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<article article-type="research-article" dtd-version="1.1" xml:lang="en" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance">

	<front>
		<journal-meta>
			<journal-id journal-id-type="publisher-id">Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria (INIA)</journal-id>
			<journal-title-group>
				<journal-title>Forest System</journal-title>
				<abbrev-journal-title>For Syst</abbrev-journal-title>
			</journal-title-group>
			<issn pub-type="epub">2171-9845</issn>
			<publisher>
				<publisher-name>Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria (INIA)</publisher-name>
			</publisher>
		</journal-meta>
		<article-meta>
			<article-id pub-id-type="publisher-id">20521</article-id>
			<article-id pub-id-type="doi">10.5424/fs/2023323-20521</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>SHORT COMMUNICATION</subject>
				</subj-group>
			</article-categories>

			<title-group>
				<article-title>A low altitude forest dwelling Pyrenean chamois population increases the potential habitat of this subspecies</article-title>
			</title-group>

			<contrib-group>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-8273-3141</contrib-id>
					<name>
						<surname>Herrero</surname>
						<given-names>Juan</given-names>
					</name>
					<aff id="aff1"><institution>Technical School, University of Zaragoza, </institution><addr-line>22071 Huesca, </addr-line><country>Spain.</country></aff>
				</contrib>

				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-2474-7729</contrib-id>
					<name>
						<surname>García-Serrano</surname>
						<given-names>Alicia</given-names>
					</name>
					<aff id="aff2"><institution>Ega Wildlife Consultants, </institution><addr-line>Sierra de Vicor 31, 50003 Zaragoza, </addr-line><country>Spain.</country></aff>
				</contrib>

				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0009-0002-8091-0516</contrib-id>
					<name>
						<surname>Félez</surname>
						<given-names>Carlos</given-names>
					</name>
					<aff id="aff3"><institution>Oficina Comarcal Agroambiental del Gobierno de Aragón, </institution><addr-line>Samper 2-4 bajos, 22340 Boltaña, </addr-line><country>Spain.</country></aff>
				</contrib>				

				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0009-0001-2850-9656</contrib-id>
					<name>
						<surname>Herrero</surname>
						<given-names>Fernando</given-names>
					</name>
					<aff id="aff3"><institution>Oficina Comarcal Agroambiental del Gobierno de Aragón, </institution><addr-line>Samper 2-4 bajos, 22340 Boltaña, </addr-line><country>Spain.</country></aff>
				</contrib>

				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0009-0007-3222-9431</contrib-id>
					<name>
						<surname>Machuca</surname>
						<given-names>Alberto</given-names>
					</name>
					<aff id="aff3"><institution>Oficina Comarcal Agroambiental del Gobierno de Aragón, </institution><addr-line>Samper 2-4 bajos, 22340 Boltaña, </addr-line><country>Spain.</country></aff>
				</contrib>			

				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0009-0007-3262-3455</contrib-id>
					<name>
						<surname>Ponz</surname>
						<given-names>Bosco</given-names>
					</name>
					<aff id="aff3"><institution>Oficina Comarcal Agroambiental del Gobierno de Aragón, </institution><addr-line>Samper 2-4 bajos, 22340 Boltaña, </addr-line><country>Spain.</country></aff>
				</contrib>

				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0009-0002-2936-5372</contrib-id>
					<name>
						<surname>Sancho</surname>
						<given-names>Santiago</given-names>
					</name>
					<aff id="aff3"><institution>Oficina Comarcal Agroambiental del Gobierno de Aragón, </institution><addr-line>Samper 2-4 bajos, 22340 Boltaña, </addr-line><country>Spain.</country></aff>
				</contrib>				

			</contrib-group>			


			<pub-date pub-type="epub">
				<day>24</day>
				<month>08</month>
				<year>2023</year>
			</pub-date>			
			<pub-date pub-type="collection">
				<month>11</month>
				<year>2023</year>
			</pub-date>
			<volume>32</volume>
			<issue>3</issue>
			<elocation-id>eSC03</elocation-id>
			<history>
				<date date-type="received">
					<day>06</day>
					<month>06</month>
					<year>2023</year>
				</date>
				<date date-type="accepted">
					<day>24</day>
					<month>08</month>
					<year>2023</year>
				</date>
				<date date-type="pub">
					<day>24</day>
					<month>08</month>
					<year>2023</year>
				</date>
			</history>			
			<permissions>
				<copyright-statement>&#xa9;2023 CSIC</copyright-statement>
				<copyright-year>2023</copyright-year>
				<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by/4.0/">
					<license-p>This is an open access article distributed under the terms of the Creative Commons Attribution 4.0 International (CC BY 4.0) License.</license-p>
				</license>
			</permissions>

			<self-uri xlink:href="https://doi.org/10.5424/fs/2023323-20521"/>
			<abstract>
				<title>Aim of study:</title>
				<p>To demonstrate if a population of Pyrenean chamois <em>Rupicapra p. pyrenaica</em> can live at low altitudes all year long. This could enlarge dramatically its potential habitat.</p>
				<title>Area of study:</title>
				<p>A hunting ground in Sobrarbe County, Pyrenees, Spain, in 2022.</p>
				<title>Material and methods:</title>
				<p>We interviewed local hunters to find out when the presence of the subspecies in the area dates back. We performed block counts from April to December in the target area for calculating the size of the population.</p>
				<title>Main results:</title>
				<p>There was a presence of at least 15 years of the subspecies living at 600 m asl. The population was at least 18 animals, reproduces, and shows a normal demographic structure of kids, yearlings, adult females, and adult males.</p>
				<title>Research highlights:</title>
				<p>Low-altitude chamois populations living in forests could be a result of important ecological changes in high mountain pastures producing migrations to newly suitable areas as low-altitude forests. The potential habitat of the subspecies should be broadened.</p>
			</abstract>

			<kwd-group>
				<kwd>global warming;</kwd>
				<kwd>expansion;</kwd>
				<kwd>potential habitat;</kwd>
				<kwd>wild ungulates;</kwd>
				<kwd><em>Rupicapra p. pyrenaica</em></kwd>
			</kwd-group>
			</article-meta>
			<funding-group id="fw-01">
				<award-group id="aw1">
					<funding-source>This research is part of the long-term monitoring of wild ungulates in Aragon, Spain, supported by its Regional Government</funding-source>
				</award-group>				
			</funding-group>		
	</front>
	
	<body>
		<sec id="sec1" sec-type="intro">
			<title>Introduction</title>

			<p>Wild ungulates have experienced an extraordinary increase in number and expansion in occupancy in Europe (Apollonio et al., 2010) and in the Pyrenees (Gortázar et al., 2000; González et al., 2013) in the last decades. Southern chamois <em>Rupicapra pyrenaica</em> is one of these species. This mountain ungulate is widespread, abundant, and increasing. It is adapted to high elevations and cold climates (Corlatti et al., 2022), occupying altitudes between 1,000 and 2,800 m asl. It is endemic to Southern Europe and has three subspecies: <em>Rupicapra pyrenaica parva</em>, living in the Cantabrian Mountains, <em>Rupicapra pyrenaica ornata</em>, in the Appenines and <em>Rupicapra p. pyrenaica</em> in the Pyrenees (Herrero et al., 2020). In 2002 the population of Pyrenean chamois was estimated at around 53,000 animals, comprising one single metapopulation (Herrero et al., 2004). Even if there are no complete figures of its historical distribution in the whole Pyrenees, in the mid-19th century it was marginal. Since the mid-20th century, the population experienced an extraordinary expansion and today occupies the whole mountain chain (Gortázar et al., 2000).</p>

			<p>This endemic Pyrenean subspecies is found in alpine meadows, rocky areas, forested valleys, and lower slopes in mountainous regions. Pyrenean chamois is a mixed grazer or intermediate feeder (García-González &amp; Cuartas, 1996) adapted to high mountain environments with pastures, scrubs, and forests. The subspecies has occupied its potential habitat in the mountain range and has been subjected to sporadic outbreaks which produced important mortality (Arnal et al., 2013; Corlatti et al., 2022). It generally stays above 1,800 m in alpine meadows during the warmer months of the year (Corlatti et al., 2022). Some populations have started to inhabit forests permanently (Herrero et al., 1996), even if their ecology is poorly known. Expansion continues nowadays on the border of the distribution of the subspecies, creating new subpopulations (Antón et al., 2019) and showing sporadic presence in marginal areas (Sampere et al., 2023).</p>

			<p>It is considered of Least Concern under IUCN criteria, listed in Appendix III of the Bern Convention and Annex V of the EU Habitat Directive (Herrero et al., 2020) and suitable to be hunted in the whole Pyrenees except for the Navarre region, Spain. Nowadays it is a relevant primary consumer, important huntable trophy and a touristic attraction. Reintroductions have been frequent in the French part of the Pyrenees (Corlatti et al., 2022).</p>

			<p>In February 2022 we received the testimony of two local hunters on the presence of Pyrenean chamois living for a long period (since 2007, 15 years) in a forest area of Southcentral Pyrenees (Fig. 1). They were asking the Regional Government for a chamois quota for their hunting ground. As there was no previous information on this issue, we decided to undertake a one-year monitoring system to increase the knowledge of this particular situation. In this short communication, we would like to describe the demographic characteristics of the subpopulation and comment on the possibilities that this new situation creates.</p>

			<fig id="f1">
				<label>Figure 1</label>
				<caption>
					<title> Localization of study area in Southern Pyrenees, Spain.</title>
				</caption>
				<graphic id="gra-1" xlink:href="img/eSC03-fig1.jpg"/>
			</fig>				
		
		</sec> <!--/sec 1-->

		<sec id="sec2" sec-type="materials|methods">
			<title>Material and methods</title>

			<p>We performed block counts (Berducou et al., 1986, Herrero et al., 2011) of the area where Pyrenean chamois was supposed to dwell once in April (before parturition), June (after parturition), September and November (rut) during 2022. Four operators walked in the area after sunrise following a planned route using binoculars, spotting scopes, mobile phones and cameras, for a period of around 4 h. When Pyrenean chamois individuals were sighted, animals were registered in a sheet. Double counts were avoided gathering together information at the end of each survey, considering sex, age, localization, and timing. Occasional sightings were also recorded.</p>
	
				<table-wrap id="t1">
					<label>Table 1</label>
					<caption>
						<title>Sightings of Pyrenean chamois in the study area during 2022</title>
					</caption>
					<table>
						<thead>
							<tr>
								<th align="center">Date</th>
								<th align="center">Source</th>
								<th align="center">Males</th>
								<th align="center">Females</th>
								<th align="center">Kids</th>
								<th align="center">Yearlings</th>
								<th align="center">Adults</th>
								<th align="center">Indet.</th>
								<th align="center">Total</th>
							</tr>
						</thead>
						<tbody>
						    <tr>
						      <td align="center">February</td>
						      <td align="center">Hunters</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">2</td>
						      <td align="center"></td>
						      <td align="center">15</td>
						      <td align="center"></td>
						      <td align="center">17</td>
						    </tr>
						    <tr>
						      <td align="center">March</td>
						      <td align="center">Casual</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">7</td>
						      <td align="center">7</td>
						    </tr>
						    <tr>
						      <td align="center">April</td>
						      <td align="center">Count</td>
						      <td align="center">5</td>
						      <td align="center">2</td>
						      <td align="center">2</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">9</td>
						    </tr>
						    <tr>
						      <td align="center">June</td>
						      <td align="center">Casual</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">4</td>
						    </tr>
						    <tr>
						      <td align="center">July</td>
						      <td align="center">Count</td>
						      <td align="center"></td>
						      <td align="center">4</td>
						      <td align="center">2</td>
						      <td align="center">5</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">11</td>
						    </tr>
						    <tr>
						      <td align="center">August</td>
						      <td align="center">Count</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">10-12</td>
						      <td align="center">10-12</td>
						    </tr>
						    <tr>
						      <td align="center">September</td>
						      <td align="center">Count</td>
						      <td align="center"></td>
						      <td align="center">3</td>
						      <td align="center"></td>
						      <td align="center">7*</td>
						      <td align="center"></td>
						      <td align="center">4</td>
						      <td align="center">14</td>
						    </tr>
						    <tr>
						      <td align="center">November</td>
						      <td align="center">Count</td>
						      <td align="center"></td>
						      <td align="center">5</td>
						      <td align="center">3</td>
						      <td align="center">1</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">9</td>
						    </tr>
						    <tr>
						      <td align="center">Estimate</td>
						      <td align="center"></td>
						      <td align="center">5</td>
						      <td align="center">5</td>
						      <td align="center">3</td>
						      <td align="center">5</td>
						      <td align="center"></td>
						      <td align="center"></td>
						      <td align="center">18</td>
						    </tr>
						</tbody>						
					</table>
					<table-wrap-foot>
						<fn id="TFN1">
							<p>*: some were probably kids.</p>
						</fn>
					</table-wrap-foot>					
				</table-wrap>			

			<sec id="sec2.1">
				<title>Study area</title>

				<p>The area has strong slopes of loam substrate. Forest tree cover is composed mainly by <em>Pinus halepensis</em>, but other species such <em>Quercus × cerrioides</em>, <em>Pinus nigra</em> or <em>Pinus sylvestris</em> are locally common. <em>Medicago sativa</em> pastures are present too. Total surface surveyed was around 1,000 ha. Other ungulates in the area were domestic sheep <em>Ovis aries</em>, wild boar <em>Sus scrofa</em>, and roe deer <em>Capreolus capreolus</em>. Wild boar is usually hunted in battues and roe deer in still hunts. No interference with chamois occurred during battues due to the proper behavior of hunting dogs. The entire study area is under Natura 2000 Network protection because of its Aleppo pine autochthonous forests, among other values (Code ES2410055).</p>

				<p>The source of this subpopulation of chamois could be the nearby Sierra Ferrera (2,295 m), with a population of around 150 chamois, stable since 2000 (García-Serrano &amp; Herrero, 2022).</p>

			</sec> <!--sec2.1-->	

		</sec> <!--/sec 2-->

		<sec id="sec3" sec-type="results">
			<title>Results and discussion</title>

			<p>Table 1 shows the results of the survey (Fig. 2). There were sighted at least 18 chamois during the whole study period. Movements from the neighboring Sierra Ferrera could not be discarded. The subpopulational stock is very low compared to other Pyrenean subpopulation (Herrero et al., 2004), but the population structure is comparable to other forest populations (Garin &amp; Herrero, 1997). Even habitat use and grouping are comparable to other forest dwelling populations of the species (Herrero et al., 1996, 2002). The occupied area is around 2 km2 and the density is 9 chamois km-2, moving in an altitudinal range of 600-650 m asl.</p>

			<fig id="f2">
				<label>Figure 2</label>
				<caption>
					<title>Above: adult female and kid in the loam. Below: group of chamois in the loam surrounded by Aleppo pine; from left to right: kid, yearling, adult female and adult female. </title>
				</caption>
				<graphic id="gra-2" xlink:href="img/eSC03-fig2.jpg"/>
			</fig>			

			<p>Even if this altitudinal and habitat pattern can be considered as an exception, Pyrenean chamois could inhabit forests with strong slopes below 700 m permanently. This would mean a significant increase in its distribution range surpassing the current consideration of the species as an exclusively high mountain dweller. In any case, steep slopes are a fundamental requirement for the reproduction of chamois since females need vertical areas to give birth and have their young protected from terrestrial predators when they cannot flee (Pérez-Barbería &amp; Nores, 1994).</p>

			<p>It has to be considered that global warming (Lovari et al., 2020), encroachment (García-Ruíz et al., 2021) and forest increase (Lasanta &amp; Vicente-Serrano, 2007) are reducing chamois habitat and quality. The occupation of these low elevation areas can represent an alternative to their traditional habitat. This preliminary work is the first document that shows a Pyrenean chamois population living all year long for a long period under 1000 m of altitude. Additional work is needed for corroborating this potential trend and studying the variables involved in this process.</p>
		</sec><!--//sec3-->
		
	</body>
	<back>
		<author-notes>
			<title>Authors’ contributions</title>
			<fn>Conceptualization: J. Herrero</fn>
			<fn>Data curation: J. Herrero, A. García-Serrano</fn>
			<fn>Formal analysis: J. Herrero, A. García-Serrano</fn>
			<fn>Funding acquisition: A. García-Serrano</fn>
			<fn>Investigation: J. Herrero, Carlos Félez, Fernando Herrero, A. Machuca, B. Ponz, S. Sancho</fn>
			<fn>Methodology: J. Herrero, Carlos Félez, Fernando Herrero, A. Machuca, B. Ponz, S. Sancho</fn>
			<fn>Project administration: A. García-Serrano</fn>
			<fn>Resources: A. García-Serrano</fn>
			<fn>Software: A. García-Serrano</fn>
			<fn>Supervision: J. Herrero, A. García-Serrano</fn>
			<fn>Validation: J. Herrero, A. García-Serrano</fn>
			<fn>Visualization: J. Herrero, A. García-Serrano</fn>
			<fn>Writing – original draft: J. Herrero</fn>
			<fn>Writing – review &amp; editing: J. Herrero, A. García-Serrano</fn>
		</author-notes>

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