ref-list>
<fn-group>
<fn id="n1" fn-type="custom" custom-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/312939">Ana Butron</ext-link>, Spanish Council for Scientific Research (MBG-CSIC), Spain</p></fn>
<fn id="n2" fn-type="custom" custom-type="reviewed-by">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/117603">Ali R. Bandani</ext-link>, University of Tehran, Iran</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3287600">Ginson Riungu</ext-link>, Kenya Agricultural and Livestock Research Organization, Kenya</p></fn>
</fn-group>
</back>
</article>