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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.13-2.2</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>Artículo</subject>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>Predation of <italic>Gecarcinus quadratus</italic> (Decapoda: Gecarcinidae) by <italic>Aramides cajaneus</italic> (Gruiformes: Rallidae) in Costa Rica</article-title>
				<trans-title-group xml:lang="es">
					<trans-title>Depredación de <italic>Gecarcinus quadratus</italic> (Decapoda: Gecarcinidae) por <italic>Aramides cajaneus</italic> (Gruiformes: Rallidae) en Costa Rica</trans-title>
				</trans-title-group>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author">
					<name>
						<surname>Villegas-Retana</surname>
						<given-names>Sergio</given-names>
					</name>
					<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
				</contrib>
				<contrib contrib-type="author">
					<name>
						<surname>Picado-Masís</surname>
						<given-names>Jorge</given-names>
					</name>
					<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
				</contrib>
			</contrib-group>
			<aff id="aff1">
				<label>1</label>
				<institution content-type="original"> Escuela de Biología, Universidad de Costa Rica, San Pedro de Montes de Oca, 11501-2060 San José, Costa Rica. sergio.villegas@ucr.ac.cr ORCID: http://orcid.org/0000-0002-9579-1479</institution>
				<institution content-type="normalized">Universidad de Costa Rica</institution>
				<institution content-type="orgdiv1">Escuela de Biología</institution>
				<institution content-type="orgname">Universidad de Costa Rica</institution>
				<addr-line>
					<city>San Pedro de Montes de Oca</city>
				</addr-line>
				<country country="CR">Costa Rica</country>
				<email>sergio.villegas@ucr.ac.cr</email>
			</aff>
			<aff id="aff2">
				<label>2</label>
				<institution content-type="original"> Professional certified tour guide, Manuel Antonio National Park, Quepos, 60601 Puntarenas, Costa Rica. picadomasisjorge@gmail.com, ORCID: https://orcid.org/0000-0003-4493-9346</institution>
				<addr-line>
					<city>Quepos</city>
				</addr-line>
				<country country="CR">Costa Rica</country>
				<email>picadomasisjorge@gmail.com</email>
			</aff>
			<pub-date pub-type="collection">
				<season>Jul-Dec</season>
				<year>2021</year>
			</pub-date>
			<volume>13</volume>
			<issue>2</issue>
			<fpage>23</fpage>
			<lpage>28</lpage>
			<history>
				<date date-type="received">
					<day>12</day>
					<month>07</month>
					<year>2021</year>
				</date>
				<date date-type="rev-recd">
					<day>17</day>
					<month>08</month>
					<year>2021</year>
				</date>
				<date date-type="accepted">
					<day>19</day>
					<month>08</month>
					<year>2021</year>
				</date>
			</history>
			<permissions>
				<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by-nc-nd/3.0/" xml:lang="en">
					<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><italic>Gecarcinus quadratus</italic> is a land crab preyed upon by some species, but there is little information about the genera and species of these predators. This scientific note presents the first record of <italic>Aramides cajaneus</italic> preying upon <italic>G. quadratus</italic>. This observation occurred during daylight in May 2021 in Costa Rica, and contributes to the knowledge about the natural history, behavior and ecology of these species.</p>
			</abstract>
			<trans-abstract xml:lang="es">
				<title>RESUMEN</title>
				<p><italic>Gecarcinus quadratus</italic> es un cangrejo terrestre depredado por algunas especies, sin embargo, hay poca información en el nivel de género y especie de sus predadores. Esta nota científica presenta el primer registro de <italic>Aramides cajaneus</italic> depredando a <italic>G. quadratus</italic>. La observación ocurrió durante el día, en mayo del 2021, en Costa Rica y contribuye al conocimiento sobre la historia natural, comportamiento y la ecología de estas especies.</p>
			</trans-abstract>
			<kwd-group xml:lang="en">
				<title>Keywords:</title>
				<kwd>behavior</kwd>
				<kwd>diet</kwd>
				<kwd>Gray-cowled Wood-Rail</kwd>
				<kwd>Halloween Crab</kwd>
				<kwd>natural history</kwd>
			</kwd-group>
			<kwd-group xml:lang="es">
				<title>Palabras clave:</title>
				<kwd>comportamiento</kwd>
				<kwd>dieta</kwd>
				<kwd>Cangrejo de Halloween</kwd>
				<kwd>Rascón Cuelligrís</kwd>
				<kwd>historia natural</kwd>
			</kwd-group>
			<counts>
				<fig-count count="1"/>
				<table-count count="0"/>
				<equation-count count="0"/>
				<ref-count count="17"/>
				<page-count count="6"/>
			</counts>
		</article-meta>
	</front>
	<body>
		<sec sec-type="intro">
			<title>INTRODUCTION</title>
			<p><italic>Gecarcinus quadratus</italic> (de Saussure, 1853) is a terrestrial crab that inhabits neotropical coastal forests, and is distributed from Florida, through the Antilles and Central America to Guyana on the Atlantic Coast, and from Mexico to Peru on the Pacific Coast (<xref ref-type="bibr" rid="B1">Burggren &amp; McMahon, 1988</xref>; <xref ref-type="bibr" rid="B8">Sherman, 2002</xref>). It is fossorial, its populations (of up to six crabs m<sup>-2</sup>) extend approximately 600 m inland from the shore in coastal forests, and its activity outside the burrow increases during the rainy season (<xref ref-type="bibr" rid="B16">Von Prahl &amp; Manjarrés, 1984</xref>;<xref ref-type="bibr" rid="B8"> Sherman, 2002</xref>; <xref ref-type="bibr" rid="B9">Sherman, 2003</xref>). It has been reported as being preyed upon by <italic>Crocodylus acutus</italic> (Cuvier, 1807), <italic>Procyon lotor</italic> (Linnaeus, 1758), some birds and perhaps other crabs (<xref ref-type="bibr" rid="B2">Carrillo et al. 2001</xref>; <xref ref-type="bibr" rid="B6">Perger et al. 2013</xref>; <xref ref-type="bibr" rid="B3">Cupul-Magaña et al. 2015</xref>; <xref ref-type="bibr" rid="B15">Villagrán-Colón et al. 2016</xref>). <italic>G. quadratus</italic> may have developed adaptations to high levels of predation (Perger, 2013), but there is little information about the genera and species of these predators.</p>
			<p>On the other hand, <italic>Aramides cajaneus</italic> (Statius Muller, 1776) is a bird distributed from southwestern Costa Rica southwards to northern Argentina, from lowlands up to 2,300 m.a.s.l. (<xref ref-type="bibr" rid="B13">Stiles &amp; Skutch, 1989</xref>; <xref ref-type="bibr" rid="B14">Taylor, 2020</xref>). It inhabits wet ground along the sides of streams, as well as swamps, secondary growth and deciduous woodlands, mangroves, marshes and marshy thickets, rice and cane fields, seasonal pools and scrubby pastures, as long as they include wet and shaded areas (<xref ref-type="bibr" rid="B13">Stiles &amp; Skutch, 1989</xref>; <xref ref-type="bibr" rid="B14">Taylor, 2020</xref>). Its diet is diverse and includes mostly invertebrates (including crabs), but also small vertebrates such as frogs, as well as seeds and fruits (<xref ref-type="bibr" rid="B13">Stiles &amp; Skutch, 1989</xref>; <xref ref-type="bibr" rid="B14">Taylor, 2020</xref>). It also feeds on the feces of giant otters in latrines (<xref ref-type="bibr" rid="B5">Leuchtenberger et al. 2012</xref>).</p>
			<p>Among the crab species reported as prey of <italic>A. cajaneus</italic> are: <italic>Uca rapax</italic> (Smith, 1870), <italic>U. mordax</italic> (Smith, 1870)<italic>, U. leptodactyla</italic> (Rathbun in Rankin, 1898)<italic>, U. uruguayensis</italic> (Nobili, 1901), and <italic>Callinectes danae</italic> (Smith, 1869; <xref ref-type="bibr" rid="B10">Silva &amp; Olmos 2015</xref>). It has also been observed consuming droppings of horses and <italic>P. lotor</italic>; in the latter, perhaps looking for remains of crab exoskeletons, possibly of the genera <italic>Gecarcinus</italic> and <italic>Cardiosma</italic> (<xref ref-type="bibr" rid="B11">Skutch, 1994</xref>; <xref ref-type="bibr" rid="B12">Solano-Ugalde, 2005</xref>). Nevertheless, there are no reports of <italic>A. cajaneus</italic> preying on <italic>G. quadratus</italic>. The following report represents the first record of an event of this kind.</p>
		</sec>
		<sec sec-type="materials|methods">
			<title>MATERIALS AND METHODS</title>
			<p>The event occurred on the El Perezoso Trail, at Manuel Antonio National Park, Quepos, Puntarenas, Costa Rica (9° 23' N, 84° 8' W; 23 m.a.s.l.). On 24 May 2021, at 08 00 h, the second author observed an individual of <italic>A. cajaneus</italic> preying on an individual of <italic>G. quadratus</italic> during a guided tour. The crab was coming out of the forest and the wood-rail followed, pecking at it. When they reached the trail, the crab tried to cross it and escape, but the wood-rail took advantage to get close to it, catching one of its walking legs with its bill and flipping it on its back. It began to hammer at the crab with its bill to break its shell, while the crab tried to defend itself by raising its chelipeds (<xref ref-type="fig" rid="f1">Fig. 1</xref>). Finally, the bird killed the crab and carried it into the forest as a large group of people approached the site. The whole event lasted at least 2 minutes. </p>
			<p>
				<fig id="f1">
					<label>Fig. 1</label>
					<caption>
						<title><italic>Aramides cajaneus</italic> preying on <italic>Gecarcinus quadratus</italic>, Manuel Antonio National Park, Costa Rica, 2021 (Photos from a video recorded by Jorge Picado; the pictures should be viewed from left to right in each row and the rows should be viewed from top to bottom)</title>
					</caption>
					<graphic xlink:href="1659-407X-revmar-13-02-23-gf1.jpg"> </graphic>
				</fig>
			</p>
			<p>Fig. 1. <italic>Aramides cajaneus</italic> depredando a <italic>Gecarcinus quadratus</italic>, Parque Nacional Manuel Antonio, Costa Rica, 2021 (Secuencia de fotos del vídeo grabado por Jorge Picado, ver de izquierda a derecha y de arriba hacia abajo)</p>
		</sec>
		<sec sec-type="results|discussion">
			<title>RESULTS AND DISCUSSION</title>
			<p>Land crabs are preyed upon by a wide range of vertebrates (<xref ref-type="bibr" rid="B1">Burggren &amp; McMahon, 1988</xref>). Many of them are birds, which have been identified at the species level (<xref ref-type="bibr" rid="B1">Burggren &amp; McMahon, 1988</xref>; <xref ref-type="bibr" rid="B17">Wolcott &amp; Wolcott, 1982</xref>; <xref ref-type="bibr" rid="B10">Silva &amp; Olmos 2015</xref>). In the case <italic>G. quadratus</italic>, we only found two predators identified to this level (<xref ref-type="bibr" rid="B2">Carrillo et al. 2001</xref>; <xref ref-type="bibr" rid="B3">Cupul-Magaña et al. 2015</xref>). Birds have also been reported as predators of <italic>G. quadratus</italic>, but there is no specific data on their taxonomy; this is the first study in which an avian predator is identified at the species level.</p>
			<p>Some crab species are also an important part of the diet of <italic>A. cajaneus</italic> (<xref ref-type="bibr" rid="B10">Silva &amp; Olmos 2015</xref>; <xref ref-type="bibr" rid="B14">Taylor, 2020</xref>). In mangroves, for example, this bird preys heavily on crabs (<xref ref-type="bibr" rid="B14">Taylor, 2020</xref>). In this sense, it could be that this crab is a regular part of the diet of <italic>A. cajaneus</italic>, along with other crab species (<xref ref-type="bibr" rid="B10">Silva &amp; Olmos 2015</xref>), but it is necessary to carry out further studies to determine its diet in more detail. It is possible that the wood-rail took advantage of the crab crossing the trail to hunt it.</p>
			<p>The wood-rail hammered its prey with its bill to destroy its shell, a behavior which has been reported when this species preys on snails (<xref ref-type="bibr" rid="B14">Taylor, 2020</xref>). While the crab tried to defend itself by raising its chelipeds, it was unable to use them successfully or use autonomy in response to the attack, which is a defensive system used by <italic>G. quadratus</italic> to defend itself effectively while wandering far from a burrow or retreat (<xref ref-type="bibr" rid="B7">Robinson et al. 1970</xref>). Likewise, the Halloween Crab is nocturnal (<xref ref-type="bibr" rid="B8">Sherman, 2002</xref>; <xref ref-type="bibr" rid="B9">Sherman, 2003</xref>), and <italic>A. cajaneus</italic> apparently often feeds at night (<xref ref-type="bibr" rid="B14">Taylor, 2020</xref>), but this observation occurred at 08:00 h. However, both species can also engage in diurnal activity (<xref ref-type="bibr" rid="B16">Von Prahl &amp; Manjarrés, 1984</xref>; <xref ref-type="bibr" rid="B12">Solano-Ugalde, 2005</xref>), so it is possible that this was an opportunistic event. </p>
			<p>Knowledge about predation is one of the initial steps to understand the ecology and ethology of animals (<xref ref-type="bibr" rid="B4">Curio, 2012</xref>). Identifying species of predators provides basic information to study its impact on the life histories of prey and their defense mechanisms, as well as the hunting behavior and diet of predators (<xref ref-type="bibr" rid="B7">Robinson et al. 1970</xref>; Curio, 2012; <xref ref-type="bibr" rid="B6">Perger et al. 2013</xref>). The evidence presented here contributes to the knowledge about the natural history, behavior, and ecology of both <italic>G. quadratus</italic> and <italic>A. cajaneus</italic>.</p>
		</sec>
	</body>
	<back>
		<ack>
			<title>ACKNOWLEDGEMENTS</title>
			<p>We gratefully acknowledge to Ingo Wehrtmann for suggestions and recommendations for preparing the manuscript. We also want to thank the two anonymous reviewers for suggestions that improved the manuscript.</p>
		</ack>
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