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   <front>
      <journal-meta>
         <journal-id journal-id-type="publisher-id">revmar</journal-id>
         <journal-title-group>
            <journal-title>Revista Ciencias Marinas y Costeras</journal-title>
            <abbrev-journal-title abbrev-type="publisher">Rev. Cienc. Mar. Cost</abbrev-journal-title>
         </journal-title-group>
         <issn pub-type="ppub">1659-455X</issn>
         <issn pub-type="epub">1659-407X</issn>
         <publisher>
            <publisher-name>Universidad Nacional, Costa Rica</publisher-name>
         </publisher>
      </journal-meta>
      <article-meta>
         <article-id pub-id-type="doi">10.15359/revmar.14-2.2</article-id>
         <article-categories>
            <subj-group subj-group-type="heading">
               <subject>Artículo</subject>
            </subj-group>
         </article-categories>
         <title-group>
            <article-title>First record of the Fiddler Crab, Minuca osa from the Eastern Montijo Gulf, Panama</article-title>
            <trans-title-group xml:lang="es">
               <trans-title>Primer reporte del cangrejo violinista, Minuca osa en el oriente del golfo de Montijo, Panamá</trans-title>
            </trans-title-group>
         </title-group>
         <contrib-group>
            <contrib contrib-type="author">
               <contrib-id contrib-id-type="orcid">0000-0002-0279-8621</contrib-id>
               <name>
                  <surname>Lombardo-González</surname>
                  <given-names>Roberto C.</given-names>
               </name>
               <xref ref-type="aff" rid="aff1">
                  <sup>1 </sup>
               </xref>
            </contrib>
            <aff id="aff1">
               <label>1</label>
               <institution content-type="original"> Universidad de Panamá, Centro Regional Universitario de Veraguas. Centro de Capacitación, Investigación y Monitoreo de la Biodiversidad en el Parque Nacional Coiba. roberto.lombardo@up.ac.pa* ORCID: https://orcid.org/0000-0002-0279-8621</institution>
               <institution content-type="orgname">Universidad de Panamá</institution>
               <institution content-type="orgdiv1">Centro Regional Universitario de Veraguas</institution>
               <institution content-type="orgdiv2">Centro de Capacitación, Investigación y Monitoreo de la Biodiversidad en el Parque Nacional Coiba</institution>
               <email>roberto.lombardo@up.ac.pa</email>
            </aff>
         </contrib-group>
         <pub-date pub-type="collection">
            <season>Jul-Dec</season>
            <year>2022</year>
         </pub-date>
         <volume>14</volume>
         <issue>2</issue>
         <fpage>27</fpage>
         <lpage>36</lpage>
         <history>Recepción:<date date-type="received">
               <day>30</day>
               <month>08</month>
               <year>2022</year>
            </date>Corregido:<date date-type="rev-recd">
               <day>05</day>
               <month>10</month>
               <year>2022</year>
            </date>Aceptado:<date date-type="accepted">
               <day>10</day>
               <month>10</month>
               <year>2022</year>
            </date>
         </history>
         <permissions>
            <license xml:lang="en" license-type="open-access"
                     xlink:href="https://creativecommons.org/licenses/by-nc-nd/3.0/">
               <license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution License</license-p>
            </license>
         </permissions>
         <abstract>
            <title>ABSTRACT</title>
            <p>Minuca osa had been previously thought to be endemic to Pacific Costa Rica (Golfo Dulce). However, specimens fitting the description were sampled in April 2021 and August 2022 in two sites in the Eastern Montijo Gulf, Panama. Based on the morphological character correspondence between the specimens collected and the original species description, it can be concluded that Minuca osa is also present in the Eastern Montijo Gulf.</p>
         </abstract>
         <trans-abstract xml:lang="es">
            <title>RESUMEN</title>
            <p>Anteriormente, se pensaba que Minuca osa era endémica del Pacífico de Costa Rica (golfo Dulce); sin embargo, en abril 2021 y agosto 2022, se colectaron especímenes que se ajustaban a la descripción de la especie en dos sitios del oriente del golfo de Montijo, Panamá. Con base en la correspondencia de caracteres morfológicos, se concluye que Minuca osa también está presente en el oriente del golfo de Montijo.</p>
         </trans-abstract>
         <kwd-group xml:lang="en">
            <title>Keywords:</title>
            <kwd>tuberculate ridge</kwd>
            <kwd>Uca</kwd>
            <kwd>dactyl</kwd>
            <kwd>chela</kwd>
            <kwd>pollex</kwd>
         </kwd-group>
         <kwd-group xml:lang="es">
            <title>Palabras clave:</title>
            <kwd>borde tuberculado</kwd>
            <kwd>Uca</kwd>
            <kwd>dáctilo</kwd>
            <kwd>quela</kwd>
            <kwd>pollex</kwd>
         </kwd-group>
         <counts>
            <fig-count count="2"/>
            <table-count count="2"/>
            <equation-count count="0"/>
            <ref-count count="16"/>
            <page-count count="10"/>
         </counts>
      </article-meta>
   </front>
   <body>
      <sec sec-type="intro">
         <title>INTRODUCTION</title>
         <p>Fiddler crabs (Ocypodidae: Uca Leach, 1814) are distributed along most tropical and semitropical coastlines of the world (<xref ref-type="bibr" rid="B15">Rosenberg, 2020</xref>).</p>
         <p>Research on fiddler crabs in Panama has been published on various topics, such as, functional morphology in <italic>Petruca panamensis</italic>, <italic>Leptuca terpsichores </italic>and <italic>L. beebei </italic>(<xref ref-type="bibr" rid="B11">Lim &amp; Goh, 2021</xref>), vision in <italic>L. stenodactylus </italic>(see <xref ref-type="bibr" rid="B5">How <italic>et al</italic>. 2015</xref>), sexual selection (<xref ref-type="bibr" rid="B4">Dennenmoser &amp; Christy, 2013</xref>), construction of structures (<xref ref-type="bibr" rid="B13">Pardo <italic>et al</italic>. 2020</xref>) in <italic>L. terpsichores </italic>and <italic>L. beebei</italic>, and larval ecology in <italic>L. terpsichores </italic>
            <xref ref-type="bibr" rid="B2">(Christy, 2003</xref>), and <italic>L. deichmanni </italic>(<xref ref-type="bibr" rid="B8">Kerr, 2015</xref>). Even though fiddler crabs are widely distributed in Panama, sampling in all the aforementioned studies was conducted exclusively at the Pacific entrance of the Panama Canal, whereas studies from other provinces are lacking.</p>
         <p>There are currently 35 recognized fiddler crab species in Panama. On the Atlantic coast, there are seven species, distributed into three genera: Minuca with four species, Uca with two, and a single species of Leptuca (<italic>L. thayeri</italic>). In the Pacific, there are 28 species grouped into four genera, of which sixteen are in the genus Leptuca, six within Uca, five within Minuca, and a single species of Petruca (<italic>P. panamensis</italic>) (<xref ref-type="bibr" rid="B14">Rosenberg, 2014</xref>).</p>
         <p>In the Ponuga River, Eastern Montijo Gulf, Veraguas Province, fiddler crabs were observed waving their claw in synchrony. The major claw, bright red-orange with a white tip, was the most striking feature of these fiddler crabs. A species fitting this morphological characteristic was described by <xref ref-type="bibr" rid="B10">Landstorfer &amp; Schubart (2010</xref>) from the Golfo Dulce, Pacific coast of Costa Rica <xref ref-type="fig" rid="f1">(<bold>Fig. 1A</bold>
            </xref>). The authors named the new species, <italic>Minuca osa</italic>, and suggested that it might be endemic. Based on geographical proximity of Golfo Dulce and our study area, it seemed plausible that <italic>M. osa </italic>could also be found in Panama <xref ref-type="fig" rid="f1">(<bold>Fig. 1B, C</bold>
            </xref>). Therefore, the objective of this study was to identify the specimens found in the Eastern Montijo Gulf and report on the possible geographical extension of <italic>M. osa </italic>distribution range.</p>
      </sec>
      <sec sec-type="materials|methods">
         <title>MATERIALS AND METHODS</title>
         <bold> </bold>
         <p>Sampling was conducted on April 27th, 2021, and August 13th and 14th, 2022 at two sites on the Eastern Montijo Gulf (<xref ref-type="fig" rid="f1">Fig. 1</xref>D); the Ponuga River (07° 51ʼ 51.3756ʼʼ N,-081° 00ʼ 52.6248ʼʼ W) and Mata Oscura (07° 27ʼ 29.0124ʼʼ N, -080° 55ʼ 12.8532ʼʼ W). Fiddler crabs (n = 22) were caught by hand, rinsed, and later stored in a freezer unit. Morphological characters of interest were photographed; specimens’ carapace width and length, as well as chela height and length, were measured (mean ± SD; 0.01 mm), and taxonomical keys (<xref ref-type="bibr" rid="B10">Landstorfer &amp; Schubart, 2010</xref>; Cra- ne, 2015; <xref ref-type="bibr" rid="B16">Shih <italic>et al</italic>. 2016</xref>; <xref ref-type="bibr" rid="B15">Rosenberg, 2020</xref>) were used for the identification of the collected material. Morphological characters between Golfo Dulce <xref ref-type="bibr" rid="B10">(Landstorfer &amp; Schubart, 2010</xref>) and the Eastern Montijo Gulf were compared with the t-test.</p>
         <p>
            <fig id="f1">
               <label>Fig. 1</label>
               <caption>
                  <title>Contexto geográfico y ubicaciones de reportes de <italic>Minuca osa </italic>en el Pacífico Tropical Oriental. A: sitio de procedencia (rojo) de los especímenes en la descripción original de <italic>Minuca osa </italic>(<xref ref-type="bibr" rid="B10">Landstorfer &amp; Schubart, 2010</xref>). Imágenes preliminares de <italic>Minuca osa </italic>macho (B) y hembra (C) del oriente del golfo de Montijo. D: Sitios de muestreo georeferenciados de Ponuga (norte) y Mata Oscura (sur) (amarillo); escala: 1:250,000. Proyección UTM (Universal Mercator Traverse 17 N). Datum horizontal: Sistema Geodésico Mundial 1984 (WGS-84). Datum vertical: Nivel medio del mar del geoide WGS-84</title>
               </caption>
               <graphic xlink:href="a02v14n2image001.jpg"/>
               <attrib>First record of the Fiddler Crab, <italic>Minuca osa </italic>from the Eastern Montijo Gulf, Panama</attrib>
            </fig>
         </p>
      </sec>
      <sec sec-type="results">
         <title>RESULTS</title>
         <p>Features of the genus <italic>Minuca </italic>were found in 22 individuals averaging 21.44 ± 2.96 mm and 14.60 ± 1.62 mm in carapace width and length, respectively; 14 were from Ponuga and eight from Mata Oscura. Specimens were larger than those from Golfo Dulce, except in major chela height, where no difference was detected (<xref ref-type="table" rid="t1">Table 1</xref>). In general, compared to other Minuca species, sampled individuals from Ponuga and Mata Oscura can be considered large within the genus <xref ref-type="table" rid="t2">(<bold>Table 2</bold>
            </xref>).</p>
         <p>
            <table-wrap id="t1">
               <label>Table 1</label>
               <caption>
                  <title>Morphological character comparisons (mm) between <italic>Minuca osa</italic> specimens collected in the Eastern Montijo Gulf (Panama) and those noted in the original species description by <xref ref-type="bibr" rid="B10">Landstorfer and Schubart (2010</xref>) at Golfo Dulce (Costa Rica). EMG: Eastern Montijo Gulf. GD: Golfo Dulce. CW: carapace width. CL: carapace length. QH: major chela height. QL: major chela length </title>
               </caption>
               <table>
                  <colgroup>
                     <col/>
                     <col/>
                     <col/>
                     <col/>
                     <col/>
                     <col/>
                     <col/>
                  </colgroup>
                  <tbody>
                     <tr>
                        <td align="left">EMG-CW</td>
                        <td align="center">22</td>
                        <td align="center">21.44 ± 2.96</td>
                        <td align="center">12.66</td>
                        <td align="center">25.98</td>
                        <td align="center">4.21; 18</td>
                        <td align="center">0.001</td>
                     </tr>
 
                     <tr>
                        <td align="left">GD-CW</td>
                        <td align="center">12</td>
                        <td align="center">16.22 ± 3.70</td>
                        <td align="center">9.28</td>
                        <td align="center">21.10</td>
                        <td align="center"> </td>
                        <td align="center"> </td>
                     </tr>
 
                     <tr>
                        <td align="left">EMG-CL</td>
                        <td align="center">22</td>
                        <td align="center">14.60 ± 1.62</td>
                        <td align="center">11.10</td>
                        <td align="center">17.89</td>
                        <td align="center">4.01; 15</td>
                        <td align="center">0.001</td>
                     </tr>
 
                     <tr>
                        <td align="left">GD-CL</td>
                        <td align="center">12</td>
                        <td align="center">11.32 ± 2.57</td>
                        <td align="center">6.12</td>
                        <td align="center">14.70</td>
                        <td align="center"> </td>
                        <td align="center"> </td>
                     </tr>
 
                     <tr>
                        <td align="left">EMG-QH</td>
                        <td align="center">22</td>
                        <td align="center">11.04 ± 2.34</td>
                        <td align="center">4.09</td>
                        <td align="center">14.66</td>
                        <td align="center">1.55; 13</td>
                        <td align="center">0.146</td>
                     </tr>
 
                     <tr>
                        <td align="left">GD-QH</td>
                        <td align="center">7</td>
                        <td align="center">9.76 ± 1.76</td>
                        <td align="center">7.83</td>
                        <td align="center">11.90</td>
                        <td align="center"> </td>
                        <td align="center"> </td>
                     </tr>
 
                     <tr>
                        <td align="left">EMG-QL</td>
                        <td align="center">22</td>
                        <td align="center">30.27 ± 7.80</td>
                        <td align="center">7.13</td>
                        <td align="center">42.23</td>
                        <td align="center">2.31; 18</td>
                        <td align="center">0.033</td>
                     </tr>
 
                     <tr>
                        <td align="left">GD-QL</td>
                        <td align="center">8</td>
                        <td align="center">24.41 ± 5.42</td>
                        <td align="center">18.32</td>
                        <td align="center">31.05</td>
                        <td align="center"> </td>
                        <td align="center"> </td>
                     </tr>
 
                     <tr>
                        <td align="left">EMG-Front</td>
                        <td align="center">22</td>
                        <td align="center">7.47 ± 1.04</td>
                        <td align="center">4.42</td>
                        <td align="center">9.09</td>
                        <td align="center">3.85; 13</td>
                        <td align="center">0.002</td>
                     </tr>
 
                     <tr>
                        <td align="left">GD-Front</td>
                        <td align="center">10</td>
                        <td align="center">5.59 ± 1.37</td>
                        <td align="center">3.11</td>
                        <td align="center">7.17</td>
                        <td align="center"> </td>
                        <td align="center"> </td>
                     </tr>
                  </tbody>
               </table>
            </table-wrap>
         </p>
         <p>
            <table-wrap id="t2">
               <label>Table 2</label>
               <caption>
                  <title>Carapace width differences between <italic>Minuca osa </italic>from the Eastern Montijo Gulf (mean = 21.44 mm), Golfo Dulce (Pacific Costa Rica) and other Minuca species reported in Panama</title>
               </caption>
               <graphic xlink:href="a02v14n2image002.jpg"/>
            </table-wrap>
         </p>
         <p>Morphological characters of the genus found in sampled specimens include: H-like depression in the cardiac-mesogastric region, carapace width to front ratio of approximately 35% (wide front; <xref ref-type="fig" rid="f2">Fig. 2A</xref>), with long anterolateral margins curving inwards <xref ref-type="fig" rid="f2">(<bold>Fig. 2A, D</bold>
            </xref>), small suborbital crenulations increasing towards outer angle <xref ref-type="fig" rid="f2">(<bold>Fig. 2B</bold>
            </xref>), narrow gape minor chelipeds with small serrations <xref ref-type="fig" rid="f2">(<bold>Fig. 2C</bold>
            </xref>), and two postero-lateral striae in the carapace <xref ref-type="fig" rid="f2">(<bold>Fig. 2D</bold>
            </xref>).</p>
         <p>A total of 16 males and six females exhibited traits that fit the description of <italic>Minuca osa</italic>. Specifically, segments of the pleon were not fused, carapace surface with small pits; no pubescence or setae on carapace <xref ref-type="fig" rid="f2">(<bold>Fig. 2A, D</bold>
            </xref>). There was pile with few long setae in the ambulatories dorsal margin of the carpus and propodus, in agreement with that reported for <italic>M. osa </italic>
            <xref ref-type="fig" rid="f2">(<bold>Fig. 2E</bold>
            </xref>). The dorsal margins of ambulatory meri in males had a soft dorsal tuberculate ridge <xref ref-type="fig" rid="f2">(<bold>Fig. 2F</bold>
            </xref>); dactyls of ambulatory legs have three distinct rows of short setae <xref ref-type="fig" rid="f2">(<bold>Fig. 2G</bold>
            </xref>). In males, the external surface and dorsal edge of the manus was tuberculate, with a minuscule hollow space, proximal to the predactylar groove. Tubers in the major chela diminished on lower manus and were minuscule on pollex and dactyl <xref ref-type="fig" rid="f2">(<bold>Fig. 2H</bold>
            </xref>). The pollex was curved upwards at the tip; both, the pollex and dactyl were flattened. The dactyl was longer than the pollex and was curved at approx. 90° downward at tip <xref ref-type="fig" rid="f2">(<bold>Fig. 2H, K</bold>, <bold>M</bold>
            </xref>). The pollex showed three rows of tubercles, and one large carina beyond the midpoint <xref ref-type="fig" rid="f2">(<bold>Fig. 2I</bold>
            </xref>). There was one large subdistal tooth at the gape midrow of the pollex, forming three distinct tubercles (white) on the tip <xref ref-type="fig" rid="f2">(<bold>Fig. 2J, K, M</bold>
            </xref>). The dactyl had three to four larger tubercles (<bold>Fig. 2K, M</bold>). The inner surface of the manus had a ridge of dense tubercles lining the margin of the carpal cavity, warping in direction to the palm with tubercles growing in size and ending in a patch in the central section of the manus, almost connecting to the oblique tuberculate edge <xref ref-type="fig" rid="f2">(<bold>Fig. 2L</bold>
            </xref>). The ventral manus tuberculate ridge was oblique with a high apex, sloping into pollex ventral surface. This ridge had distinctive tubercles decreasing in size towards the pollex with increased irregularity. The pollex and manus articulation had a deep sulcus which ran between the pollex ventral margin and two predactylar tuberculate ridges; the nearest was divergent and curled into the inner dorsal tuberculate ridge of the pollex <xref ref-type="fig" rid="f2">(<bold>Fig. 2L, M</bold>
            </xref>). The carpal surface (dorsal) of major chelipeds bearded four to five sharp tubers, and the merus showed numerous sharp tubercles on the dorsal edge <xref ref-type="fig" rid="f2">(<bold>Fig. 2N</bold>
            </xref>). The gonopod had two flanges and a thumb; the posterior flange was longer than the anterior. The genital pore and flanges were covered by extension of the shaft. Setae were on the opposite side of the flanges, on thumb and tip of gonopod <xref ref-type="fig" rid="f2">(<bold>Fig. 2O</bold>
            </xref>).</p>
         <p>
            <fig id="f2">
               <label>Fig. 2</label>
               <caption>
                  <title>
                     <italic>Minuca osa </italic>diagnostic features from the Eastern Montijo Gulf, Veraguas, Panama. A: carapace. B: orbital cavity. C: minor chelipeds, left female (outer side); right male (inner surface). D: carapace striae. E: pubescence in carpus and propodus of ambulatories. F: ambulatory merus tuberculate ridge. G: ambulatories dactylus setae rows. H: major chela. I: pollex tubercle rows and large carina beyond midpoint. J: large tubercles at pollex tip. K: dactyl edge tubercles. L-M: manus, palm, and pollex tubercular ridges. N: large chela carpus and merus tubercles. O: right gonopod, anterolateral (left) and posteromesial (right). (Scale bar: A, D, I-L, N = 5 mm; B, C, F, G = 2 mm; H, M = 10 mm; E = 0.5 mm; O = 0.2 mm)</title>
               </caption>
               <graphic xlink:href="a02v14n2image003.jpg"/>
            </fig>
         </p>
      </sec>
      <sec sec-type="discussion">
         <title>DISCUSSION</title>
         <bold> </bold>
         <p>
            <italic>Minuca burgersi</italic>, <italic>M. vocator, </italic>and <italic>M. ecuadoriensis </italic>are morphologically similar to <italic>M. osa </italic>(<xref ref-type="bibr" rid="B10">Landstorfer &amp; Schubart, 2010</xref>; <xref ref-type="bibr" rid="B15">Rosenberg, 2020</xref>); however, <italic>M. burgersi </italic>and <italic>M. vocator </italic>are only distributed in the Western Atlantic, while <italic>M. ecuadoriensis </italic>is only found in the Eastern Pacific (<xref ref-type="bibr" rid="B12">Masunari <italic>et al</italic>. 2020</xref>; <xref ref-type="bibr" rid="B15">Rosenberg, 2020</xref>). Furthermore, <italic>M. burgersi </italic>also differs from our specimens in that the oblique tuberculate ridge on the major palm has small tubercles (not a single row); tubercle ridge does not diminish distally on outer pollex <xref ref-type="bibr" rid="B3">(Crane, 2015</xref>; <xref ref-type="bibr" rid="B16">Shih <italic>et al</italic>. 2016</xref>). In <italic>Minuca vocator </italic>and <italic>M. ecuadoriensis</italic>, the carapace as well as the ambulatories on all segments have a profuse pile (except dactyls) in an irregular pattern (see <xref ref-type="bibr" rid="B12">Masunari <italic>et al</italic>. 2020</xref>), while our specimens had no such pile. In <italic>M. ecuadoriensis</italic>, the oblique ridge in the major palm has a very low apex; tubercles are small and irregular, sometimes vestigial or absent. In contrast, the apex in <italic>M. osa </italic>is high and tubercles are large (<xref ref-type="bibr" rid="B10">Landstorfer &amp; Schubart, 2010</xref>; <xref ref-type="bibr" rid="B3">Crane, 2015</xref>); all of our specimens showed the latter. Considering its behavior and coloration, <italic>M. osa </italic>is a conspicuous fiddler crab species; thus, it is unclear why it was not been identified other than in Golfo Dulce <xref ref-type="bibr" rid="B10">(Landstorfer &amp; Schubart, 2010</xref>) prior to this report. Based on the morphological differences between <italic>M. osa, M. burgersi, M. vocator, </italic>and <italic>M. ecuadoriensis</italic>, it can be concluded that <italic>M. osa </italic>is also present in the Eastern Montijo Gulf, Panama</p>
      </sec>
   </body>
   <back>
      <ack>
         <title>ACKNOWLEDGEMENTS</title>
         <p>Our gratitude goes to the anonymous reviewers for their comments to improve the manuscript and to John Christy and Fumio Takeshita for their preliminary discussions. We would also like to thank Jaime Rivera for his cartographic assistance, as well as Jacinto Rodríguez and the Cantarana Hotel staff for their hospitality</p>
      </ack>
      <ref-list>
         <title>REFERENCES</title>
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                     <surname>Negreiros-Fransozo</surname>
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               </person-group>
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            </element-citation>
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