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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">90</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:4EBD0208-8328-5D69-8C44-EC50939C0967</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:52E200AF-408A-423E-8797-5BFC4FDE0F99</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Herpetozoa</journal-title>
        <abbrev-journal-title xml:lang="en">Herpetozoa</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1013-4425</issn>
      <issn pub-type="epub">2682-955X</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/herpetozoa.39.e178597</article-id>
      <article-id pub-id-type="publisher-id">178597</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Short Communication</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Anguidae</subject>
          <subject>Squamata</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Evolutionary biology</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Central America and the Caribbean</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>It's not all about size: sexual size dimorphism and caudal autotomy of Campbell's Alligator Lizard, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> Brodie &amp; Savage, 1993</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Torres-Almazán</surname>
            <given-names>Mónica</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0009-1995-1467</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Schrei Godoy</surname>
            <given-names>Klaus Thomas</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0003-1373-2139</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Lock</surname>
            <given-names>Brad A.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0000-4402-6251</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Escobedo-Galván</surname>
            <given-names>Armando H.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-1514-1255</uri>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>González-Maya</surname>
            <given-names>José F.</given-names>
          </name>
          <email xlink:type="simple">jfgonzalezmaya@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-8942-5157</uri>
          <xref ref-type="aff" rid="A4">4</xref>
          <xref ref-type="aff" rid="A5">5</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Fundación para las Especies Amenazadas de Guatemala – FUNDESGUA, Lote 21 A Labor de Castilla II zona 9 de Mixco, Guatemala, Guatemala</addr-line>
        <institution>Fundación para las Especies Amenazadas de Guatemala – FUNDESGUA</institution>
        <addr-line content-type="city">Guatemala</addr-line>
        <country>Guatemala</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Oklahoma City Zoo and Botanical Garden, 2000 Remington Place, Oklahoma City, OK 73111, USA</addr-line>
        <institution>Oklahoma City Zoo and Botanical Garden</institution>
        <addr-line content-type="city">Oklahoma City</addr-line>
        <country>United States of America</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Centro Universitario de la Costa, Universidad de Guadalajara, Av. Universidad 203, 48280 Puerto Vallarta, Jalisco, Mexico</addr-line>
        <institution>Centro Universitario de la Costa, Universidad de Guadalajara</institution>
        <addr-line content-type="city">Puerto Vallarta</addr-line>
        <country>Mexico</country>
        <uri content-type="ror">https://ror.org/043xj7k26</uri>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">Área Académica en Biología de la Conservación, Departamento de Ciencias Ambientales, DCBS, Universidad Autónoma Metropolitana Unidad Lerma, Av. De las Garzas 10, Lerma de Villada, Estado de México, 52005, Mexico</addr-line>
        <institution>Área Académica en Biología de la Conservación, Departamento de Ciencias Ambientales, DCBS, Universidad Autónoma Metropolitana Unidad Lerma</institution>
        <addr-line content-type="city">Lerma de Villada</addr-line>
        <country>Mexico</country>
      </aff>
      <aff id="A5">
        <label>5</label>
        <addr-line content-type="verbatim">ProCAT Colombia/Costa Rica, Calle 118 # 16-61, Of. 602, Bogotá, Colombia</addr-line>
        <institution>ProCAT Colombia/Costa Rica</institution>
        <addr-line content-type="city">Bogotá</addr-line>
        <country>Colombia</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: José F. González-Maya (<email xlink:type="simple">jfgonzalezmaya@gmail.com</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Johannes Foufopoulos</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>09</day>
        <month>09</month>
        <year>2026</year>
      </pub-date>
      <volume>39</volume>
      <fpage>367</fpage>
      <lpage>372</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/E557867D-FF63-5DA9-A524-C6114092B676">E557867D-FF63-5DA9-A524-C6114092B676</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/0E847863-F484-4C69-BF30-6750EE3A49A1">0E847863-F484-4C69-BF30-6750EE3A49A1</uri>
      <history>
        <date date-type="received">
          <day>20</day>
          <month>11</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>16</day>
          <month>06</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Mónica Torres-Almazán, Klaus Thomas Schrei Godoy, Brad A. Lock, Armando H. Escobedo-Galván, José F. González-Maya</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <self-uri content-type="zoobank" xlink:type="simple">https://zoobank.org/0E847863-F484-4C69-BF30-6750EE3A49A1</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>Sexual dimorphism and caudal autotomy are poorly documented in the lizard genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic>. We analyzed sexual size dimorphism and caudal autotomy in 54 adult Campbell’s Alligator Lizards (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic>) from Jalapa, Guatemala. We measured snout–vent length (<abbrev xlink:title="snout–vent length">SVL</abbrev>), head width (<abbrev xlink:title="head width">HW</abbrev>), and body mass (<abbrev xlink:title="body mass">BM</abbrev>) for 29 males and 25 females. While both sexes showed similar <abbrev xlink:title="snout–vent length">SVL</abbrev> and <abbrev xlink:title="body mass">BM</abbrev>, the males had significantly broader heads than females relative to body size and body mass. In addition, caudal autotomy was similar between males and females. This monomorphism in body size, coupled with dimorphism in head width, suggests that sexual selection, likely driven by male–male combat or the need to grasp females during mating, acts strongly on male head morphology in this species.</p>
      </abstract>
      <kwd-group>
        <label>Key Words</label>
        <kwd>Allometry</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="subfamily" reg="Gerrhonotinae">Gerrhonotinae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>Guatemala</kwd>
        <kwd>head morphology</kwd>
        <kwd>intrasexual selection</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p>Sexual dimorphism in body traits is widespread across animal species. Lizards serve as models for investigating the evolutionary and ecological consequences of sexual size dimorphism (<abbrev xlink:title="sexual size dimorphism">SSD</abbrev>) across traits including body size, head dimensions, and limb length (<xref ref-type="bibr" rid="B32">Tinkle and Dunham 1986</xref>; <xref ref-type="bibr" rid="B19">Jiménez-Arcos et al. 2017</xref>; <xref ref-type="bibr" rid="B22">Liang et al. 2021</xref>), providing insights into relative growth and form of morphological traits (<xref ref-type="bibr" rid="B27">Scharf and Meiri 2013</xref>; <xref ref-type="bibr" rid="B14">Goldberg et al. 2024</xref>). Males often exhibit proportionately larger heads due to male–male agonistic behaviours (<xref ref-type="bibr" rid="B30">Shine 1989</xref>), while females may show longer trunk lengths related to offspring production (<xref ref-type="bibr" rid="B27">Scharf and Meiri 2013</xref>; <xref ref-type="bibr" rid="B26">Pérez-Quintero et al. 2019</xref>). These variations can lead to hyperallometric or hypoallometric relationships between traits and body size, favouring intraspecific phenotypic divergence driven by different reproductive roles between sexes (<xref ref-type="bibr" rid="B9">Cox et al. 2007</xref>).</p>
      <p>The alligator lizards of the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Gerrhonotinae">Gerrhonotinae</tp:taxon-name-part></tp:taxon-name> currently comprise five genera distributed across North and Central America (<xref ref-type="bibr" rid="B37">Vitt and Caldwell 2014</xref>; <xref ref-type="bibr" rid="B3">Blair et al. 2022</xref>). Historically, the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> Gray, 1838, was characterized as an exclusively arboreal lineage endemic to Mesoamerican highland forests, ranging from central Mexico to western Panama (<xref ref-type="bibr" rid="B8">Clause et al. 2024</xref>). However, recent genomic evidence reveals that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> is nested within, or closely allied with, terrestrial species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesaspis">Mesaspis</tp:taxon-name-part></tp:taxon-name></italic> Cope, 1877, rendering both genera non-monophyletic under their current definitions (<xref ref-type="bibr" rid="B16">Gutiérrez-Rodríguez et al. 2021</xref>). Consequently, the expanded circumscription of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> now includes 39 species; while the majority remain arboreal, several taxa exhibit terrestrial habits (<xref ref-type="bibr" rid="B33">Uetz et al. 2025</xref>). Most of the arboreal species are characterized by a very striking colour pattern, characterized by bright colours in shades of blue, green, cream, brown, and black, which allow them to blend in with the epiphytic plants in their habitat (<xref ref-type="bibr" rid="B6">Campbell and Frost 1993</xref>). At present, a comprehensive understanding of ecological and life-history traits of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> populations remains limited and anecdotal, where ecological studies are mainly concentrated on <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="taeniata">taeniata</tp:taxon-name-part></tp:taxon-name></italic> (Wiegmann, 1828) (<xref ref-type="bibr" rid="B35">Villamar-Duque et al. 2019</xref>; <xref ref-type="bibr" rid="B12">Fierro-Estrada et al. 2023</xref>). Overall, there has been little analysis of morphological traits in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> species, particularly regarding sexual size dimorphism have been conducted in less than 15% of the 39 described <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> species (<xref ref-type="bibr" rid="B21">Lemos-Espinal et al. 2001</xref>; <xref ref-type="bibr" rid="B15">González-Porter et al. 2015</xref>; <xref ref-type="bibr" rid="B10">Cruz-Elizalde et al. 2020</xref>).</p>
      <p>In this study, we analysed the sexual size dimorphism and caudal autotomy of Campbell’s Alligator Lizard, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> Brodie &amp; Savage, 1993, a Critically Endangered arboreal lizard micro-endemic to the pine–oak and cloud forests of southeastern Guatemala. The species inhabits old-growth oak forests dominated by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Quercus">Quercus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="peduncularis">peduncularis</tp:taxon-name-part></tp:taxon-name></italic> Née, 1801 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Quercus">Q.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brachystachys">brachystachys</tp:taxon-name-part></tp:taxon-name></italic> Bentham, 1842 where it shows a strong association with the epiphytic bromeliad <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tillandsia">Tillandsia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="usneoides">usneoides</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1753) which serves as its primary microhabitat (<xref ref-type="bibr" rid="B1">Ariano-Sánchez and Torres-Almazán 2010</xref>). We predicted sexual dimorphism in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic>, with allometric differences between sexes. Specifically, we expected males to have broader heads than females, consistent with previous studies (<xref ref-type="bibr" rid="B21">Lemos-Espinal et al. 2001</xref>; <xref ref-type="bibr" rid="B35">Villamar-Duque et al. 2019</xref>; <xref ref-type="bibr" rid="B15">González-Porter et al. 2015</xref>; <xref ref-type="bibr" rid="B10">Cruz-Elizalde et al. 2020</xref>).</p>
    </sec>
    <sec sec-type="materials|methods" id="sec2">
      <title>Methods and materials</title>
      <p>We conducted diurnal surveys near Potrero Carrillo, Department of Jalapa, Guatemala (Fig. <xref ref-type="fig" rid="F1">1</xref>). Between 2014 and 2016, we captured 54 individuals (29 adult males, 25 adult females) throughout the year. Fieldwork and specimen collection were performed under Research Licenses No. 036/2014 and No. 010/2015, and Collection Permits Serie A No. 000972 and No. 002013. After capturing the individuals, we recorded the following morphological measurements with an electronic caliper: snout–vent length (<abbrev xlink:title="snout–vent length">SVL</abbrev>) of individuals measured from the tip of the snout to the anterior margin of the cloaca; head width (<abbrev xlink:title="head width">HW</abbrev>), the maximum width of the head, and body mass (<abbrev xlink:title="body mass">BM</abbrev>) measured using an analytical balance (± 0.01 g; <xref ref-type="bibr" rid="B10">Cruz-Elizalde et al. 2020</xref>). To ensure reproductive maturity, only adult individuals with a snout–vent length (<abbrev xlink:title="snout–vent length">SVL</abbrev>) ≥ 105 mm were included in the analyses. This threshold is conservative compared to the original species description, which reports adult sizes beginning at approximately 93–94 mm <abbrev xlink:title="snout–vent length">SVL</abbrev> (Brodie and Savage 1993). The sex of individuals was determined by manually everting hemipenes. We also recorded whether the individual had undergone caudal autotomy. Finally, each lizard was individually identified utilizing a passive integrated transponder tag (12.5 × 2.1 mm, 118 mg; Avid FriendChip PIT tags, Avid Identification Systems) before releasing it at the capture site. On each new capture, we first passed the PIT tag reader over different parts of the body to avoid duplicate measurements.</p>
      <fig id="F1">
        <object-id content-type="doi">10.3897/herpetozoa.39.e178597.figure1</object-id>
        <object-id content-type="arpha">F2581065-EC31-5336-A7B8-F6FEA5391F0F</object-id>
        <label>Figure 1.</label>
        <caption>
          <p>Location of the field locality were <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> individuals were captured in Potrero Carrillo and distribution range according to IUCN Red List of Threatened Species (<xref ref-type="bibr" rid="B2">Ariano-Sánchez et al. 2013</xref>), department of Jalapa, Guatemala.</p>
        </caption>
        <graphic xlink:href="herpetozoa-39-367-g001.jpg" id="oo_1768375.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1768375</uri>
        </graphic>
      </fig>
      <p>We examined the allometric relationships between <abbrev xlink:title="snout–vent length">SVL</abbrev> and morphological traits (<abbrev xlink:title="head width">HW</abbrev> and <abbrev xlink:title="body mass">BM</abbrev>) for each sex. To test the isometric relation (null hypothesis, <xref ref-type="bibr" rid="B7">Charnov 1993</xref>), we considered a slope with a 95% confidence interval (<abbrev xlink:title="confidence interval">CI</abbrev>) including the expected value of 1.0 to show isometry, while slopes with a 95% <abbrev xlink:title="confidence interval">CI</abbrev> below or above the expected value were interpreted as hypoallometric or hyperallometric relationships, respectively (<xref ref-type="bibr" rid="B11">Fairbairn 1997</xref>). To estimate sexual size dimorphism, we initially conducted a t-student test to compare <abbrev xlink:title="snout–vent length">SVL</abbrev> between sexes. Subsequently, an analysis of covariance (<abbrev xlink:title="analysis of covariance">ANCOVA</abbrev>) was used to assess sexual dimorphism adjusted to body size using <abbrev xlink:title="snout–vent length">SVL</abbrev> as the covariate. We used a chi-squared test to determine differences in the caudal autotomy between sexes. We present mean values ± 1 standard deviation (<abbrev xlink:title="standard deviation">SD</abbrev>). All analyses were performed using PAST 5.1 (<xref ref-type="bibr" rid="B18">Hammer et al. 2001</xref>).</p>
    </sec>
    <sec sec-type="Results" id="sec3">
      <title>Results</title>
      <p>Of the 54 <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> analysed for this study, females exhibited a hypoallometric relationship for <abbrev xlink:title="snout–vent length">SVL</abbrev>–<abbrev xlink:title="head width">HW</abbrev> and <abbrev xlink:title="body mass">BM</abbrev>–<abbrev xlink:title="head width">HW</abbrev> (slope &lt; 1), while the <abbrev xlink:title="snout–vent length">SVL</abbrev>–<abbrev xlink:title="body mass">BM</abbrev> showed a hyperallometric relationship (slope &gt; 1; Table <xref ref-type="table" rid="T1">1</xref>). Meanwhile, for males, <abbrev xlink:title="snout–vent length">SVL</abbrev>–<abbrev xlink:title="body mass">BM</abbrev> were consistent with slope = 1; both <abbrev xlink:title="snout–vent length">SVL</abbrev>–<abbrev xlink:title="head width">HW</abbrev> and <abbrev xlink:title="body mass">BM</abbrev>–<abbrev xlink:title="head width">HW</abbrev> exhibited hypoallometry (Table <xref ref-type="table" rid="T1">1</xref>). The <abbrev xlink:title="snout–vent length">SVL</abbrev> of males ranged from 89.0 to 129.8 mm (mean 105.6 ± 10.7 mm <abbrev xlink:title="standard deviation">SD</abbrev>), while females exhibited <abbrev xlink:title="snout–vent length">SVL</abbrev> ranging from 90.0 to 122.0 mm (mean 106.1 ± 8.6 mm <abbrev xlink:title="standard deviation">SD</abbrev>). The sexes showed a similar body size (t<sub>1,52</sub> = 0.25, <italic>p</italic> = 0.79). Females and males showed similar <abbrev xlink:title="body mass">BM</abbrev> (F<sub>1,51</sub> = 1.05, <italic>p</italic> = 0.30), while males exhibited broader heads than females (F<sub>1,51</sub> = 59.70, <italic>p</italic> &lt; 0.01; Fig. <xref ref-type="fig" rid="F2">2</xref>). The <abbrev xlink:title="head width">HW</abbrev> of males ranged from 18.2 to 26.2 mm (mean 23.3 ± 1.9 mm <abbrev xlink:title="standard deviation">SD</abbrev>), while females exhibited <abbrev xlink:title="head width">HW</abbrev> ranging from 16.0 to 24.0 mm (mean 18.8 ± 1.8 mm <abbrev xlink:title="standard deviation">SD</abbrev>). The <abbrev xlink:title="body mass">BM</abbrev> of females ranged from 13.5 to 39.1 g (mean 27.3 ± 6.1 g <abbrev xlink:title="standard deviation">SD</abbrev>), and the <abbrev xlink:title="body mass">BM</abbrev> of males ranged from 15.4 to 45.0 g (mean 28.6 ± 7.1 g <abbrev xlink:title="standard deviation">SD</abbrev>). Both sexes showed a similar body mass (t<sub>1,52</sub> = 0.61, <italic>p</italic> = 0.53). The frequency of caudal autotomy was not different between males and females, 4.17% and 4.88%, respectively (χ<sup>2</sup> = 1.16, d.f. = 1, <italic>p</italic> = 0.28).</p>
      <fig id="F2">
        <object-id content-type="doi">10.3897/herpetozoa.39.e178597.figure2</object-id>
        <object-id content-type="arpha">0C665964-F69D-5887-ADD1-2ACB7B9FC178</object-id>
        <label>Figure 2.</label>
        <caption>
          <p>Female (<bold>a</bold>) and male (<bold>b</bold>) of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> captured at the study site in the locality of Potrero Carrillo, department of Jalapa, Guatemala.</p>
        </caption>
        <graphic xlink:href="herpetozoa-39-367-g002.jpg" id="oo_1768376.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1768376</uri>
        </graphic>
      </fig>
      <table-wrap id="T1" position="float" orientation="portrait">
        <label>Table 1.</label>
        <caption>
          <p>Allometric relations between body size and morphological traits of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic>. <abbrev xlink:title="no significant">N.S.</abbrev> = no significant.</p>
        </caption>
        <table>
          <tbody>
            <tr>
              <th rowspan="1" colspan="1">
                <bold>Independent variable</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>Dependent variable</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>n</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>R<sup>2</sup></bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>p</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>Slope (a)</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>95% CL</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>Result</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>Y intercept</bold>
              </th>
            </tr>
            <tr>
              <td rowspan="1" colspan="9">Females</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <abbrev xlink:title="snout–vent length">SVL</abbrev>
              </td>
              <td rowspan="1" colspan="1">Body Mass</td>
              <td rowspan="1" colspan="1">25</td>
              <td rowspan="1" colspan="1">0.49</td>
              <td rowspan="1" colspan="1">&lt;0.001</td>
              <td rowspan="1" colspan="1">2.08</td>
              <td rowspan="1" colspan="1">1.01–2.91</td>
              <td rowspan="1" colspan="1">Hyperallometry</td>
              <td rowspan="1" colspan="1">-2.78</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1"/>
              <td rowspan="1" colspan="1">Head Width</td>
              <td rowspan="1" colspan="1">25</td>
              <td rowspan="1" colspan="1">0.04</td>
              <td rowspan="1" colspan="1">0.289</td>
              <td rowspan="1" colspan="1">0.25</td>
              <td rowspan="1" colspan="1">-0.21–0.72</td>
              <td rowspan="1" colspan="1">
                <abbrev xlink:title="no significant">N.S.</abbrev>
              </td>
              <td rowspan="1" colspan="1">0.75</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <abbrev xlink:title="body mass">BM</abbrev>
              </td>
              <td rowspan="1" colspan="1">Head Width</td>
              <td rowspan="1" colspan="1">25</td>
              <td rowspan="1" colspan="1">0.04</td>
              <td rowspan="1" colspan="1">0.317</td>
              <td rowspan="1" colspan="1">0.08</td>
              <td rowspan="1" colspan="1">-0.22–0.24</td>
              <td rowspan="1" colspan="1">
                <abbrev xlink:title="no significant">N.S.</abbrev>
              </td>
              <td rowspan="1" colspan="1">1.15</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="9">Males</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <abbrev xlink:title="snout–vent length">SVL</abbrev>
              </td>
              <td rowspan="1" colspan="1">Body Mass</td>
              <td rowspan="1" colspan="1">29</td>
              <td rowspan="1" colspan="1">0.39</td>
              <td rowspan="1" colspan="1">&lt;0.001</td>
              <td rowspan="1" colspan="1">1.57</td>
              <td rowspan="1" colspan="1">0.75–2.46</td>
              <td rowspan="1" colspan="1">Isometry</td>
              <td rowspan="1" colspan="1">-1.74</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1"/>
              <td rowspan="1" colspan="1">Head Width</td>
              <td rowspan="1" colspan="1">29</td>
              <td rowspan="1" colspan="1">0.41</td>
              <td rowspan="1" colspan="1">&lt;0.001</td>
              <td rowspan="1" colspan="1">0.56</td>
              <td rowspan="1" colspan="1">0.26–0.85</td>
              <td rowspan="1" colspan="1">Hypoallometry</td>
              <td rowspan="1" colspan="1">0.19</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <abbrev xlink:title="body mass">BM</abbrev>
              </td>
              <td rowspan="1" colspan="1">Head Width</td>
              <td rowspan="1" colspan="1">29</td>
              <td rowspan="1" colspan="1">0.36</td>
              <td rowspan="1" colspan="1">&lt;0.001</td>
              <td rowspan="1" colspan="1">0.21</td>
              <td rowspan="1" colspan="1">0.11–0.31</td>
              <td rowspan="1" colspan="1">Hypoallometry</td>
              <td rowspan="1" colspan="1">1.03</td>
            </tr>
          </tbody>
        </table>
      </table-wrap>
    </sec>
    <sec sec-type="Discussion" id="sec4">
      <title>Discussion</title>
      <p>Females showed a positive relationship between body size and body mass, meaning that females have higher body weight than expected as body size increases, but we did not observe differences in weight compared to males. On the other hand, we not only observed a significant allometric relationship between body size and head width in males, but also wider heads compared to females, which suggests that head width in males is under intrasexual selective pressure. Our results support monomorphism in body size and body mass of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic>. Based on the literature and our results, the sexual size monomorphism in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> species may maximize reproductive performance at similar body sizes (<xref ref-type="bibr" rid="B35">Villamar-Duque et al. 2019</xref>; <xref ref-type="bibr" rid="B10">Cruz-Elizalde et al. 2020</xref>).</p>
      <p>Males of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> showed on average 19.3% wider heads than females. Some studies have also shown that males have wider heads than females; for example, 14.5% in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="monticola">monticola</tp:taxon-name-part></tp:taxon-name></italic> (Cope, 1878), 11.3% in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="graminea">graminea</tp:taxon-name-part></tp:taxon-name></italic> (Cope, 1864), and 10.2% in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="taeniata">taeniata</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B34">Vial and Stewart 1989</xref>; <xref ref-type="bibr" rid="B21">Lemos-Espinal et al. 2001</xref>; <xref ref-type="bibr" rid="B10">Cruz-Elizalde et al. 2020</xref>). However, in available data for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> obtained prior to our study, <xref ref-type="bibr" rid="B1">Ariano-Sánchez and Torres-Almazán (2010)</xref> reported an <abbrev xlink:title="head width">HW</abbrev> mean of 26.0 ± 1.9 mm for males and 16.1 ± 2.1 mm for females, which showed males had on average 38.0% wider heads than females. These observations could be related to courtship behaviors such as neck and head bites during the courtship period. <xref ref-type="bibr" rid="B17">Gvozdík and Van Damme (2003)</xref> observed that sexual dimorphism in head size among adult lacertilians arises from differential growth in the postorbital region of the head, which is more pronounced in males. This area serves as the insertion point for the jaw abductor muscles, which are responsible for bite force. Some studies have shown that sexual selection may favour larger heads in males (resulting in greater bite force) if differences in male–male combat ability translate into the capacity to maintain territories and win intrasexual agonistic encounters, thereby increasing access to females (<xref ref-type="bibr" rid="B36">Vitt 1983</xref>; <xref ref-type="bibr" rid="B20">Lappin et al. 2006</xref>). For instance, Formanowicz et al. (1990) examined the interaction patterns between and within sexes of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="vasconcelosii">vasconcelosii</tp:taxon-name-part></tp:taxon-name></italic> (Bocourt, 1872) under laboratory conditions. The authors reported nine behaviour patterns, of which bites from head to tail were the most frequent in both male–male and female–female interactions. We observed antagonistic encounters in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic>, mainly between individuals of the same sex, although occasionally also between sexes. This behavior is characterized by one individual chasing another, often culminating in biting between the contestants. These interactions were opportunistic observations in the wild, during which one individual was confirmed as a male; this consisted of a chase, and no biting was observed as the individuals quickly fled. Additional observations occurred during field handling in rare instances when individuals were temporarily placed together in the same collection bag, although individuals were normally kept separately. In these cases, encounters involved chasing and biting and appeared more common between individuals of the same sex. Animals were separated immediately to prevent injury. During mating, male <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">Abronia</tp:taxon-name-part></tp:taxon-name></italic> typically restrain females with a firm bite on the left or right side of the head. For example, <xref ref-type="bibr" rid="B28">Schmidt-Ballardo and Mendoza-Quijano (1999)</xref> observed a male grasping a female on the temporal region in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mixteca">mixteca</tp:taxon-name-part></tp:taxon-name></italic> for 8 hours. <xref ref-type="bibr" rid="B29">Schmidt-Ballardo et al. (2015)</xref> observed that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="deppii">deppii</tp:taxon-name-part></tp:taxon-name></italic> copulated for approximately 18 h. <xref ref-type="bibr" rid="B31">Solano-Zavaleta et al. (2017)</xref> observed captive male <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="antauges">antauges</tp:taxon-name-part></tp:taxon-name></italic> maintaining a mate-holding grip on the left side of the female’s head for nearly four hours. In comparison, similar mate-holding behavior in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Elgaria">Elgaria</tp:taxon-name-part></tp:taxon-name></italic> has been documented to persist for over 24 hours (Nguyen et al. 2020). This behavior suggests that larger-headed males may have a greater ability to control females, potentially increasing their reproductive success.</p>
      <p>Some studies have shown that the incidence of tail loss in males is likely due to intrasexual agonistic encounters (<xref ref-type="bibr" rid="B38">Vitt et al. 1974</xref>). We observed a low frequency of caudal autotomy in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic>, particularly when compared to species like <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="vasconcelosii">vasconcelosii</tp:taxon-name-part></tp:taxon-name></italic>, which has been reported to exhibit regenerated tails in 72.2% of captured individuals (Formanowicz et al. 1990). The low frequency of caudal autotomy observed in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Abronia">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="campbelli">campbelli</tp:taxon-name-part></tp:taxon-name></italic> may reflect several non-mutually exclusive factors, including reduced intraspecific aggression, variable population densities associated with its endangered status, or an inherently lower autotomy threshold, possibly because the prehensile tail represents a particularly valuable locomotor asset whose loss would be highly costly. Notably, low autotomy rates do not necessarily imply that antagonistic interactions are directed away from the tail. As <xref ref-type="bibr" rid="B25">Paﬁlis et al. (2019)</xref> and <xref ref-type="bibr" rid="B4">Brock et al. (2015)</xref> have shown, inducing tail loss in a competitor may itself be selectively advantageous, as it removes a valuable locomotor structure and could reduce the rival’s competitive ability. As <xref ref-type="bibr" rid="B23">Martin (1955)</xref> noted, the loss of this “fifth limb” significantly impairs arboreal performance, making its preservation essential for survival in the canopy. Ultimately, our findings highlight how morphological constraints and social behavior intersect in these endangered lizards, underscoring the urgent need for further natural history research to inform their conservation.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We are especially grateful to José Cristian Ramirez López, our field assistant, for his dedication, effort, and commitment throughout the fieldwork. We also thank the local communities and property owners who allowed us to conduct this study on their lands and helped us appreciate this remarkable species in its natural habitat.</p>
    </ack>
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    <sec sec-type="Additional information" id="sec5">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec6">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec7">
        <title>Ethical statement</title>
        <p>No ethical statement is reported. The permission to handle were conducted under Guatemalan National Council for Protected Areas (CONAP): collection permits Serie A No. 000972 (2014–2015) and No. 003265 (2016–2017), and research permits No. 00174-B (2014–2015) and No. 00317-B (2016–2017).</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec8">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.</p>
        <p>No AI tools were used in the preparation of this manuscript.</p>
      </sec>
      <sec sec-type="Funding" id="sec9">
        <title>Funding</title>
        <p>This study was supported by the Georgia Tech School of Biology and Elizabeth Smithgall Watts Endowment Fund, Fresno Chaffee Zoo's Wildlife Conservation Fund, Zoologische Gesellschaft für Arten- und Populationsschutz (ZGAP), Greenville Zoo Conservation Fund, Taronga Field Conservation Grants, Satch Krantz Conservation Fund, PPG Conservation and Sustainability Fund, Sophie Danforth Conservation Fund, Disney Conservation Fund, Mohamed bin Zayed Species Conservation Fund, Oklahoma City Zoo and Botanical Garden, Phoenix Zoo, and ERM Foundation.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec10">
        <title>Author contributions</title>
        <p>Conceptualization: MT-A, KTSG, BAL. Data curation: MT-A. Funding acquisition: MT-A, BAL. Investigation: MT-A, KTSG, BAL. Methodology: MT-A, KTSG, BAL. Resources: MT-A, BAL. Project administration: MT-A. Validation: MT-A, KTSG, BAL, AHE-G, JFG-M. Writing – original draft: MT-A, AHE-G, JFG-M. Writing – review and editing: MT-A, BAL, AHE-G, JFG-M.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec11">
        <title>Author ORCIDs</title>
        <p>M. Torres-Almazán <ext-link xlink:href="https://orcid.org/0009-0009-1995-1467" ext-link-type="uri">https://orcid.org/0009-0009-1995-1467</ext-link></p>
        <p>K.T. Schrei Godoy <ext-link xlink:href="https://orcid.org/0009-0003-1373-2139" ext-link-type="uri">https://orcid.org/0009-0003-1373-2139</ext-link></p>
        <p>B.A. Lock <ext-link xlink:href="https://orcid.org/0009-0000-4402-6251" ext-link-type="uri">https://orcid.org/0009-0000-4402-6251</ext-link></p>
        <p>A.H. Escobedo-Galván <ext-link xlink:href="https://orcid.org/0000-0003-1514-1255" ext-link-type="uri">https://orcid.org/0000-0003-1514-1255</ext-link></p>
        <p>J.F. González-Maya <ext-link xlink:href="https://orcid.org/0000-0002-8942-5157" ext-link-type="uri">https://orcid.org/0000-0002-8942-5157</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec12">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text.</p>
      </sec>
    </sec>
  </back>
</article>
