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<front>
<journal-meta>
<journal-id journal-id-type="index">3220</journal-id>
<journal-title-group>
<journal-title specific-use="original" xml:lang="es">Revista de la Sociedad Entomológica Argentina</journal-title>
<abbrev-journal-title abbrev-type="publisher" xml:lang="es">RSEA</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1851-7471</issn>
<publisher>
<publisher-name>Sociedad Entomológica Argentina</publisher-name>
<publisher-loc>
<country>Argentina</country>
<email>gsanblas@mendoza-conicet.gob.ar</email>
</publisher-loc>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="art-access-id" specific-use="redalyc">322083084003</article-id>
<article-id pub-id-type="doi">10.25085/rsea.840408</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Artículos</subject>
</subj-group>
</article-categories>
<title-group>
<article-title xml:lang="en">
<bold>Wild Bee Diversity in Parque Nacional Volcán Isluga: Assessing Plant-Bee Interactions in a High-Altitude Desert</bold>
</article-title>
<trans-title-group>
<trans-title xml:lang="es">
<bold>Explorando la diversidad de abejas silvestres en desierto de gran altitud del Parque Nacional Volcán Isluga</bold>
</trans-title>
</trans-title-group>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="no">
<name name-style="western">
<surname>PINHEIRO-COSTA</surname>
<given-names>Bruna K.</given-names>
</name>
<xref ref-type="aff" rid="aff1"/>
<xref ref-type="aff" rid="aff2"/>
<email>bruna_kpc@yahoo.com.br</email>
</contrib>
<contrib contrib-type="author" corresp="no">
<name name-style="western">
<surname>CORTÉS-RIVAS</surname>
<given-names>Benito</given-names>
</name>
<xref ref-type="aff" rid="aff3"/>
<xref ref-type="aff" rid="aff4"/>
<email>benitocortes.r@gmail.com</email>
</contrib>
<contrib contrib-type="author" corresp="no">
<name name-style="western">
<surname>PÁDUA</surname>
<given-names>Diego G.</given-names>
</name>
<xref ref-type="aff" rid="aff5"/>
<email>paduadg@gmail.com</email>
</contrib>
<contrib contrib-type="author" corresp="no">
<name name-style="western">
<surname>MOREIRA-MUÑOZ</surname>
<given-names>Andrés</given-names>
</name>
<xref ref-type="aff" rid="aff6"/>
<email>andres.moreira@pucv.cl</email>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name name-style="western">
<surname>ARAUJO</surname>
<given-names>Rodrigo O.</given-names>
</name>
<xref ref-type="corresp" rid="corresp1"/>
<xref ref-type="aff" rid="aff7"/>
<email>araujorodrigodeoliveira@gmail.com</email>
</contrib>
</contrib-group>
<aff id="aff1">
<institution content-type="original">Programa de Doctorado en Salud Ecosistémica, Centro de Investigación de Estudios Avanzados del Maule, Universidad Católica del Maule, Talca, Chile</institution>
<country country="CL">Chile</country>
<institution-wrap>
<institution content-type="orgname">Universidad Católica del Maule</institution>
</institution-wrap>
</aff>
<aff id="aff2">
<institution content-type="original">Laboratorio de Ecología de Abejas, Departamento de Ciencias Biológicas y Químicas, Facultad de Ciencias Básicas, Universidad Católica del Maule, Talca, Chile</institution>
<country country="CL">Chile</country>
<institution-wrap>
<institution content-type="orgname">Universidad Católica del Maule</institution>
</institution-wrap>
</aff>
<aff id="aff3">
<institution content-type="original">Laboratorio de Ecología de Abejas, Departamento de Ciencias Biológicas y Químicas, Facultad de Ciencias Básicas, Universidad Católica del Maule, Talca, Chile</institution>
<country country="CL">Chile</country>
<institution-wrap>
<institution content-type="orgname">Universidad Católica del Maule</institution>
</institution-wrap>
</aff>
<aff id="aff4">
<institution content-type="original">Laboratorio de Entomología General y Aplicada, Centro de Investigación de Estudios Avanzados del Maule, Vicerrectoría de Investigación y Postgrado, Universidad Católica del Maule, Talca, Chile</institution>
<country country="CL">Chile</country>
<institution-wrap>
<institution content-type="orgname">Universidad Católica del Maule</institution>
</institution-wrap>
</aff>
<aff id="aff5">
<institution content-type="original">Laboratorio de Entomología General y Aplicada, Centro de Investigación de Estudios Avanzados del Maule, Vicerrectoría de Investigación y Postgrado, Universidad Católica del Maule, Talca, Chile</institution>
<country country="CL">Chile</country>
<institution-wrap>
<institution content-type="orgname">Universidad Católica del Maule</institution>
</institution-wrap>
</aff>
<aff id="aff6">
<institution content-type="original">Instituto de Geografía, Pontificia Universidad Católica de Valparaíso, Valparaíso, Chile</institution>
<country country="CL">Chile</country>
<institution-wrap>
<institution content-type="orgname">Pontificia Universidad Católica de Valparaíso</institution>
</institution-wrap>
</aff>
<aff id="aff7">
<institution content-type="original">Laboratorio de Entomología General y Aplicada, Centro de Investigación de Estudios Avanzados del Maule, Vicerrectoría de Investigación y Postgrado, Universidad Católica del Maule, Talca, Chile</institution>
<country country="CL">Chile</country>
<institution-wrap>
<institution content-type="orgname">Universidad Católica del Maule</institution>
</institution-wrap>
</aff>
<author-notes>
<corresp id="corresp1">
<email>araujorodrigodeoliveira@gmail.com</email>
</corresp>
</author-notes>
<pub-date pub-type="epub-ppub">
<year>2025</year>
</pub-date>
<volume>84</volume>
<issue>4</issue>
<elocation-id>e0408</elocation-id>
<history>
<date date-type="received" publication-format="dd mes yyyy">
<day>15</day>
<month>05</month>
<year>2025</year>
</date>
<date date-type="accepted" publication-format="dd mes yyyy">
<day>25</day>
<month>09</month>
<year>2025</year>
</date>
</history>
<permissions>
<ali:free_to_read/>
</permissions>
<abstract xml:lang="en">
<title>Abstract</title>
<p>Bees are highly responsive to floral resource availability and environmental variability, making them valuable biological indicators of habitat degradation, land-use change, and agricultural expansion. Despite the harsh conditions of arid and high-altitude ecosystems, these environments support remarkable bee diversity and endemism. However, bee diversity, as well as the identity of floral resources that support insect populations, remains underexplored, particularly in the high-altitude ecosystems of northern Chile, which are underrepresented in the scientific literature. This study assesses bee diversity and plant-bee interactions in a high-altitude desert ecosystem in the Volcán Isluga National Park, located in the Tarapacá region, an area of scientific interest due to mining activities. Fieldwork was conducted in two pre-Andean villages between May and June 2023. A total of 71 bees from 19 morphospecies, 13 genera, and four families were collected. The family Apidae was the most abundant, followed by Halictidae, Colletidae, and Megachilidae. Bees interacted with 18 plant species, with Fabaceae and Asteraceae being the most represented families. These findings highlight the role of local flora in supporting bee communities and underscore the need for conservation strategies to maintain ecosystem services and ensure sustained pollination services.<bold/>
</p>
</abstract>
<trans-abstract xml:lang="es">
<title>Resumen</title>
<p>Las abejas son sensibles a la disponibilidad de recursos florales y a los cambios ambientales, lo que las convierte en valiosos indicadores biológicos de la degradación del hábitat, los cambios en el uso del suelo y la expansión agrícola. Sin embargo, la diversidad de las abejas, así como la identificación de los recursos florales que sustentan las poblaciones de estos insectos, siguen sin estar suficientemente estudiadas, especialmente en los ecosistemas de gran altitud del norte de Chile, que están poco representados en la literatura científica. Este estudio evalúa la diversidad de abejas y las interacciones planta-abeja en el ecosistema desértico de gran altitud del Parque Nacional Volcán Isluga, ubicado en la región de Tarapacá, una zona de interés científico debido a las actividades mineras. El trabajo de campo se realizó en dos localidades preandinas entre mayo y junio de 2023. Se colectaron un total de 71 abejas de 19 morfoespecies, 13 géneros y cuatro familias. La familia Apidae fue la más abundante, seguida de Halictidae, Colletidae y Megachilidae. Las abejas interactuaron con 18 especies de plantas, siendo Fabaceae y Asteraceae las familias más representadas. Estos hallazgos resaltan el papel de la flora local en el apoyo a las comunidades de abejas y subrayan la necesidad de estrategias de conservación para mantener los servicios ecosistémicos y garantizar la gestión sostenible de los servicios de polinización.</p>
</trans-abstract>
<kwd-group xml:lang="en">
<title>Keywords</title>
<kwd>Bee conservation</kwd>
<kwd>Chilean bees</kwd>
<kwd>Desertic zone</kwd>
<kwd>Pollination</kwd>
</kwd-group>
<kwd-group xml:lang="es">
<title>Palabras clave</title>
<kwd>Abejas chilenas</kwd>
<kwd>Conservación de abejas</kwd>
<kwd>Polinización</kwd>
<kwd>Zona desértica</kwd>
</kwd-group>
<counts>
<fig-count count="3"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="71"/>
</counts>
<custom-meta-group>
<custom-meta>
<meta-name>redalyc-journal-id</meta-name>
<meta-value>3220</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec>
<title>
<bold>INTRODUCTION</bold>
</title>
<p>Bees play an important role in ecosystems as pollinators, contributing to plant reproduction, agricultural productivity, and biodiversity conservation (<xref ref-type="bibr" rid="redalyc_322083084003_ref29">Klein et al., 2007</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref55">Potts et al., 2010</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref47">Ollerton et al., 2011</xref>). Due to their sensitivity to environmental change and floral resource availability, they are valuable bioindicators of habitat degradation and land-use change (<xref ref-type="bibr" rid="redalyc_322083084003_ref45">Odanaka &amp; Rehan, 2019</xref>). Despite the increasing global recognition of pollinators' importance, significant knowledge gaps persist regarding their distribution and diversity, particularly in arid and high-altitude ecosystems (<xref ref-type="bibr" rid="redalyc_322083084003_ref4">Arroyo et al., 1982</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref23">Gonzalez &amp; Engel, 2004</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref25">Henríquez-Piskulich et al., 2020</xref>).</p>
<p>In arid and high-altitude ecosystems, bee diversity and distribution patterns are influenced by extreme environmental conditions such as low temperatures, high solar radiation, and scarce water availability (<xref ref-type="bibr" rid="redalyc_322083084003_ref1">Abrahamczyk et al., 2011</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref51">Orr et al., 2021</xref>). These ecosystems tend to support specialized bee communities adapted to harsh climatic conditions, often characterized by a dominance of ground-nesting species and seasonal activity patterns closely linked to the availability of floral resources (<xref ref-type="bibr" rid="redalyc_322083084003_ref41">Minckley et al., 2000</xref>). In addition, the fragmented nature of the vegetation and the presence of endemic plant species contribute to unique plant-bee interactions (<xref ref-type="bibr" rid="redalyc_322083084003_ref4">Arroyo et al., 1982</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref41">Minckley et al., 2000</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref24">Henríquez-Piskulich et al., 2018</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref35">Luna et al., 2024</xref>) that are particularly vulnerable to environmental disturbances, making their conservation a priority (<xref ref-type="bibr" rid="redalyc_322083084003_ref24">Henríquez-Piskulich et al., 2018</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref64">Rodríguez et al., 2021</xref>).</p>
<p>In Chile, bee diversity has mainly been studied in the central and southern regions, while the northern high-altitude ecosystems remain underrepresented in the scientific literature (<xref ref-type="bibr" rid="redalyc_322083084003_ref42">Montalva &amp; Ruz, 2010;</xref>
<xref ref-type="bibr" rid="redalyc_322083084003_ref25">Henríquez-Piskulich et al., 2020</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref33">López-Aliste et al., 2021</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref36">Marshall et al., 2023</xref>). The northern region of Chile is characterized by an arid and semi-arid climate, with extreme variations in temperature and solar radiation (<xref ref-type="bibr" rid="redalyc_322083084003_ref5">Arroyo &amp; Cavieres, 2013</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref2">Anthelme et al., 2014</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref9">Cavieres et al., 2025</xref>). As a result, these areas support an ecosystem that is favorable to diversity and evolution of endemic forms (<xref ref-type="bibr" rid="redalyc_322083084003_ref51">Orr et al., 2021</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref36">Marshall et al., 2023</xref>).</p>
<p>The Volcán Isluga National Park, located in the Tarapacá Region, is a protected area characterized by high-altitude desert ecosystems that harbor a rich diversity of native flora and Andean vegetation (<xref ref-type="bibr" rid="redalyc_322083084003_ref14">CONAF, 1988</xref>). This Park has been declared of scientific interest, which restricts mining activities (<xref ref-type="bibr" rid="redalyc_322083084003_ref13">CHILE, 1983</xref>), and natural resource extraction and land use are strictly regulated under a conservation framework that aims to sustain biodiversity and maintain key ecosystem services provided by the local species (<xref ref-type="bibr" rid="redalyc_322083084003_ref49">Ormazabal, 1993</xref>). However, studies on the diversity and ecological interactions of insects in this region remain limited. Bees are the main pollinators of flowering plants (<xref ref-type="bibr" rid="redalyc_322083084003_ref47">Ollerton et al., 2011</xref>), supporting essential ecological processes such as plant reproduction, fruit production, and ecological succession (<xref ref-type="bibr" rid="redalyc_322083084003_ref29">Klein et al., 2007</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref55">Potts et al., 2010)</xref>. In this sense, this study aims to evaluate the diversity and composition of wild bees and provide a list of plant-bee interactions in Vólcan Isluga National Park. Based on these results, we discuss the relevance of the understanding of bee diversity and their interactions with native plant species as a key component for the conservation of pollination services.</p>
</sec>
<sec>
<title>
<bold>MATERIAL AND METHODS </bold>
</title>
<sec>
<title>
<bold>Study site</bold>
</title>
<p>We conducted field sampling during May and June 2023, at two pre-Andean villages of Chiapa and Jaina located on Volcán Isluga National Park (19°15' S, 43°31' W), Provincia de Iquique, in the Tarapacá Region, Chile. This Park covers a total area of 174,744 ha and encompasses a variety of desert vegetation zones adapted to high-altitude and arid climatic conditions (<xref ref-type="fig" rid="gf1">Fig. 1</xref>). The park's elevation ranges from 800 to 5,550 m (<xref ref-type="bibr" rid="redalyc_322083084003_ref14">CONAF, 1988</xref>). The climate in the study areas is predominantly classified as Hot Desert (BWh), with a moderate influence from the Cold Desert climate (BWk) (<xref ref-type="bibr" rid="redalyc_322083084003_ref69">Sarricolea et al., 2017</xref>). In general, summers are warm, while winters are cold and dry, with temperatures ranging from -20 °C to 10 °C. Rainfall is variable throughout the year, reaching 50 to 350 mm, primarily between December and March (<xref ref-type="bibr" rid="redalyc_322083084003_ref14">CONAF, 1988</xref>). Volcán Isluga National Park is a legally protected area in which mineral exploration and exploitation are restricted by the Mining Code Regulations (<xref ref-type="bibr" rid="redalyc_322083084003_ref13">Chile, 1983</xref>; Law 18,248). In addition to its legal status as a biodiversity refuge, the park is recognised as a zone of development for Aymara indigenous communities, highlighting its socio-ecological importance in northern Chile (<xref ref-type="bibr" rid="redalyc_322083084003_ref74">Toledo et al., 2017</xref>).</p>
<p>The most representative vegetation zones include the tropical Andean low shrubland (literal translation of Matorral bajo tropical andino), dominated by <italic>Parastrephia lepidophylla</italic> (Weed) Cabrera and <italic>P. quadrangularis </italic>(Meyen) Cabrera (Asteraceae), commonly known as <italic>tolar</italic>, which occurs above 3,000 meters above sea level. Between 2,500 and 3,000 meters, the landscape is characterized by the same vegetation type, with dominance of the endemic <italic>Fabiana ramulosa</italic> (Wedd.) Hunz &amp; Barboza (Solanaceae) and <italic>Diplostephium meyenii </italic>Wedd (Asteraceae). The northeastern section of the National Park corresponds to high-altitude zones with tropical Andean low shrublands (Matorral bajo tropical andino) dominated by <italic>Parastrephia lucida</italic> (Meyen) Cabrera (Asteraceae) and <italic>Azorella compacta </italic>Phil. (Apiaceae), and in some areas this species is accompanied by <italic>Pycnophyllum molle</italic> J. Remy (Caryophyllaceae). In contrast, the southwestern section includes tropical Andean desert low shrubland (<italic>Matorral bajo desértico tropical andino</italic>), characterized by <italic>Atriplex imbricata</italic> (Moq.) D. Dietr. (Amaranthaceae) and <italic>Aloysia deserticola </italic>(Phil.) Lu-Irving &amp; N.O'Leary (Verbenaceae); tropical Andean spiny forest (<italic>Bosque espinoso tropical andino</italic>), with species such as <italic>Browningia candelaris</italic> (Meyen) Britton &amp; Rose and <italic>Corryocactus brevistylus </italic>(K. Schum. ex Vaupel) Britton &amp; Rose (Cactaceae), representing the columnar cactus belt; and interior tropical Andean desert low shrubland (<italic>Matorral bajo desértico tropical interior</italic>), dominated by <italic>Adesmia atacamensis </italic>Phil. (Fabaceae) and <italic>Silvaea salsoloides </italic>(Barnéoud) Teillier &amp; S.T. Ibáñez (Montiaceae), marking the transition to lower, more arid zones (<xref ref-type="bibr" rid="redalyc_322083084003_ref34">Luebert &amp; Pliscoff, 2017</xref>).</p>
<p>
<fig id="gf1">
<label>
<bold>Figure 1</bold>
</label>
<caption>
<title>
<bold>Study site within the Volcán Isluga National Park, Tarapacá Region, northern Chile</bold>
</title>
<p>The map illustrates different vegetation types adapted to high-altitude and arid climatic conditions. Green dots indicate nearby settlements (villages/hamlets) used as geographic references.</p>
</caption>
<alt-text>Figure 1 Study site within the Volcán Isluga National Park, Tarapacá Region, northern Chile</alt-text>
<graphic xlink:href="322083084003_gf2.png" position="anchor" orientation="portrait">
<alt-text>Figure 1 Study site within the Volcán Isluga National Park, Tarapacá Region, northern Chile</alt-text>
</graphic>
</fig>
</p>
<p>The vegetation map was produced using vegetation belt data downloaded from <underline>www.geoportal.cl</underline> (accessed on May 01, 2025) (originally described by <xref ref-type="bibr" rid="redalyc_322083084003_ref34">Luebert &amp; Pliscoff, 2017</xref>) and processed in ArcGIS Pro 2.8.8 software.</p>
</sec>
<sec>
<title>
<bold>Wild bee sampling </bold>
</title>
<p>We employed two complementary sampling methods during May and June 2023 field seasons. The combination of active and passive approaches was selected to maximize the likelihood of capturing exploratory data on the richness and abundance of the regional bee community, as recommended in pollinator survey protocols (e.g., <xref ref-type="bibr" rid="redalyc_322083084003_ref82">Westphal et al., 2008</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref66">Roulston et al., 2007</xref>).</p>
<p>For the passive collection, we used yellow pan traps (Moericke traps) consisting of 4.5 cm high, 10 cm diameter plastic bowls filled with 150 mL of water mixed with a few drops detergent to break the surface tension (Wang et al., 2017). A total of five traps were installed simultaneously along a 350 m fixed transect for five consecutive days, resulting in 25 individual trap deployments. The traps were placed at ground level in open areas near flowering vegetation, spaced approximately 50 m apart to maximize the likelihood of capturing floral visitors. Each trap was left in the field for 24 h, yielding a cumulative sampling effort of 600 trap-hours.</p>
<p>In addition, active searches were conducted along transects between 09:00 and 18:00, with two researchers conducting a total of 45 hours per observer. Six sampling points were randomly selected, ensuring a minimum distance of 1,500 meters between points to minimize spatial clustering. At each point, observers were not restricted to a fixed transect line but moved slowly in search of flowering plants, taking care not to stray more than 600 meters from the designated location (adapted from <xref ref-type="bibr" rid="redalyc_322083084003_ref82">Westphal et al., 2008</xref>). This transect walk approach was chosen to maximize the likelihood of encountering flowering individuals during the sampling period, given the presence of empty areas without flowering plants (<xref ref-type="bibr" rid="redalyc_322083084003_ref82">Westphal et al., 2008</xref>). Direct searching along variable transects can enhance the representativeness of bee foraging activity across temporal and spatial scales, especially in fragmented or arid environments, compared to fixed transect corridors (<xref ref-type="bibr" rid="redalyc_322083084003_ref15">Dafni et al., 2005</xref>). However, it should be noted that sampling effort per plant species (limited to individual plants) and the presence or abundance of plants across transects could not be standardized or controlled. Collectors remained in front of a flowering individual for approximately five minutes and captured all visiting bees using an entomological net. A plant-bee interaction matrix was constructed based on all interactions between bees and flowering plants recorded by the observers during the surveys. The sampling points and pan traps are provided in Supplementary Table S1.</p>
<p>
<fig id="gf2">
<label>
<bold>Figure 2</bold>
</label>
<caption>
<title>
<bold>Representative bee specimens collected during this study</bold>
</title>
<p>a. <italic>Alloscirtetica</italic>
<italic> weyrauchi</italic> Michener, LaBerge &amp; Moure, 1955. b. <italic>Anthophora</italic>
<italic> arequipensis</italic> Bréthes, 1920. c. <italic>Centris buchholzi</italic> Herbst, 1918. d. <italic>Centris sacsayhuaman</italic> Vivallo, 2020. e. <italic>Xylocopa viridigastra</italic> Lepeletier, 1841. f. <italic>Caenohalictus</italic>
<italic> cuprellus</italic> (Vachal, 1903). g. <italic>Lasioglossum</italic> sp. Scale bar= a, b, c, f, g: 2 mm; d, e: 5 mm</p>
</caption>
<alt-text>Figure 2 Representative bee specimens collected during this study</alt-text>
<graphic xlink:href="322083084003_gf3.png" position="anchor" orientation="portrait">
<alt-text>Figure 2 Representative bee specimens collected during this study</alt-text>
</graphic>
</fig>
</p>
<p>The specimens observed during this study were collected, prepared, and deposited in the Entomological Collection of the Laboratorio de Ecología de Abejas, Universidad Católica del Maule (UCM). Bees were identified to the lowest taxonomic level possible, usually species, using interactive keys and reference literature, including Michener et al. (<xref ref-type="bibr" rid="redalyc_322083084003_ref40">1955</xref>); Chiappa et al. (<xref ref-type="bibr" rid="redalyc_322083084003_ref11">1990</xref>); Toro (<xref ref-type="bibr" rid="redalyc_322083084003_ref76">1999</xref>); Rojas &amp; Toro (<xref ref-type="bibr" rid="redalyc_322083084003_ref65">2000</xref>); Michener (<xref ref-type="bibr" rid="redalyc_322083084003_ref39">2007</xref>); and Ferrari (<xref ref-type="bibr" rid="redalyc_322083084003_ref20">2017</xref>). Benito Cortés-Rivas (author) conducted the identification and validation of the specimens by comparing them with reference material from the entomological collections of the Laboratorio de Ecología de Abejas at the Universidad Católica del Maule. Experienced bee taxonomist Felipe Vivallo (Museo Nacional do Rio de Janeiro, Brazil) provided confirmation for specimens from <italic>Centris</italic> genus. All plant species were identified at the species level by A. Moreira-Muñoz (author). Insect sociability was recorded based on the reference literature (<xref ref-type="bibr" rid="redalyc_322083084003_ref37">Melander, 1902</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref21">Gerling et al., 1989</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref38">Michener, 1964</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref76">Toro et al., 1999;</xref>
<xref ref-type="bibr" rid="redalyc_322083084003_ref10">Chiappa et al., 2000</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref70">Silveira et al., 2002</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref22">González &amp; Chávez, 2004</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref53">Packer, 2006</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref16">Davis et al., 2012</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref43">Montalva et al., 2012;</xref>
<xref ref-type="bibr" rid="redalyc_322083084003_ref62">Richards &amp; Packer, 2021</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref33">López-Aliste et al., 2021</xref>).</p>
<p>A heatmap was constructed using the “ggplot2” package in R (<xref ref-type="bibr" rid="redalyc_322083084003_ref61">R Core Team, 2023</xref>), based on the abundance matrix of bee visitors and host plants to visualize the interactions between bee species and their associated plant species. To contextualize our findings within the broader knowledge of bee diversity in northern Chile, we conducted a comparative analysis with published records for the Tarapacá region. Specifically, we used the checklist compiled by Toro (<xref ref-type="bibr" rid="redalyc_322083084003_ref75">1986</xref>); López-Aliste et al. (<xref ref-type="bibr" rid="redalyc_322083084003_ref33">2021</xref>).</p>
<p>Bees' images were captured with a Leica S9i stereomicroscope using an LED illumination dome (<xref ref-type="bibr" rid="redalyc_322083084003_ref28">Kawada &amp; Buffington, 2016</xref>) and the Leica Application Suite x extended-focus software. Images stacking was done using Helicon Focus 8.1.0, followed by editing in Adobe Photoshop 2020.</p>
</sec>
</sec>
<sec>
<title>
<bold>RESULTS</bold>
</title>
<p>A total of 71 bees representing 19 morphospecies in 13 genera and four families were collected during the sampling periods (<xref ref-type="fig" rid="gf2">Figure 2</xref>; <xref ref-type="table" rid="gt1">Table I</xref>). The Apidae family was the most abundant, comprising 35.5 % of the specimens (n= 27 specimens; 7 species), followed by Halictidae (26.3 %; n= 20 individuals; 4 species), Colletidae (17.1 %; n= 13 individuals; 3 species), and Megachilidae (14.5 %; n= 11 individuals; 5 species). No individuals from the family Andrenidae were recorded.</p>
<p>Among the collected specimens, Apidae exhibited the highest species richness and abundance, with notable species including <italic>Alloscirtetica weyrauchi</italic> Michener<italic>, </italic>LaBerge &amp; Moure, 1955, <italic>Anthophora arequipensis </italic>Bréthes, 1920, and multiple <italic>Centris</italic> species (<italic>C. buchholzi </italic>Herbst, 1918<italic>, C. escomeli </italic>Cockerell, 1926<italic>, C. sacsayhuaman </italic>Vivallo, 2020<italic>,</italic> and<italic> C. miluni</italic> Vivallo, 2020), as well as <italic>Xylocopa viridigastra</italic> Lepeletier, 1841 (<xref ref-type="fig" rid="gf3">Fig. 3</xref>). The most abundant species were <italic>A. arequipensis</italic> (n= 12 specimens, 16.9 %), <italic>Caenohalictus cuprellus</italic> (Vachal, 1903) (n= 10 specimens, 14.1 %, <xref ref-type="table" rid="gt1">Table I</xref>). Bee species exhibited distinct social behaviors, including solitary, cleptoparasitic and variations of social patterns behaviour (<xref ref-type="table" rid="gt1">Table I</xref>).</p>
<p>
<table-wrap id="gt1">
<label>Table I</label>
<caption>
<title>
<bold>List of bee morphospecies collected in Volcán Isluga National Park, including their ecology and records of floral interactions</bold>
</title>
</caption>
<alt-text>Table I List of bee morphospecies collected in Volcán Isluga National Park, including their ecology and records of floral interactions</alt-text>
<graphic xlink:href="322083084003_gt2.png" position="anchor" orientation="portrait">
<alt-text>Table I List of bee morphospecies collected in Volcán Isluga National Park, including their ecology and records of floral interactions</alt-text>
</graphic>
<table-wrap-foot>
<fn id="fn4" fn-type="other">
<label>
<italic>Notes</italic>
</label>
<p>
<italic>Classification of bee species by body size: S= small (5-8 mm), M= medium (9-16 mm), G= large (&gt; 17 mm), based on body width</italic>
</p>
</fn>
</table-wrap-foot>
</table-wrap>
</p>
<p>
<fig id="gf3">
<label>
<bold>Figure 3.</bold>
</label>
<caption>
<title>
<bold>Abundance visitation matrix. Heatmap showing the abundance and interaction between bee and plant species recorded in Volcán Isluga</bold>
<bold>National Park</bold>
</title>
<p>The vertical columns represent bee species, while each horizontal row corresponds to a plant species. The color intensity of the heatmap reflects the number of observed floral visits, with darker shades indicating higher interaction frequencies</p>
</caption>
<alt-text>Figure 3. Abundance visitation matrix. Heatmap showing the abundance and interaction between bee and plant species recorded in Volcán Isluga National Park</alt-text>
<graphic xlink:href="322083084003_gf4.png" position="anchor" orientation="portrait">
<alt-text>Figure 3. Abundance visitation matrix. Heatmap showing the abundance and interaction between bee and plant species recorded in Volcán Isluga National Park</alt-text>
</graphic>
</fig>
</p>
<p>Regarding the sampling method, only one specimen of <italic>Lasioglossum</italic> sp. 1 (Halictidae) was collected using yellow pan traps, while the remaining individuals were recorded using the active sweep net method (<xref ref-type="table" rid="gt1">Table I</xref>). Bees were recorded interacting with 18 plant species belonging to 9 botanical families. Fabaceae (n= 7 species; 18 specimens) and Asteraceae (n= 5 species; 18 specimens) showed the highest number of plants species, while Malvaceae, represented by one single species, was the most abundant family (27 specimens). The remaining families, Calceolariaceae, Caryophyllaceae, Francoaceae, Lamiaceae, Loasaceae, Solanaceae, and Vivianiaceae, were each represented by a single species (<xref ref-type="fig" rid="gf3">Fig. 3</xref>).</p>
<p>Among the 19 bees morphospecies recorded in our study, 12 had previously documented distributions for the Tarapacá region. Notably, our survey recorded <italic>Centris miluni</italic> and <italic>Centris sacsayhuaman</italic>, two species that were only recently described (<xref ref-type="bibr" rid="redalyc_322083084003_ref79">Vivallo et al., 2020</xref>), as well as <italic>Caupolicana albiventris</italic>, <italic>Xeromelissa wilmattae</italic>, and <italic>Nothanthidium bizonatum</italic>, which were previously documented only by Toro (<xref ref-type="bibr" rid="redalyc_322083084003_ref75">1986</xref>). Another relevant record, <italic>Anthidium funereum</italic>, had only been documented previously for the Arica region (<xref ref-type="bibr" rid="redalyc_322083084003_ref75">Toro, 1986</xref>).</p>
</sec>
<sec>
<title>
<bold>DISCUSSION</bold>
</title>
<p>Our findings highlight the diversity and abundance of wild bees in Volcán Isluga National Park, the first assessment of plant-bee interactions in this high-altitude desert ecosystem in this region. Our exploration of plant–bee interactions revealed that 19 bee species interacted with 18 native plant species, primarily from the Fabaceae and Asteraceae families. These results suggest that the local flora may play a key role in supporting solitary bees, particularly within the Apidae and Halictidae families.</p>
<p>According to López-Aliste et al. (<xref ref-type="bibr" rid="redalyc_322083084003_ref33">2021</xref>), the study region has a low frequency of documented bee records. In their most recent survey of the Tarapacá region, 26 bee species were reported, ten of which were also observed in our study (See <xref ref-type="table" rid="gt1">Table I</xref>). The presence of <italic>Centris miluni, Centris sacsayhuaman, Caupolicana albiventris, Xeromelissa wilmattae</italic>, and <italic>Nothanthidium bizonatum</italic> in our survey underscores the Tarapacá region's understudied bee diversity. Several of these species have been recently described or were previously documented only in distant locations (<xref ref-type="bibr" rid="redalyc_322083084003_ref79">Vivallo et al., 2020</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref75">Toro, 1986</xref>), which suggests that their geographic ranges are likely broader than currently recognized. It is also important to highlight that the identification of some morphospecies was limited to the genus level due to outdated or incomplete taxonomic references for certain groups (e.g., <italic>Lasioglossum</italic>). This limitation reflects the taxonomic challenges documented in recent literature, particularly for morphologically difficult and cryptic taxa such as <italic>Lasioglossum</italic> (Halictidae) (<xref ref-type="bibr" rid="redalyc_322083084003_ref32">Landaverde-González et al., 2017</xref>).</p>
<p>Studies in arid desert ecosystems have revealed high bee diversity of interest (<xref ref-type="bibr" rid="redalyc_322083084003_ref3">Argueta-Guzmán et al., 2022</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref73">Thapa‐Magar et al., 2022</xref>). The species share adaptations that allow them to survive in arid, high-altitude conditions where climatic extremes (<xref ref-type="bibr" rid="redalyc_322083084003_ref67">Sakagami &amp; Munakata, 1972</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref18">Eickwort et al., 1996</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref59">Purcell et al., 2015</xref>). These adaptations include specific body traits such as increased hair density for thermoregulation (<xref ref-type="bibr" rid="redalyc_322083084003_ref78">Turnley et al., 2025</xref>) and predominantly solitary behaviour (<xref ref-type="bibr" rid="redalyc_322083084003_ref44">Moss &amp; While, 2021</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref52">Ostwald et al., 2024</xref>). In addition to arid conditions, the availability of floral resources, both spatially and temporally (<xref ref-type="bibr" rid="redalyc_322083084003_ref71">Snyder et al., 2006</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref60">Pueyo et al., 2008</xref>), appear to play an important role in determining bee richness and occurrence in these ecosystems (<xref ref-type="bibr" rid="redalyc_322083084003_ref64">Rodríguez et al., 2021</xref>).</p>
<p>In Volcán Isluga, most of the recorded species were solitary and visited especially native plants, such as <italic>Tarasa operculata</italic> (Malvaceae), <italic>Grindelia tarapacana </italic>(Asteraceae) and <italic>Lupinus oreophilus </italic>(Fabaceae). These species are typically found in the herbaceous and shrubby vegetation of the desert Matorral system, which occurs on the dry western slopes of the Andes. Some species, such as <italic>Tarasa operculata</italic>, are adapted to higher altitudes (ranging from 1,600 to 2,300 m) and to conditions with limited water availability (<xref ref-type="bibr" rid="redalyc_322083084003_ref12">Chicalla-Rios, 2021</xref>). Native bees were observed collecting pollen and other floral resources, indicating that the presence of these plant species in the desert landscape may serves as key resources for sustaining bee communities and maintaining ecosystem stability.</p>
<p>Furthermore, our study presents limitations that may affect the generalizability of its findings. Although it provides an initial insight into plant-bees interaction, we did not quantify the influence of plant abundance and diversity across spatial heterogeneity gradients on plant-bee interactions. The availability and distribution of floral resources can directly impact bee foraging behavior (<xref ref-type="bibr" rid="redalyc_322083084003_ref19">Escobedo-Kenefic et al., 2020</xref>), resource partitioning, and the overall structure of pollination networks (generalist or specialist) (<xref ref-type="bibr" rid="redalyc_322083084003_ref6">Bendel et al., 2019</xref>). Although we combined different collection methods, our study was conducted in a limited time period, which restricts our ability to capture seasonal variations in bee activity and floral resource availability.  Since the composition and level of specialization of plant-bee interactions are strongly influenced by phenological cycles (<xref ref-type="bibr" rid="redalyc_322083084003_ref50">Oertli et al., 2005</xref>), future studies should incorporate multiseasonal sampling to adequately characterize community dynamics and assess the effect of temporal variability on pollination networks in high-altitude desert ecosystems (<xref ref-type="bibr" rid="redalyc_322083084003_ref6">Bendel et al., 2019</xref>).</p>
<p>Combining active and passive sampling methods provides a more comprehensive assessment of bee diversity and plant-bee interactions (<xref ref-type="bibr" rid="redalyc_322083084003_ref81">Wilson et al., 2016</xref>). However, the success of each type of method can vary depending on climatic conditions, seasonality, suitable habitat, trap color and insect body size (<xref ref-type="bibr" rid="redalyc_322083084003_ref57">Prendergast et al., 2020</xref>). In our study, the effectiveness of passive collection methods, such as pan traps, was notably lower, recording only a single bee species. This result probably reflects the limitations of trap color selection, and the small number of samples taken, which may have reduced the ability to capture the full diversity of bee species present in the study area. To overcome these limitations, some studies have employed multi-colored pan traps combined with standardized active surveys, which improves the representativeness of bee community overviews in terms of both diversity and abundance (<xref ref-type="bibr" rid="redalyc_322083084003_ref80">Vereecken et al., 2021</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref26">Hutchinson et al., 2022</xref>; <xref ref-type="bibr" rid="redalyc_322083084003_ref83">Xie et al., 2024</xref>). However, these combined approaches also increase the logistical effort and financial cost of sampling and monitoring, which can limit their feasibility in remote or resource-constrained environments (<xref ref-type="bibr" rid="redalyc_322083084003_ref7">Bell et al., 2023)</xref>.</p>
<p>Despite these limitations, our study provides an important baseline characterization of plant-bee interactions in a high-altitude desert ecosystem in northern Chile. Many of the recorded interactions occur within legally protected areas, such as national parks, which help safeguard these ecological relationships. However, on a regional scale, the park is located in a zone of potential conflict due to the proximity of mining activities to protected areas (<xref ref-type="bibr" rid="redalyc_322083084003_ref74">Toledo et al., 2017</xref>). Our findings underscore the importance of ongoing monitoring and conservation efforts in the region and highlight the value of Isluga Volcano as a refuge for pollinator communities and other forms of biodiversity.</p>
</sec>
<sec>
<title>
<bold>CONCLUSION</bold>
</title>
<p>Overall, our finding suggests that bee diversity in the region is underestimated and likely higher than previously reported. The presence of previously unrecorded species further underscores the importance of documenting plant-bee interactions. These interactions can provide valuable insights into local ecological processes such as pollination and ecosystem resilience. Future research should focus on analyzing plant-bee networks, including their temporal and spatial dynamics, as well as assessing the effectiveness of bees as pollinators. Incorporating measures of pollination effectiveness, beyond simple floral visitation, will allow a better understanding of the distinct roles that different floral visitors play within the community and inform conservation strategies in these high-altitude desert ecosystems (<xref ref-type="bibr" rid="redalyc_322083084003_ref68">Santiago‐Hernández et al., 2019</xref>).</p>
</sec>
</body>
<back>
<ack>
<title>Acknowledgments</title>
<p>MOREIRA-MUÑOZ (as principal investigator) and ARAUJO (as coinvestigator) thank Agencia Nacional de Investigación y Desarrollo (ANID), Chile, for funding this study through the FONDECYT Regular nº 1221879 project. PINHEIRO-COSTA thanks UCM for the doctoral fellowship provided.</p>
</ack>
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<fn-group>
<title>Notes</title>
<fn id="fn5" fn-type="other">
<label>
<bold>DATA STATEMENT</bold>
</label>
<p>Supplementary material associated with this manuscript is available at <underline>https://doi.org/10.5281/zenodo.17042033</underline>
</p>
</fn>
<fn id="fn6" fn-type="other">
<label>
<bold>COMPETING INTERESTS</bold>
</label>
<p>The authors declare that they have no competing financial interests or personal relationships that could have appeared to influence the work reported in this paper</p>
</fn>
<fn id="fn7" fn-type="other">
<label>
<bold>AUTHORS CONTRIBUTIONS</bold>
</label>
<p>BCR, DGP, AMM and ROA conducted the fieldwork in Parque Nacional Volcán Isluga, including bee and plant sampling and specimen collection. BCR performed the morphological identification of bee specimens and specimen curation. AMM conducted the plant identification. BKPC processed ecological data and performed statistical analyses. BCR and BKPC drafted the original version of the manuscript. DGP and ROA reviewed and edited the manuscript, refining ecological interpretations and ensuring consistency across sections. AMM and ROA conceived the study, coordinated logistics, and obtained the necessary research permits and funding. All authors reviewed and approved the final version of the manuscript.</p>
</fn>
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