<?xml version="1.0" encoding="UTF-8"?><!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "http://dtd.nlm.nih.gov/publishing/3.0/journalpublishing3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" dtd-version="3.0" xml:lang="en" article-type="research article">
 <front>
  <journal-meta>
   <journal-id journal-id-type="publisher-id">
    oje
   </journal-id>
   <journal-title-group>
    <journal-title>
     Open Journal of Ecology
    </journal-title>
   </journal-title-group>
   <issn pub-type="epub">
    2162-1985
   </issn>
   <issn publication-format="print">
    2162-1993
   </issn>
   <publisher>
    <publisher-name>
     Scientific Research Publishing
    </publisher-name>
   </publisher>
  </journal-meta>
  <article-meta>
   <article-id pub-id-type="doi">
    10.4236/oje.2025.157025
   </article-id>
   <article-id pub-id-type="publisher-id">
    oje-144174
   </article-id>
   <article-categories>
    <subj-group subj-group-type="heading">
     <subject>
      Articles
     </subject>
    </subj-group>
    <subj-group subj-group-type="Discipline-v2">
     <subject>
      Earth 
     </subject>
     <subject>
       Environmental Sciences
     </subject>
    </subj-group>
   </article-categories>
   <title-group>
    Invisible Architecture of Tephritids (Diptera): A Chemically Mediated Exploration of Ecological Structure in Tropical Orchards of Northern Cameroon
   </title-group>
   <contrib-group>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Didi Gaëlle
      </surname>
      <given-names>
       Mokam
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff1"> 
      <sup>1</sup>
     </xref>
    </contrib>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Aoutougour
      </surname>
      <given-names>
       Ndakabo
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff2"> 
      <sup>2</sup>
     </xref>
    </contrib>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Ezechiel
      </surname>
      <given-names>
       Awono
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff2"> 
      <sup>2</sup>
     </xref>
    </contrib>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Albert
      </surname>
      <given-names>
       Ngakou
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff1"> 
      <sup>1</sup>
     </xref>
    </contrib>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Ulrich Tanegang
      </surname>
      <given-names>
       Lambou
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff2"> 
      <sup>2</sup>
     </xref>
    </contrib>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Théodore Wilfried Ounguetou
      </surname>
      <given-names>
       Antani
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff2"> 
      <sup>2</sup>
     </xref>
    </contrib>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Désirée Chantal
      </surname>
      <given-names>
       Aléné
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff2"> 
      <sup>2</sup>
     </xref> 
     <xref ref-type="aff" rid="aff3"> 
      <sup>3</sup>
     </xref>
    </contrib>
    <contrib contrib-type="author" xlink:type="simple">
     <name name-style="western">
      <surname>
       Champlain
      </surname>
      <given-names>
       Djieto-Lordon
      </given-names>
     </name> 
     <xref ref-type="aff" rid="aff2"> 
      <sup>2</sup>
     </xref>
    </contrib>
   </contrib-group> 
   <aff id="aff1">
    <addr-line>
     aLaboratory of Zoology, Department of Biological Science, Faculty of Science, University of Ngaoundéré, Ngaoundéré, Cameroon
    </addr-line> 
   </aff> 
   <aff id="aff2">
    <addr-line>
     aLaboratory of Zoology, Department of Animal Biology and Physiology, Faculty of Science, University of Yaoundé I, Yaoundé, Cameroon
    </addr-line> 
   </aff> 
   <aff id="aff3">
    <addr-line>
     aFaculty of Science, University of Ebolowa, Ebolowa, Cameroon
    </addr-line> 
   </aff> 
   <pub-date pub-type="epub">
    <day>
     23
    </day> 
    <month>
     07
    </month>
    <year>
     2025
    </year>
   </pub-date> 
   <volume>
    15
   </volume> 
   <issue>
    07
   </issue>
   <fpage>
    435
   </fpage>
   <lpage>
    449
   </lpage>
   <history>
    <date date-type="received">
     <day>
      6,
     </day>
     <month>
      July
     </month>
     <year>
      2025
     </year>
    </date>
    <date date-type="published">
     <day>
      20,
     </day>
     <month>
      July
     </month>
     <year>
      2025
     </year> 
    </date> 
    <date date-type="accepted">
     <day>
      20,
     </day>
     <month>
      July
     </month>
     <year>
      2025
     </year> 
    </date>
   </history>
   <permissions>
    <copyright-statement>
     © Copyright 2014 by authors and Scientific Research Publishing Inc. 
    </copyright-statement>
    <copyright-year>
     2014
    </copyright-year>
    <license>
     <license-p>
      This work is licensed under the Creative Commons Attribution International License (CC BY). http://creativecommons.org/licenses/by/4.0/
     </license-p>
    </license>
   </permissions>
   <abstract>
    Understanding the ecology of tephritid communities is essential for developing sustainable pest management strategies in tropical fruit-based agroecosystems. This study assessed relative abundance and the efficacy of four male-specific semiochemical lures (methyl eugenol (ME), cue-lure (CU), terpinyl acetate (TA), and trimedlure (TR)), across two agroecological zones (AEZs) in Cameroon: the Sudano-Sahelian savannah (AEZ 1) and the High Guinea savannah (AEZ 2). Nine tephritid species were recorded, including two previously unreported in these AEZs (Ceratitis punctata and C. fasciventris). Despite a shared species pool, community structure differed significantly between AEZs. In AEZ 2, Bactrocera dorsalis accounted for over 90% of captures, reflecting its invasive dominance. This was amplified by the high specificity of ME, which almost exclusively attracted B. dorsalis. Conversely, AEZ 1 supported a more balanced assemblage dominated by B. dorsalis, Zeugodacus cucurbitae, and Ceratitis cosyra, particularly in dry AEZ rich in cucurbit crops. Thus, CU was effective in capturing cucurbit-associated species (Z. cucurbitae, Dacus spp.), while TR and TA attracted a broader range of Ceratitis species. These findings highlight the value of multi-lure trap systems in accurately revealing tephritid community composition and support the need for agroecologically tailored Integrated Pest Management (IPM). Promoting the use of semiochemical surveillance in smallholder systems will require institutional support to offset lure costs. Such strategies can improve early detection and targeted control of key tephritid pests, ultimately enhancing crop protection in vulnerable agricultural landscapes of various sub-Saharan Africa countries. 
   </abstract>
   <kwd-group> 
    <kwd>
     Male Lure
    </kwd> 
    <kwd>
      Bactrocera dorsalis
    </kwd> 
    <kwd>
      Ceratitis cosyra
    </kwd> 
    <kwd>
      Zeugodacus cucurbitae
    </kwd> 
    <kwd>
      Agroecosystems
    </kwd> 
    <kwd>
      Pest Surveillance
    </kwd>
   </kwd-group>
  </article-meta>
 </front>
 <body>
  <sec id="s1">
   <title>1. Introduction</title>
   <p>Fruit flies (Diptera: Tephritidae) are among the most economically significant phytophagous pests worldwide, infesting a wide range of fruits and vegetables and causing substantial pre- and post-harvest losses <xref ref-type="bibr" rid="scirp.144174-1">
     [1]
    </xref>-<xref ref-type="bibr" rid="scirp.144174-3">
     [3]
    </xref>. Their destructive potential is attributed to a combination of biological traits, including broad host range <xref ref-type="bibr" rid="scirp.144174-4">
     [4]
    </xref>, high fecundity, and strong dispersal ability <xref ref-type="bibr" rid="scirp.144174-5">
     [5]
    </xref>-<xref ref-type="bibr" rid="scirp.144174-7">
     [7]
    </xref>. Damage occurs when female oviposit into host fruit, and larval feeding rapidly degrades fruit tissue, leading to unmarketable produce <xref ref-type="bibr" rid="scirp.144174-7">
     [7]
    </xref>-<xref ref-type="bibr" rid="scirp.144174-9">
     [9]
    </xref>.</p>
   <p>In sub-Saharan Africa, tephritid outbreaks are driven not only by the abundance of cultivated and wild host plants, weak surveillance infrastructure, and inadequate management strategies <xref ref-type="bibr" rid="scirp.144174-10">
     [10]
    </xref> <xref ref-type="bibr" rid="scirp.144174-11">
     [11]
    </xref>, but also by biological invasions. Two highly aggressive invasive tephritid species Bactrocera dorsalis and Zeugodacus cucurbitae, both of Asian origin have rapidly colonized much of the region, and fundamentally may reshaping tephritid communities <xref ref-type="bibr" rid="scirp.144174-12">
     [12]
    </xref> <xref ref-type="bibr" rid="scirp.144174-13">
     [13]
    </xref>.</p>
   <p>In Cameroon, tephritid infestations are widespread and severe. Overripe and attacked fruits often accumulate in orchards, markets, and disposal sites, serving as persistent reservoirs that sustain local tephritid populations <xref ref-type="bibr" rid="scirp.144174-12">
     [12]
    </xref>. Economically important species belong primarily to four genera: Bactrocera, Ceratitis, Dacus, and Zeugodacus <xref ref-type="bibr" rid="scirp.144174-9">
     [9]
    </xref> <xref ref-type="bibr" rid="scirp.144174-13">
     [13]
    </xref>. Among these, Bactrocera dorsalis and Ceratitis cosyra are the major pests of mango <xref ref-type="bibr" rid="scirp.144174-14">
     [14]
    </xref> and guava <xref ref-type="bibr" rid="scirp.144174-15">
     [15]
    </xref> <xref ref-type="bibr" rid="scirp.144174-16">
     [16]
    </xref>, whereas Dacus bivittatus, Dacus ciliatus, Dacus punctatifrons, and Zeugodacus cucurbitae inflict considerable damage on cucurbit crops <xref ref-type="bibr" rid="scirp.144174-9">
     [9]
    </xref> <xref ref-type="bibr" rid="scirp.144174-13">
     [13]
    </xref>.</p>
   <p>Beyond direct yield losses, tephritid infestations also generate substantial indirect costs, particularly due to quarantine restrictions and trade barriers. The continued reliance on broad-spectrum insecticides is becoming increasingly unsustainable because of the development of resistance, growing environmental and human health concerns, the accumulation of toxic residues on fruits, and the inaccessibility of immature stages of tephritids that remain concealed within host tissues <xref ref-type="bibr" rid="scirp.144174-5">
     [5]
    </xref> <xref ref-type="bibr" rid="scirp.144174-17">
     [17]
    </xref>-<xref ref-type="bibr" rid="scirp.144174-20">
     [20]
    </xref>. As a result, Integrated Pest Management (IPM) strategies incorporating semiochemical attractants such as methyl eugenol (ME), cue-lure (CL), trimedlure (TML), and terpinyl acetate (TA), have gained importance, both for surveillance and, in some cases, for population suppression <xref ref-type="bibr" rid="scirp.144174-3">
     [3]
    </xref> <xref ref-type="bibr" rid="scirp.144174-21">
     [21]
    </xref>-<xref ref-type="bibr" rid="scirp.144174-23">
     [23]
    </xref>. Among these, ME is the most widely validated and operationalized semiochemical within IPM programs, particularly through the Male Annihilation Technique (MAT), owing to its exceptional attractiveness and specificity for Bactrocera dorsalis males <xref ref-type="bibr" rid="scirp.144174-24">
     [24]
    </xref>. Despite their proven effectiveness, the adoption of these tools remains limited in Cameroon, primarily due to a lack of awareness, accessibility, purchasing capacity, and context-specific recommendations tailored to smallholder farmers, most of whom operate under resource-constrained conditions.</p>
   <p>Developing effective IPM programs requires a comprehensive understanding of the local ecology of tephritid communities, including species composition, abundance patterns, and species-specific responses to male-targeted lures. This study seeks to address critical knowledge gaps in tephritid ecology in Cameroon by: 1) characterizing the diversity of male lures-responsive tephritid species, 2) assessing their ecological community structure, and 3) evaluating the efficacy and selectivity of major male lures across the studied agroecosystems.</p>
  </sec><sec id="s2">
   <title>2. Materials and Methods</title>
   <sec id="s2_1">
    <title>2.1. Study Area and Sampling Period</title>
    <p>Field data were collected during two years, from 2021 to 2022, in five orchards located in two agroecological zones (AEZs) of Cameroon. These orchards, previously described in <xref ref-type="bibr" rid="scirp.144174-2">
      [2]
     </xref>, were not treated with insecticides. Detailed agroclimatic and edaphic profiles of each AEZ are also available in <xref ref-type="bibr" rid="scirp.144174-2">
      [2]
     </xref>.</p>
    <p>AEZ 1 (Sudano-Sahelian Savannah) included two orchards situated in Ngong and Mbé, while AEZ 2 (High Guinea Savannah) comprised three orchards located in Malang, Manwi, and Marza. Each selected orchard covered at least five hectares and featured mixed tropical fruit trees, with mango (Mangifera indica L.) and guava (Psidium guajava L.) as the dominant species. All orchards were accessible and suitable for long-term ecological monitoring.</p>
   </sec>
   <sec id="s2_2">
    <title>2.2. Trap and Lure Configurations</title>
    <p>A total of four tephritid traps were installed at each sampling point, each baited with one of four male-specific lures: methyl-eugenol (ME), cue-lure (CU), trimedlure (TR), and terpinyl acetate (TA) <xref ref-type="bibr" rid="scirp.144174-21">
      [21]
     </xref> <xref ref-type="bibr" rid="scirp.144174-25">
      [25]
     </xref> <xref ref-type="bibr" rid="scirp.144174-26">
      [26]
     </xref>. Every trap contained the organophosphate insecticide dichlorvos (2,2-dichlorovinyl dimethyl phosphate, DDVP) as the killing agent. Traps were hung about 1.5 m above ground on sturdy fruit-tree branches. To minimize odor interference and edge effects, trap placement followed a standardized layout: the first trap was positioned 30 m inside the orchard boundary, and the remaining traps were set at 50 m intervals <xref ref-type="bibr" rid="scirp.144174-27">
      [27]
     </xref>. The <xref ref-type="fig" rid="fig1">
      Figure 1
     </xref> illustrates the typical trap configuration.</p>
   </sec>
   <sec id="s2_3">
    <title>2.3. Sampling and Trap Maintenance</title>
    <p>A total of 16 traps was installed in each orchard (4 replicates × 4 lure types). The lures and DDVP insecticide were replaced every six weeks, resulting in eight complete sampling cycles per year in each orchard <xref ref-type="bibr" rid="scirp.144174-28">
      [28]
     </xref>. Sterile gloves were used</p>
    <fig id="fig1" position="float">
     <label>Figure 1</label>
     <caption>
      <title>Figure 1. Tephri-traps baited with male-specific lures to tephritids in five orchards in Cameroon during the 2021-2022 sampling period. From left to right, the traps contain: methyl eugenol (ME), cue-lure (CU), terpinyl acetate (TA), and trimedlure (TR), each paired with a DDVP-based insecticide.</title>
     </caption>
     <graphic mimetype="image" position="float" xlink:type="simple" xlink:href="https://html.scirp.org/file/1381776-rId22.jpeg?20250723115326" />
    </fig>
    <p>during all handling procedures to prevent cross-contamination between attractants. To reduce ant predation and interference from other organisms, a thick layer of automotive grease was applied to the branches used to suspend the traps <xref ref-type="bibr" rid="scirp.144174-29">
      [29]
     </xref>. Captured tephritid specimens were collected weekly and individually preserved in labeled entomological tubes containing 70% ethanol. All samples were then transported to the Zoology Laboratory of the University of Ngaoundéré for taxonomic identification.</p>
   </sec>
   <sec id="s2_4">
    <title>2.4. Identification Procedure</title>
    <p>Tephritid specimens were sorted under a stereomicroscope and identified to species level using standard morphological keys <xref ref-type="bibr" rid="scirp.144174-30">
      [30]
     </xref>. Additional identifications were carried out to confirm initial determinations at the French Agency for Food, Environmental and Occupational Health &amp; Safety (ANSES) in Montpellier, France, and at the Royal Museum for Central Africa (RMCA) in Tervuren, Belgium.</p>
   </sec>
   <sec id="s2_5">
    <title>2.5. Assessment of Alpha Diversity</title>
    <p>Alpha diversity was assessed using a tripartite approach that included: rarefaction analysis (species accumulation curves), rank-abundance (Whittaker) curves, and complementary diversity indices <xref ref-type="bibr" rid="scirp.144174-31">
      [31]
     </xref>.</p>
    <p>Rarefaction curves were used to evaluate sampling completeness, with a 95% asymptotic plateau considered indicative of adequate sampling effort <xref ref-type="bibr" rid="scirp.144174-32">
      [32]
     </xref>.</p>
    <p>Species abundance distributions were visualized using rank-abundance curves, where log-transformed abundance is plotted against species rank, in order to analyze patterns of dominance and evenness <xref ref-type="bibr" rid="scirp.144174-33">
      [33]
     </xref>.</p>
    <p>Quantitative assessment of diversity was based on three complementary indices:</p>
    <p>where p<sub>i</sub> is the proportion of individuals belonging to tephritid species i, and S is the total species richness <xref ref-type="bibr" rid="scirp.144174-31">
      [31]
     </xref> <xref ref-type="bibr" rid="scirp.144174-34">
      [34]
     </xref> <xref ref-type="bibr" rid="scirp.144174-35">
      [35]
     </xref>.</p>
    <p>The Shannon-Wiener index ranges from 0 to ln(S), indicating maximum diversity. Higher values of H' and 1 − D reflect greater diversity and more balanced communities. Pielou’s evenness index, ranging from 0 to 1, indicates more equitable species distribution when values are closer to 1.</p>
   </sec>
   <sec id="s2_6">
    <title>2.6. Assessment of Beta Diversity</title>
    <p>Beta diversity between AEZs and orchards was evaluated using the Jaccard similarity index:</p>
    <p>
     <math xmlns="http://www.w3.org/1998/Math/MathML"> <mrow> 
       <mi>
         J 
       </mi> 
       <mrow> 
        <mo>
          ( 
        </mo> 
        <mrow> 
         <mtext>
           A 
         </mtext> 
         <mo>
           , 
         </mo> 
         <mtext>
           B 
         </mtext> 
        </mrow> 
        <mo>
          ) 
        </mo> 
       </mrow> 
       <mo>
         = 
       </mo> 
       <mfrac> 
        <mi>
          a 
        </mi> 
        <mrow> 
         <mi>
           a 
         </mi> 
         <mo>
           + 
         </mo> 
         <mi>
           b 
         </mi> 
         <mo>
           + 
         </mo> 
         <mi>
           c 
         </mi> 
        </mrow> 
       </mfrac> 
      </mrow> 
     </math>;</p>
    <p>where a is the number of species shared by orchards A and B, b is the number of species unique to orchard A, and c is the number of species unique to orchard B <xref ref-type="bibr" rid="scirp.144174-36">
      [36]
     </xref>. The index ranges from 0 (no species shared) to 1 (identical species composition). Pairwise similarity values were compiled into a matrix and visualized as a heatmap using a yellow-orange-red (YlOrRd) gradient scale from 0 to 100% to to illustrate the degree of species overlap.</p>
   </sec>
   <sec id="s2_7">
    <title>2.7. Evaluation of Lure Efficacy and Selectivity</title>
    <p>To evaluate lure efficacy, the mean number of tephritids captured per trap and per sampling day was calculated for each male lure across all orchards. Statistical comparisons of abundance were performed using analysis of variance (ANOVA), followed by Tukey’s post hoc test (p &lt; 0.05), using XLSTAT software.</p>
    <p>Selectivity was assessed by calculating the relative abundance of each species per male lure. The proportion of total captures attributed to each tephritid species was used to determine lure specificity and performance for targeted surveillance.</p>
   </sec>
  </sec><sec id="s3">
   <title>3. Results</title>
   <sec id="s3_1">
    <title>3.1. Alpha Diversity</title>
    <p>Species accumulation curves plateaued across all five orchards, indicating that the sampling effort, measured by cumulative sampling days, was sufficient to capture the local pool of male tephritid species and to support robust ecological analyses (<xref ref-type="fig" rid="fig2">
      Figure 2
     </xref>). The asymptotic trend consistently observed in the curves of each orchard confirms that further sampling would likely yield only marginal gains in species detection.</p>
    <p>The Whittaker diagram highlighted patterns of dominance within the tephritid communities studied across the AEZs. In particular, in AEZ 2, the steep slope of the curves reflected a highly unbalanced community, overwhelmingly dominated by Bactrocera dorsalis (<xref ref-type="fig" rid="fig3">
      Figure 3
     </xref>). Conversely, the flatter curves observed in AEZ 1, notably in Ngong, indicated a more equitable species distribution, with several major pest species contributing significantly to community structure, especially B. dorsalis, Ceratitis cosyra, and Zeugodacus cucurbitae (<xref ref-type="fig" rid="fig3">
      Figure 3
     </xref>).</p>
    <fig id="fig2" position="float">
     <label>Figure 2</label>
     <caption>
      <title>Figure 2. Species accumulation curves illustrating the cumulative tephritid species richness during the 2021-2022 sampling period in five orchards located across two agroecological zones (AEZs) in Cameroon: two orchards in AEZ 1 (Mbé and Ngong), and three in AEZ 2 (Marza, Manwi, and Malang).</title>
     </caption>
     <graphic mimetype="image" position="float" xlink:type="simple" xlink:href="https://html.scirp.org/file/1381776-rId35.jpeg?20250723115330" />
    </fig>
    <fig id="fig3" position="float">
     <label>Figure 3</label>
     <caption>
      <title>Figure 3. Rank-abundance curves of tephritid communities in five orchards across two agroecological zones (AEZs) in Cameroon: two in AEZ 1 (Mbé, Ngong), and three in AEZ 2 (Marza, Manwi, and Malang), during the 2021-2022 sampling period. Species are ranked from the most to the least abundant, with abundance expressed as Log10 (abundance + 1).</title>
     </caption>
     <graphic mimetype="image" position="float" xlink:type="simple" xlink:href="https://html.scirp.org/file/1381776-rId36.jpeg?20250723115331" />
    </fig>
    <p>Diversity indices revealed marked variations between the AEZs (<xref ref-type="fig" rid="fig4">
      Figure 4
     </xref>). Orchards located in AEZ 1 exhibited high values for the Shannon-Wiener diversity index, Simpson’s index, and Pielou’s evenness index, indicating tephritid communities that were both species-rich and evenly distributed (<xref ref-type="fig" rid="fig4">
      Figure 4
     </xref>). In contrast, diversity indices were significantly lower in AEZ 2, reflecting the numerical dominance of a single species and consequently a low evenness within each orchard (<xref ref-type="fig" rid="fig4">
      Figure 4
     </xref>).</p>
    <fig id="fig4" position="float">
     <label>Figure 4</label>
     <caption>
      <title>Figure 4. Values of diversity indices: H’, 1-D, and J’, associated with tephritid communities in five orchards located in two agroecological zones (AEZs) in Cameroon: two orchards in AEZ 1 (Mbé and Ngong) and three orchards in AEZ 2 (Marza, Manwi, and Malang), during the 2020-2021 sampling period.</title>
     </caption>
     <graphic mimetype="image" position="float" xlink:type="simple" xlink:href="https://html.scirp.org/file/1381776-rId37.jpeg?20250723115331" />
    </fig>
   </sec>
   <sec id="s3_2">
    <title>3.2. Evaluation Beta Diversity</title>
    <p>Jaccard similarity index values revealed high similarity within AEZ (J &gt; 0.85), suggesting that shared species were more abundant than unique species within each AEZ (<xref ref-type="fig" rid="fig5">
      Figure 5
     </xref>). In contrast, similarity values between AEZs were notably lower (~0.60), indicating a relatively weak ecological differentiation of tephritid communities between AEZs.</p>
    <fig id="fig5" position="float">
     <label>Figure 5</label>
     <caption>
      <title>Figure 5. Heatmap showing pairwise Jaccard similarity index among tephritid communities in five orchards across two agroecological zones (AEZs) in Cameroon: two orchards in AEZ 1 (Mbé and Ngong) and three orchards in AEZ 2 (Marza, Manwi, and Malang), during the 2021-2022 sampling period. Color intensity reflects the degree of similarity, with higher intensity indicating greater similarity.</title>
     </caption>
     <graphic mimetype="image" position="float" xlink:type="simple" xlink:href="https://html.scirp.org/file/1381776-rId38.jpeg?20250723115332" />
    </fig>
   </sec>
   <sec id="s3_3">
    <title>3.3. Lure Effectiveness</title>
    <p>A total of nine male tephritid species were recorded across all surveyed orchards. Among them, Ceratitis bremii and C. punctata were absent from AEZ 2 (<xref ref-type="fig" rid="fig6">
      Figure 6
     </xref>; Supplementary material <xref ref-type="table" rid="tableS1">
      Table S1
     </xref>).</p>
    <fig id="fig6" position="float">
     <label>Figure 6</label>
     <caption>
      <title>Figure 6. Grouped horizontal bar plots by four male-specific lures: cue-lure (CU), methyl eugenol (ME), trimedlure (TA), and terpinyl acetate (TR), across five orchards located in two agroecological zones (AEZs) of Cameroon: two orchards in AEZ 1 (Mbé and Ngong) and three orchards in AEZ 2 (Marza, Manwi, and Malang), during the 2021-2022 sampling period. Bar length represents relative abundance (%).</title>
     </caption>
     <graphic mimetype="image" position="float" xlink:type="simple" xlink:href="https://html.scirp.org/file/1381776-rId39.jpeg?20250723115333" />
    </fig>
    <p>Each male-specific lure exhibited significantly distinct selectivity profiles (p &lt; 0.05), strongly influencing species composition assessments and underscoring the strategic importance of multi-lure trapping systems for comprehensive ecological surveillance and integrated tephritid management (<xref ref-type="fig" rid="fig6">
      Figure 6
     </xref>; Supplementary material <xref ref-type="table" rid="tableS2">
      Table S2
     </xref>).</p>
    <p>Methyl eugenol (ME) demonstrated exceptionally high attractiveness and specificity toward Bactrocera dorsalis, accounting for over 99% of captures across all orchards (<xref ref-type="fig" rid="fig6">
      Figure 6
     </xref>), reflecting the species’ overwhelming numerical dominance under ME-based monitoring conditions (Supplementary material <xref ref-type="table" rid="tableS1">
      Table S1
     </xref> and <xref ref-type="table" rid="tableS2">
      Table S2
     </xref>). Nevertheless, its relative abundance varied significantly between AEZs (p &lt; 0.05), with lower representation in AEZ 1, particularly at Ngong and Mbé, compared to AEZ 2, where this species peaked with averages exceeding 1000 individuals per trapping event (Supplementary material <xref ref-type="table" rid="tableS1">
      Table S1
     </xref> and <xref ref-type="table" rid="tableS2">
      Table S2
     </xref>).</p>
    <p>Cue-lure (CU) exhibited moderate but variable performance, with significantly high capture rates at Ngong (16.76 ± 4.64) and markedly low rates at Mbé (3.69 ± 0.33) (Supplementary material <xref ref-type="table" rid="tableS2">
      Table S2
     </xref>). This lure attracted several cucurbit-associated species, notably Zeugodacus cucurbitae, Dacus punctatifrons, and D. bivittatus.</p>
    <p>Trimedlure (TR) and terpinyl acetate (TA) displayed broader and less specific attraction profiles, capturing a more diverse assemblage of Ceratitis species, including C. cosyra, C. anonae, and C. fasciventris, along with a limited number of B. dorsalis individuals (<xref ref-type="fig" rid="fig6">
      Figure 6
     </xref>). This generalist attractant behavior reinforces their complementary role in species detection, particularly for less dominant or lure-insensitive tephritids.</p>
    <p>Ceratitis cosyra and Z. cucurbitae were well represented in AEZ 1, accounting for over 30% and 25% of total captures, respectively in Ngong (Supplementary material <xref ref-type="table" rid="tableS1">
      Table S1
     </xref>). Notably, this orchard has experienced an intensification of cucurbit cultivation, particularly of sweet melon (Cucumis melo L.). In contrast, C. anonae, C. fasciventris, D. punctatifrons, and D. bivittatus, although widely distributed across sites, exhibited low responsiveness to the studied lures, with capture rates generally below 5% (Supplementary material <xref ref-type="table" rid="tableS1">
      Table S1
     </xref>).</p>
   </sec>
  </sec><sec id="s4">
   <title>4. Discussion</title>
   <p>The present study provides a comprehensive assessment of the community structure of tephritids and the performance of male-specific lures across two agroecological zones (AEZs) in Cameroon. By integrating multi-lure trapping strategies with various quantitative species diversity metrics, tephritid community studied was strongly shaped by agroecological conditions <xref ref-type="bibr" rid="scirp.144174-2">
     [2]
    </xref>, while male lure effectiveness varied considerably depending on host plants used. These findings challenge the effectiveness of standardized surveillance protocols and highlight the need for ecologically adapted Integrated Pest Management (IPM) frameworks <xref ref-type="bibr" rid="scirp.144174-37">
     [37]
    </xref> <xref ref-type="bibr" rid="scirp.144174-38">
     [38]
    </xref>.</p>
   <p>The observed saturation of species accumulation curves confirms the robustness of the sampling <xref ref-type="bibr" rid="scirp.144174-32">
     [32]
    </xref>, ensuring the reliability of ecological parameter comparisons across AEZs.</p>
   <p>
    <xref ref-type="bibr" rid="scirp.144174-"></xref>Although the incubation of attacked fruits is a labor-intensive yet accurate method for confirming host plant use <xref ref-type="bibr" rid="scirp.144174-14">
     [14]
    </xref>, male lure-based trapping offers a more inclusive and cost-effective approach for monitoring adult populations, particularly for the detection and management of major pest species <xref ref-type="bibr" rid="scirp.144174-3">
     [3]
    </xref> <xref ref-type="bibr" rid="scirp.144174-6">
     [6]
    </xref> <xref ref-type="bibr" rid="scirp.144174-24">
     [24]
    </xref> <xref ref-type="bibr" rid="scirp.144174-37">
     [37]
    </xref>. The detection approach based on methyl eugenol (ME), cue-lure (CU), terpinyl acetate (TA), and TR revealed significant, context-dependent variation in performance. This enabled the identification of nine tephritid species, including previously unreported species such as C. punctata, and C. fasciventris <xref ref-type="bibr" rid="scirp.144174-29">
     [29]
    </xref>. The absence of Perilampsis sp., Bactrocera mesomelas and Notoma biocolatum, which were recorded in previous studies, may reflect differences in male lure selectivity <xref ref-type="bibr" rid="scirp.144174-29">
     [29]
    </xref>.</p>
   <p>Despite a shared species pool, clear contrasts were observed in species diversity and community structure across AEZs.</p>
   <p>Methyl eugenol (ME) demonstrated outstanding efficacy in areas dominated by Bactrocera dorsalis, confirming its high specificity. However, in AEZ 2, this pest accounted for more than 90% of captures <xref ref-type="bibr" rid="scirp.144174-2">
     [2]
    </xref> <xref ref-type="bibr" rid="scirp.144174-22">
     [22]
    </xref> <xref ref-type="bibr" rid="scirp.144174-24">
     [24]
    </xref> <xref ref-type="bibr" rid="scirp.144174-25">
     [25]
    </xref>, indicating its ecological dominance and establishment, pattern consistent with previous studies documenting the invasion and displacement effects of B. dorsalis across sub-Saharan Africa <xref ref-type="bibr" rid="scirp.144174-11">
     [11]
    </xref> <xref ref-type="bibr" rid="scirp.144174-39">
     [39]
    </xref> <xref ref-type="bibr" rid="scirp.144174-40">
     [40]
    </xref>. This dominance was particularly pronounced in the orchards of Malang, Manwi, and Marza, suggesting a homogenized community structure under intense pest pressure <xref ref-type="bibr" rid="scirp.144174-41">
     [41]
    </xref>.</p>
   <p>By contrast, AEZ 1, characterized by dry climatic conditions and a high presence of cucurbit crops, supported a more balanced assemblage dominated by B. dorsalis, Ceratitis cosyra, and Zeugodacus cucurbitae, particularly in the orchard of Ngong. This pattern may reflect habitat heterogeneity, including broader host plant or altitude-related climatic differences that mitigate interspecific interactions and promote niche partitioning <xref ref-type="bibr" rid="scirp.144174-3">
     [3]
    </xref> <xref ref-type="bibr" rid="scirp.144174-13">
     [13]
    </xref>. The higher evenness and species richness observed in this AEZ do not corroborate previous fruit incubation studies, which identified C. cosyra as the primary mango pest in this zone <xref ref-type="bibr" rid="scirp.144174-2">
     [2]
    </xref>.</p>
   <p>Cue-lure (CU) proved essential for monitoring Z. cucurbitae and Dacus spp., especially in cucurbit-dominated zones, while trimedlure (TR) and terpinyl acetate (TA) were important for detecting Ceratitis spp., despite lower overall catch rates <xref ref-type="bibr" rid="scirp.144174-24">
     [24]
    </xref>. These results suggest the necessity of deploying mixed-lure systems in tephritid surveillance programs. Sole reliance on ME, as implemented in many B. dorsalis-infested areas, risks underrepresenting non-responsive or secondary pest species catching and thereby underestimating true community diversity and pest risk.</p>
   <p>In high-diversity settings like AEZ 1, mixed-lure approaches can enhance detection accuracy, improve early warning systems, and support more nuanced pest management strategies <xref ref-type="bibr" rid="scirp.144174-42">
     [42]
    </xref>.</p>
   <p>From a management perspective, the dominance of B. dorsalis in AEZ 2 supports the implementation of Male Annihilation Technique (MAT) using ME in conjunction with strict orchard sanitation <xref ref-type="bibr" rid="scirp.144174-42">
     [42]
    </xref>. In contrast, the complex species assemblage in AEZ 1 calls for diversified IPM approaches, including multi-lure trapping, host sanitation, and possibly augmentative biological control. The success of such strategies, however, hinges on institutional support, grower education, and access to affordable lures and traps <xref ref-type="bibr" rid="scirp.144174-10">
     [10]
    </xref>.</p>
  </sec><sec id="s5">
   <title>5. Conclusions</title>
   <p>This study highlights the spatial heterogeneity of tephritid communities across tropical agroecosystems and underscores the pivotal role of agroecological context in shaping both community structure and responsiveness to male-specific semiochemical lures. It also demonstrates the methodological complementarity between male-lure-based trapping, which is highly effective for detecting mobile adult tephritid populations.</p>
   <p>Findings from this survey reveal that no single lure, whether sex-specific or generalist, nor any standardized protocol can, in isolation, effectively capture the full species pool. Consequently, integrated and zone-specific strategies, combining a diversity of attractants, are essential for accurate surveillance and sustainable pest management.</p>
   <p>Incorporating these insights into local IPM frameworks offers a promising pathway to strengthen national phytosanitary capacity, enhance crop protection, and reduce reliance on synthetic insecticides. This is especially critical in smallholder farming systems, where limited resources and high vulnerability necessitate pest control solutions that are not only effective and affordable but also ecologically sound.</p>
  </sec><sec id="s6">
   <title>Acknowledgements</title>
   <p>We are deeply grateful to Dr. Marc De Meyer of the Royal Museum for Central Africa (RMCA) in Tervuren, Belgium, for generously providing the male lures used in this study.</p>
  </sec><sec id="s7">
   <title>Supplementary Material</title>
   <table-wrap id="table1">
    <label>
     <xref ref-type="table" rid="table1">
      Table 1
     </xref></label>
    <caption>
     <title>
      <xref ref-type="bibr" rid="scirp.144174-"></xref>Table S1. Relative abundance (%) of nine tephritid species captured in five orchards located in two agroecological zones (AEZs) of Cameroon: two orchards in AEZ 1 (Mbé and Ngong) and three orchards in AEZ 2 (Marza, Manwi, and Malang), during the 2021-2022 sampling period.</title>
    </caption>
    <table class="MsoTableGrid custom-table" border="0" cellspacing="0" cellpadding="0"> 
     <tr> 
      <td rowspan="2" class="acenter" width="39.03%"><p style="text-align:center">Tephritid species</p></td> 
      <td class="custom-bottom-td acenter" width="50.33%" colspan="3"><p style="text-align:center">High Guinea Savannah AEZ</p></td> 
      <td class="custom-bottom-td acenter" width="53.12%" colspan="2"><p style="text-align:center">Sudano Sahelian Savannah AEZ</p></td> 
     </tr> 
     <tr> 
      <td class="custom-bottom-td custom-top-td acenter" width="15.36%"><p style="text-align:center">Malang</p></td> 
      <td class="custom-bottom-td custom-top-td acenter" width="18.35%"><p style="text-align:center">Manwi</p></td> 
      <td class="custom-bottom-td custom-top-td acenter" width="16.62%"><p style="text-align:center">Marza</p></td> 
      <td class="custom-bottom-td custom-top-td acenter" width="27.26%"><p style="text-align:center">Mbé</p></td> 
      <td class="custom-bottom-td custom-top-td acenter" width="25.86%"><p style="text-align:center">Ngong</p></td> 
     </tr> 
     <tr> 
      <td class="custom-top-td acenter" width="39.03%"><p style="text-align:center">Bactrocera dorsalis</p></td> 
      <td class="custom-top-td acenter" width="15.36%"><p style="text-align:center">94.23</p></td> 
      <td class="custom-top-td acenter" width="18.35%"><p style="text-align:center">93.49</p></td> 
      <td class="custom-top-td acenter" width="16.62%"><p style="text-align:center">96.30</p></td> 
      <td class="custom-top-td acenter" width="27.26%"><p style="text-align:center">66.42</p></td> 
      <td class="custom-top-td acenter" width="25.86%"><p style="text-align:center">37.22</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Ceratitis anonae</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">0.01</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">2.88</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">0.02</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Ceratitis bremii</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">0.55</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">0.08</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Ceratitis cosyra</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">1.82</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">0.27</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">1.23</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">20.02</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">28.62</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Ceratitis fasciventris</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">1.35</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">2.27</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">0.23</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">1.14</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">2.88</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Ceratitis punctata</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">0</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">0.07</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">0</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Dacus bivittatus</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">0.61</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">1.49</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">0.96</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">1.10</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">0.37</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Dacus punctatifrons</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">1.81</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">1.49</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">1.13</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">5.59</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">0.67</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Zeugodacus cucurbitae</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">0.19</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">0.98</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">0.13</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">2.23</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">30.15</p></td> 
     </tr> 
     <tr> 
      <td class="acenter" width="39.03%"><p style="text-align:center">Total abundance</p></td> 
      <td class="acenter" width="15.36%"><p style="text-align:center">44058</p></td> 
      <td class="acenter" width="18.35%"><p style="text-align:center">28598</p></td> 
      <td class="acenter" width="16.62%"><p style="text-align:center">22673</p></td> 
      <td class="acenter" width="27.26%"><p style="text-align:center">7213</p></td> 
      <td class="acenter" width="25.86%"><p style="text-align:center">6432</p></td> 
     </tr> 
    </table>
   </table-wrap>
   <table-wrap id="table2">
    <label>
     <xref ref-type="table" rid="table2">
      Table 2
     </xref></label>
    <caption>
     <title>
      <xref ref-type="bibr" rid="scirp.144174-"></xref>Table S2. Mean number of tephritid individuals ± standard deviation captured per trap per sampled day, with minimum and maximum values in parentheses, in five orchards located in two agroecological zones (AEZs) of Cameroon: two orchards in AEZ 1 (Mbé and Ngong) and three orchards in AEZ 2 (Marza, Manwi, and Malang). Values were obtained for four male lures (CU: cue-lure; ME: methyl eugenol; TA: trimedlure; TR: terpinyl acetate), during the 2021-2022 sampling period. Superscript letters indicate statistically significant differences within each community between male lures, while numbers indicate significant differences within each male lure between communities, both based on post hoc tests (p &lt; 0.05).</title>
    </caption>
    <table class="MsoTableGrid custom-table" border="0" cellspacing="0" cellpadding="0"> 
     <tr> 
      <td rowspan="2" class="acenter"><p style="text-align:center">Male lures</p></td> 
      <td class="custom-bottom-td acenter" colspan="3"><p style="text-align:center">High Guinea Savannah (AEZ 2)</p></td> 
      <td class="custom-bottom-td acenter" colspan="2"><p style="text-align:center">Sudano-Sahelian Savannah (AEZ 1)</p></td> 
     </tr> 
     <tr> 
      <td class="custom-bottom-td custom-top-td acenter"><p style="text-align:center">Malang</p></td> 
      <td class="custom-bottom-td custom-top-td acenter"><p style="text-align:center">Manwi</p></td> 
      <td class="custom-bottom-td custom-top-td acenter"><p style="text-align:center">Marza</p></td> 
      <td class="custom-bottom-td custom-top-td acenter"><p style="text-align:center">Mbé</p></td> 
      <td class="custom-bottom-td custom-top-td acenter"><p style="text-align:center">Ngong</p></td> 
     </tr> 
     <tr> 
      <td class="custom-top-td acenter"><p style="text-align:center">CU</p></td> 
      <td class="custom-top-td acenter"><p style="text-align:center">14.46 ± 4.73</p><p style="text-align:center">(0 - 400) [c2]</p></td> 
      <td class="custom-top-td acenter"><p style="text-align:center">15.01 ± 1.95</p><p style="text-align:center">(1 - 91) [d3]</p></td> 
      <td class="custom-top-td acenter"><p style="text-align:center">8.49 ± 1.52</p><p style="text-align:center">(1 - 64) [b2]</p></td> 
      <td class="custom-top-td acenter"><p style="text-align:center">3.69 ± 0.33</p><p style="text-align:center">(0 - 33) [a2]</p></td> 
      <td class="custom-top-td acenter"><p style="text-align:center">16.76 ± 4.64</p><p style="text-align:center">(0 - 273) [e2]</p></td> 
     </tr> 
     <tr> 
      <td class="acenter"><p style="text-align:center">ME</p></td> 
      <td class="acenter"><p style="text-align:center">1415.45 ± 280.99</p><p style="text-align:center">(1 - 4807) [e4]</p></td> 
      <td class="acenter"><p style="text-align:center">1058.04 ± 198.31</p><p style="text-align:center">(1 - 4123) [d4]</p></td> 
      <td class="acenter"><p style="text-align:center">772.57 ± 147.54</p><p style="text-align:center">(1 - 3226) [c4]</p></td> 
      <td class="acenter"><p style="text-align:center">36.58 ± 6.56</p><p style="text-align:center">(0 - 377) [b4]</p></td> 
      <td class="acenter"><p style="text-align:center">22.17 ± 4.88</p><p style="text-align:center">(0 - 317) [a4]</p></td> 
     </tr> 
     <tr> 
      <td class="acenter"><p style="text-align:center">TA</p></td> 
      <td class="acenter"><p style="text-align:center">20.62 ± 3.34</p><p style="text-align:center">(1 - 134) [e3]</p></td> 
      <td class="acenter"><p style="text-align:center">14.72 ± 2.96</p><p style="text-align:center">(1 - 96) [c2]</p></td> 
      <td class="acenter"><p style="text-align:center">9.62 ± 2.80</p><p style="text-align:center">(1 - 94) [a3]</p></td> 
      <td class="acenter"><p style="text-align:center">11.01 ± 2.09</p><p style="text-align:center">(0 - 159) [b3]</p></td> 
      <td class="acenter"><p style="text-align:center">17.04 ± 3.92</p><p style="text-align:center">(0 - 250) [d3]</p></td> 
     </tr> 
     <tr> 
      <td class="acenter"><p style="text-align:center">TR</p></td> 
      <td class="acenter"><p style="text-align:center">8.38 ± 1.48</p><p style="text-align:center">(1 - 60) [e1]</p></td> 
      <td class="acenter"><p style="text-align:center">3.62 ± 0.71</p><p style="text-align:center">(1 - 24) [c1]</p></td> 
      <td class="acenter"><p style="text-align:center">3.68 ± 0.68</p><p style="text-align:center">(1 - 10) [d1]</p></td> 
      <td class="acenter"><p style="text-align:center">1.03 ± 0.23</p><p style="text-align:center">(0 - 12) [b1]</p></td> 
      <td class="acenter"><p style="text-align:center">0.21 ± 0.07</p><p style="text-align:center">(0 - 4) [a1]</p></td> 
     </tr> 
    </table>
   </table-wrap>
  </sec>
 </body><back>
  <ref-list>
   <title>References</title>
   <ref id="scirp.144174-ref1">
    <label>1</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     He, Y., Xu, Y. and Chen, X. (2023) Biology, Ecology and Management of Tephritid Fruit Flies in China: A Review. Insects, 14, Article 196. &gt;https://doi.org/10.3390/insects14020196
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref2">
    <label>2</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Mokam, D.G., Atougour, N., Tadu, Z., Aléné, D.C., Awono, E., Lontsi Tapeo, S., et al. (2024) Susceptibility of Mangifera indica (Sapindales: Anacardiaceae) Cultivars to Fruit Flies (Diptera: Tephritidae) in 2 Agroecological Zones of Cameroon. Journal of Insect Science, 24, 9. &gt;https://doi.org/10.1093/jisesa/ieae027
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref3">
    <label>3</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Scolari, F., Valerio, F., Benelli, G., Papadopoulos, N.T. and Vaníčková, L. (2021) Tephritid Fruit Fly Semiochemicals: Current Knowledge and Future Perspectives. Insects, 12, Article 408. &gt;https://doi.org/10.3390/insects12050408
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref4">
    <label>4</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Hafsi, A., Facon, B., Ravigné, V., Chiroleu, F., Quilici, S., Chermiti, B., et al. (2016) Host Plant Range of a Fruit Fly Community (Diptera: Tephritidae): Does Fruit Composition Influence Larval Performance? BMC Ecology, 16, Article No. 40. &gt;https://doi.org/10.1186/s12898-016-0094-8
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref5">
    <label>5</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Dias, N.P., Zotti, M.J., Montoya, P., Carvalho, I.R. and Nava, D.E. (2018) Fruit Fly Management Research: A Systematic Review of Monitoring and Control Tactics in the World. Crop Protection, 112, 187-200. &gt;https://doi.org/10.1016/j.cropro.2018.05.019
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref6">
    <label>6</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Metcalf, R.L. (1990) Chemical Ecology of Dacinae Fruit Flies (Diptera: Tephritidae). Annals of the Entomological Society of America, 83, 1017-1030. &gt;https://doi.org/10.1093/aesa/83.6.1017
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref7">
    <label>7</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Okolle, J.N. and Ntonifor, N.N. (2005) Field Ovipositional Behavior and Laboratory Studies on Development of Dacus punctatifrons (Diptera: Tephritidae) on Tomato. Insect Science, 12, 393-398. &gt;https://doi.org/10.1111/j.1005-295x.2005.00049.x
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref8">
    <label>8</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Migani, V., Ekesi, S. and Hoffmeister, T.S. (2013) Physiology vs. Environment: What Drives Oviposition Decisions in Mango Fruit Flies (Bactrocera invadens and Ceratitis cosyra)? Journal of Applied Entomology, 138, 395-402. &gt;https://doi.org/10.1111/jen.12038
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref9">
    <label>9</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Mokam, D.G., Djiéto-Lordon, C., Bilong Bilong, C. and Lumaret, J. (2018) Host Susceptibility and Pest Status of Fruit Flies (Diptera: Tephritidae) Attacking Cucurbits in Two Agroecological Zones of Cameroon, Central Africa. African Entomology, 26, 317-322. &gt;https://doi.org/10.4001/003.026.0317
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref10">
    <label>10</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Duyck, P., Jourdan, H. and Mille, C. (2022) Sequential Invasions by Fruit Flies (Diptera: Tephritidae) in Pacific and Indian Ocean Islands: A Systematic Review. Ecology and Evolution, 12, e8880. &gt;https://doi.org/10.1002/ece3.8880
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref11">
    <label>11</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Goergen, G., Vayssières, J., Gnanvossou, D. and Tindo, M. (2011) Bactrocera invadens (Diptera: Tephritidae), a New Invasive Fruit Fly Pest for the Afrotropical Region: Host Plant Range and Distribution in West and Central Africa. Environmental Entomology, 40, 844-854. &gt;https://doi.org/10.1603/en11017
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref12">
    <label>12</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Rwomushana, I., Ekesi, S., Gordon, I. and Ogol, C.K.P.O. (2008) Host Plants and Host Plant Preference Studies for Bactrocera invadens (Diptera: Tephritidae) in Kenya, a New Invasive Fruit Fly Species in Africa. Annals of the Entomological Society of America, 101, 331-340. &gt;https://doi.org/10.1603/0013-8746(2008)101[331:hpahpp]2.0.co;2
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref13">
    <label>13</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Qin, Y., Paini, D.R., Wang, C., Fang, Y. and Li, Z. (2015) Global Establishment Risk of Economically Important Fruit Fly Species (Tephritidae). PLOS ONE, 10, e0116424. &gt;https://doi.org/10.1371/journal.pone.0116424
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref14">
    <label>14</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Nanga Nanga, S., Hanna, R., Fotso Kuate, A., Fiaboe, K.K.M., Nchoutnji, I., Ndjab, M., et al. (2022) Tephritid Fruit Fly Species Composition, Seasonality, and Fruit Infestations in Two Central African Agro-Ecological Zones. Insects, 13, Article 1045. &gt;https://doi.org/10.3390/insects13111045
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref15">
    <label>15</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Ezechiel, A., Didi, G.M., Atougour, N., Albert, N. and Champlain, D. (2025) Diversity of Fruit Flies (Diptera: Tephritidae) Attacking Psidium guajava L. (Myrtaceae) in Two Agro-Ecological Zones of Northern Cameroon. African Journal of Agricultural Research, 21, 355-367. &gt;https://doi.org/10.5897/ajar2024.16850
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref16">
    <label>16</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Ndzana Abanda, F., Quilici, S., Vayssières, J., Kouodiekong, L. and Woin, N. (2008) Inventaire des espèces de mouches des fruits sur goyave dans la région de Yaoundé au Cameroun. Fruits, 63, 19-26. &gt;https://doi.org/10.1051/fruits:2007041
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref17">
    <label>17</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Desneux, N., Decourtye, A. and Delpuech, J. (2007) The Sublethal Effects of Pesticides on Beneficial Arthropods. Annual Review of Entomology, 52, 81-106. &gt;https://doi.org/10.1146/annurev.ento.52.110405.091440
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref18">
    <label>18</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Isman, M.B. (2006) Botanical Insecticides, Deterrents, and Repellents in Modern Agriculture and an Increasingly Regulated World. Annual Review of Entomology, 51, 45-66. &gt;https://doi.org/10.1146/annurev.ento.51.110104.151146
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref19">
    <label>19</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Jing, T., Zhang, Y., Dou, W., Jiang, X. and Wang, J. (2019) First Insights into the Intrapuparial Development of Bactrocera dorsalis (Hendel): Application in Predicting Emergence Time for Tephritid Fly Control. Insects, 10, Article 283. &gt;https://doi.org/10.3390/insects10090283
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref20">
    <label>20</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Shaurub, E.H. (2022) Review of Entomopathogenic Fungi and Nematodes as Biological Control Agents of Tephritid Fruit Flies: Current Status and a Future Vision. Entomologia Experimentalis et Applicata, 171, 17-34. &gt;https://doi.org/10.1111/eea.13244
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref21">
    <label>21</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Ebi, C. (2020) Effectiveness of Selected Parapheromones for Mass Trapping of Mango Fruit Fly, Ceratitis cosyra Walker and Oriental Fruit Fly, Bactrocera dorsalis Hendel (Diptera: Tephritidae) on Mango. Nigerian Journal of Entomology, 36, 118-129. &gt;https://doi.org/10.36108/nje/0202/63.01.41
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref22">
    <label>22</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Ndayizeye, L., Nzigidahera, B. and Theron, C.D. (2017) Effect of Parapheromones on the Capture of Fruit Flies (Diptera: Tephritidae) in Burundi. Journal of Agricultural Science and Technology A, 7, 413-425. &gt;https://doi.org/10.17265/2161-6256/2017.06.007
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref23">
    <label>23</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Wee, S. and Clarke, A.R. (2020) Male-Lure Type, Lure Dosage, and Fly Age at Feeding All Influence Male Mating Success in Jarvis’ Fruit Fly. Scientific Reports, 10, Article No. 15004. &gt;https://doi.org/10.1038/s41598-020-72209-x
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref24">
    <label>24</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Shelly, T., Fezza, T., Cook, P. and Cook, D. (2024) Attractiveness, Longevity, and Release Rates of Multilure Wafers for Trapping Males of the Oriental Fruit Fly and Melon Fly (Diptera: Tephritidae). Journal of Insect Science, 24, Article 24. &gt;https://doi.org/10.1093/jisesa/ieae095
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref25">
    <label>25</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Awarikabey, E.N., Afun, J.V.K., Billah, M.K. and Osekre, E.A. (2025) Diversity, Damage and Pheromone Specificity of Fruit Flies in the Forest-Savanna Transition Zone of Ghana. Bulletin of Entomological Research, 115, 155-165. &gt;https://doi.org/10.1017/s0007485324000750
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref26">
    <label>26</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Geurts, K., Mwatawala, M. and Meyer, M.D. (2012) Indigenous and Invasive Fruit Fly Diversity along an Altitudinal Transect in Eastern Central Tanzania. Journal of Insect Science, 12, 1-18. &gt;https://doi.org/10.1673/031.012.1201
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref27">
    <label>27</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Ekesi, S. and Billah, M.K. (2007) A Field Guide to the Management of Economically Important Tephritid Fruit Flies in Africa. ICIPE.
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref28">
    <label>28</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Shelly, T.E., Fezza, T.J. and Martin, R.M. (2024) Field Longevity of Methyl Eugenol and Cue-Lure Plugs and Associated Insecticidal Strips: Captures of Bactrocera dorsalis and Zeugodacus cucurbitae (Diptera: Tephritidae) in Hawaii. Environmental Entomology, 53, 782-788. &gt;https://doi.org/10.1093/ee/nvae064
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref29">
    <label>29</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Tinkeu, L., Ladang, D., Vayssieres, J. and Lyannaz, J. (2011) Diversité des espèces de mouches des fruits (Diptera: Tephritidae) dans un verger mixte dans la localité de Malang (Ngaoundéré, Cameroun). International Journal of Biological and Chemical Sciences, 4, 1425-1434. &gt;https://doi.org/10.4314/ijbcs.v4i5.65530
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref30">
    <label>30</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     White, I. and Elson-Harris, M. (2004) Fruit Flies of Economic Significance: Their Identification and Bionomic. CABI.
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref31">
    <label>31</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Mokam, D.G., Djiéto-Lordon, C. and Bilong Bilong, C. (2014) Patterns of Species Richness and Diversity of Insects Associated with Cucurbit Fruits in the Southern Part of Cameroon. Journal of Insect Science, 14, Article 248. &gt;https://doi.org/10.1093/jisesa/ieu110
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref32">
    <label>32</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Zou, Y., Zhao, P. and Axmacher, J.C. (2023) Estimating Total Species Richness: Fitting Rarefaction by Asymptotic Approximation. Ecosphere, 14, e4363. &gt;https://doi.org/10.1002/ecs2.4363
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref33">
    <label>33</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Avolio, M.L., Carroll, I.T., Collins, S.L., Houseman, G.R., Hallett, L.M., Isbell, F., et al. (2019) A Comprehensive Approach to Analyzing Community Dynamics Using Rank Abundance Curves. Ecosphere, 10, e02881. &gt;https://doi.org/10.1002/ecs2.2881
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref34">
    <label>34</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Morris, E.K., Caruso, T., Buscot, F., Fischer, M., Hancock, C., Maier, T.S., et al. (2014) Choosing and Using Diversity Indices: Insights for Ecological Applications from the German Biodiversity Exploratories. Ecology and Evolution, 4, 3514-3524. &gt;https://doi.org/10.1002/ece3.1155
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref35">
    <label>35</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Roswell, M., Dushoff, J. and Winfree, R. (2021) A Conceptual Guide to Measuring Species Diversity. Oikos, 130, 321-338. &gt;https://doi.org/10.1111/oik.07202
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref36">
    <label>36</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Verma, V. and Aggarwal, R.K. (2020) A Comparative Analysis of Similarity Measures Akin to the Jaccard Index in Collaborative Recommendations: Empirical and Theoretical Perspective. Social Network Analysis and Mining, 10, Article No. 43. &gt;https://doi.org/10.1007/s13278-020-00660-9
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref37">
    <label>37</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Vargas, R.I., Leblanc, L., Piñero, J.C. and Hoffman, K.M. (2014) Male Annihilation, Past, Present, and Future. In: Shelly, T., Epsky, N., Jang, E.B.J., Reyes-Flores and Vargas, R., Eds., Trapping and the Detection, Control, and Regulation of Tephritid Fruit Flies, Springer Netherlands, 493-511. &gt;https://doi.org/10.1007/978-94-017-9193-9_14
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref38">
    <label>38</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Vayssières, J.-., De Meyer, M., Ouagoussounon, I., Sinzogan, A., Adandonon, A., Korie, S., et al. (2015) Seasonal Abundance of Mango Fruit Flies (Diptera: Tephritidae) and Ecological Implications for Their Management in Mango and Cashew Orchards in Benin (Centre&amp;North). Journal of Economic Entomology, 108, 2213-2230. &gt;https://doi.org/10.1093/jee/tov143
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref39">
    <label>39</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Manrakhan, A., Venter, J.H. and Hattingh, V. (2015) The Progressive Invasion of Bactrocera dorsalis (Diptera: Tephritidae) in South Africa. Biological Invasions, 17, 2803-2809. &gt;https://doi.org/10.1007/s10530-015-0923-2
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref40">
    <label>40</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Mutamiswa, R., Nyamukondiwa, C., Chikowore, G. and Chidawanyika, F. (2021) Overview of Oriental Fruit Fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) in Africa: From Invasion, Bio-Ecology to Sustainable Management. Crop Protection, 141, Article ID: 105492. &gt;https://doi.org/10.1016/j.cropro.2020.105492
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref41">
    <label>41</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Vargas, R.I., Leblanc, L., Harris, E.J. and Manoukis, N.C. (2012) Regional Suppression of Bactrocera Fruit Flies (Diptera: Tephritidae) in the Pacific through Biological Control and Prospects for Future Introductions into Other Areas of the World. Insects, 3, 727-742. &gt;https://doi.org/10.3390/insects3030727
    </mixed-citation>
   </ref>
   <ref id="scirp.144174-ref42">
    <label>42</label>
    <mixed-citation publication-type="other" xlink:type="simple">
     Manoukis, N.C., Vargas, R.I., Carvalho, L., Fezza, T., Wilson, S., Collier, T., et al. (2019) A Field Test on the Effectiveness of Male Annihilation Technique against Bactrocera dorsalis (Diptera: Tephritidae) at Varying Application Densities. PLOS ONE, 14, e0213337. &gt;https://doi.org/10.1371/journal.pone.0213337
    </mixed-citation>
   </ref>
  </ref-list>
 </back>
</article>