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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2024.1352625</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Coccinellidae on native and introduced spruce in central Europe: conservation implications in urban areas</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Jauschov&#xe1;</surname>
<given-names>Ter&#xe9;zia</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
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</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Sarva&#x161;ov&#xe1;</surname>
<given-names>Lenka</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
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<contrib contrib-type="author">
<name>
<surname>Zach</surname>
<given-names>Peter</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Saniga</surname>
<given-names>Miroslav</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
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<contrib contrib-type="author">
<name>
<surname>Martinkov&#xe1;</surname>
<given-names>Zdenka</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
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<contrib contrib-type="author">
<name>
<surname>Hon&#x11b;k</surname>
<given-names>Alois</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Skuhrovec</surname>
<given-names>Ji&#x159;&#xed;</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Holecov&#xe1;</surname>
<given-names>Milada</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Kulfan</surname>
<given-names>J&#xe1;n</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
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<aff id="aff1">
<sup>1</sup>
<institution>Institute of Forest Ecology, Slovak Academy of Sciences</institution>, <addr-line>Zvolen</addr-line>, <country>Slovakia</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Faculty of Ecology and Environmental Sciences, Technical University in Zvolen</institution>, <addr-line>Zvolen</addr-line>, <country>Slovakia</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Crop Research Institute, Functional Diversity in Agro-Ecosystems</institution>, <addr-line>Prague</addr-line>, <country>Czechia</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Department of Zoology, Faculty of Natural Sciences, Comenius University</institution>, <addr-line>Bratislava</addr-line>, <country>Slovakia</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Panagiotis Theodorou, Martin Luther University of Halle-Wittenberg, Germany</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Olivia Sanllorente, University of Granada, Spain</p>
<p>Francisco Jos&#xe9; Cabrero-Sa&#xf1;udo, Complutense University of Madrid, Spain</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Lenka Sarva&#x161;ov&#xe1;, <email xlink:href="mailto:sarvasova@ife.sk">sarvasova@ife.sk</email>; <email xlink:href="mailto:lenka.sarvasova14@gmail.com">lenka.sarvasova14@gmail.com</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>15</day>
<month>07</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>12</volume>
<elocation-id>1352625</elocation-id>
<history>
<date date-type="received">
<day>08</day>
<month>12</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>17</day>
<month>06</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Jauschov&#xe1;, Sarva&#x161;ov&#xe1;, Zach, Saniga, Martinkov&#xe1;, Hon&#x11b;k, Skuhrovec, Holecov&#xe1; and Kulfan</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Jauschov&#xe1;, Sarva&#x161;ov&#xe1;, Zach, Saniga, Martinkov&#xe1;, Hon&#x11b;k, Skuhrovec, Holecov&#xe1; and Kulfan</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>The abundance and species composition of adult ladybird communities (Coleoptera: Coccinellidae) were investigated on two congeneric trees, native Norway spruce (<italic>Picea abies</italic>) and the introduced blue spruce (<italic>Picea pungens</italic>), at four locations in Slovakia (central Europe). For two years (2021&#x2013;2022), coccinellid adults were sampled using a standard method involving beating branches at monthly intervals from April to November. Although the species composition of the communities on both spruce species was similar, the abundance of the entire coccinellid community as well as the abundance of individual species was significantly greater on Norway spruce than on blue spruce. With the current decline of Norway spruce as a result of several negative factors, blue spruce has emerged as a suitable substitute host plant for coccinellid communities in urban areas.</p>
</abstract>
<kwd-group>
<kwd>ladybirds</kwd>
<kwd>conifers</kwd>
<kwd>congeneric plants</kwd>
<kwd>urban ecosystems</kwd>
<kwd>
<italic>Picea abies</italic>
</kwd>
<kwd>
<italic>Picea pungens</italic>
</kwd>
</kwd-group>
<counts>
<fig-count count="2"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="56"/>
<page-count count="9"/>
<word-count count="3584"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Urban Ecology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>    <p>Norway spruce (<italic>Picea abies</italic> (L.) H. Karst) is a coniferous species with a wide distribution range across central Europe, not only in naturally colonised mountain areas but also in large areas where it has been intensively planted in the past two hundred years. Due to ongoing climate change, Norway spruce, a species highly sensitive to summer droughts and warm temperatures, is currently experiencing dieback (<xref ref-type="bibr" rid="B22">Kol&#xe1;&#x159; et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B4">Boczo&#x144; et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B23">Krejza et&#xa0;al., 2021</xref>) and frequent pest outbreaks (i.e., bark beetles), leading to the periodic decline of large, forested areas (<xref ref-type="bibr" rid="B16">Jactel et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B35">Marini et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B38">Netherer et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B18">Kami&#x144;ska et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B36">Nardi et&#xa0;al., 2022</xref>). Stress factors such as air pollution, heat, water deficiency, and drought have even more intensive impacts on trees in urban environments (<xref ref-type="bibr" rid="B33">L&#xfc;ttge and Buckeridge, 2023</xref>), resulting in the disappearance of many Norway spruce trees from urban plantings (<xref ref-type="bibr" rid="B1">Anonymous, 2023</xref>).</p>
<p>With the dieback of Norway spruce stands, insect communities associated with this host plant, including communities of coccinellids (Coleoptera, Coccinellidae), have also become threatened. Coccinellids play a crucial role in limiting the abundance of phytophagous insects (<xref ref-type="bibr" rid="B12">Hodek, 1973</xref>; <xref ref-type="bibr" rid="B34">Majerus, 2016</xref>), and their absence puts host plants in urban stands at greater risk. The preservation of diverse coccinellid fauna in towns and cities has both ecological and cultural significance (<xref ref-type="bibr" rid="B49">Soares et&#xa0;al., 2022</xref>). Norway spruce is a coniferous tree in which coccinellid communities have rarely been studied (<xref ref-type="bibr" rid="B42">Reddersen and Jensen, 2002</xref>; <xref ref-type="bibr" rid="B46">Selyemov&#xe1; et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B20">Kenis et&#xa0;al., 2020</xref>). This tree species is reported as a host of diverse coccinellid communities (<xref ref-type="bibr" rid="B46">Selyemov&#xe1; et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B20">Kenis et&#xa0;al., 2020</xref>). In central Europe, the coccinellid communities on Norway spruce include species similar to those of communities on other trees, mainly the dominant native <italic>Coccinella septempunctata</italic> Linnaeus, 1758 and invasive <italic>Harmonia axyridis</italic> (Pallas, 1773) (<xref ref-type="bibr" rid="B51">Stathas et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B32">Latibari et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B19">Kempkens and Gruppe, 2018</xref>; <xref ref-type="bibr" rid="B20">Kenis et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B56">Zach et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B7">Farrow et&#xa0;al., 2022</xref>), as well as coccinellids typically occurring on conifers such as <italic>Aphidecta obliterata</italic> (Linnaeus, 1758) (<xref ref-type="bibr" rid="B6">Burmeister et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B53">Timms et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B2">Banfield-Zanin and Leather, 2016</xref>) and <italic>Exochomus quadripustulatus</italic> (Linnaeus, 1758) (<xref ref-type="bibr" rid="B51">Stathas et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B14">Holecov&#xe1; et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B37">Nedv&#x11b;d, 2020</xref>).</p>
<p>Therefore, in our study, we aimed to determine whether there are suitable substitute conifer species capable of replacing the role of the disappearing Norway spruce in the survival of these coccinellid communities. A suitable substitute species for Norway spruce, at least in urban habitats, may be the introduced blue spruce (<italic>Picea pungens</italic> Engelm). This species, which is native to North America (<xref ref-type="bibr" rid="B8">Fechner, 1990</xref>), can withstand drought better than other spruce species and is resistant to high insolation (<xref ref-type="bibr" rid="B8">Fechner, 1990</xref>). Since blue spruce is also resilient to high levels of air pollution (<xref ref-type="bibr" rid="B41">Qin et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B5">Bukharina et&#xa0;al., 2016</xref>), after its introduction to Europe, it was planted as a replacement for Norway spruce stands damaged by industrial emissions (<xref ref-type="bibr" rid="B48">Slodi&#x10d;&#xe1;k and Nov&#xe1;k, 2008</xref>; <xref ref-type="bibr" rid="B26">Kula et&#xa0;al., 2016</xref>). Due to its aesthetic qualities, blue spruce can replace Norway spruce in park stands. This tree species has been widely planted in urban parks for approximately the past 160 years (<xref ref-type="bibr" rid="B39">Pagan and Randu&#x161;ka, 1988</xref>; <xref ref-type="bibr" rid="B31">Kulfan et&#xa0;al., 2010</xref>), and its entomofauna can therefore be studied in most cities in central Europe. Blue spruce is known to be a host for numerous insect groups in Europe: xylophagous beetles (Coleoptera) (<xref ref-type="bibr" rid="B24">Kr&#x161;iak et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B28">Kula et&#xa0;al., 2012</xref>, <xref ref-type="bibr" rid="B27">2013</xref>), ladybirds (Coleoptera) (<xref ref-type="bibr" rid="B17">Jauschov&#xe1; et&#xa0;al., 2024</xref>), moths (Lepidoptera) (<xref ref-type="bibr" rid="B31">Kulfan et&#xa0;al., 2010</xref>), sawflies (Hymenoptera, Symphyta) (<xref ref-type="bibr" rid="B26">Kula et&#xa0;al., 2016</xref>) and aphids (Homoptera) (<xref ref-type="bibr" rid="B9">Fry&#x10d;, 2016</xref>; <xref ref-type="bibr" rid="B55">Wojciechowski et&#xa0;al., 2016</xref>).</p>
<p>Can blue spruce function as a host for insect communities hosted by the native Norway spruce? To answer this question, we investigated and compared the diversity of coccinellid communities on Norway spruce and blue spruce grown together in urban areas. Given their close congeneric relationships, we expected similar coccinellid communities on both spruce species in urban environments. For our study, over a two-year period, we sampled adult coccinellid communities (including Scymninae and Coccidulinae subfamilies) monthly on both spruce species in four Slovak towns representing diverse climatic regions. We then evaluated the qualitative and quantitative differences in the adult coccinellid communities occurring concurrently during the growing season on both spruce species.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Study sites</title>
<p>Coccinellids were sampled from urban plantings (parks, alleys, cemeteries, housing estates, etc.) in four towns in Slovakia, where Norway spruce and blue spruce are planted together in sufficient numbers: &#x17d;ilina (49.19805 N, 18.73806E, 363 m a.s.l.), Slia&#x10d; (48.61125 N, 19.15655E, 302 m a.s.l.), Zvolen (48.57933 N, 19.14886E, 321 m a.s.l.) and Levice (48.21743 N, 18.60141E, 160 m a.s.l.). The towns are located in different geographical regions: northern (&#x17d;ilina), central (Slia&#x10d;, Zvolen) and southern (Levice) (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). At the sites, other deciduous trees (<italic>Tilia</italic> spp., <italic>Acer</italic> spp., <italic>Quercus</italic> spp.), conifers (<italic>Pinus</italic> spp., <italic>Pseudotsuga menziesii</italic>, <italic>Abies</italic> spp., <italic>Taxus</italic> spp.) and shrubs (<italic>Forsythia</italic> &#xd7; <italic>intermedia</italic>, <italic>Juniperus</italic> spp., <italic>Sambucus nigra</italic>, <italic>Ligustrum</italic> spp.) occurred widely. The ground surface was covered with mown lawn and ornamental plants and, to a lesser extent, with asphalt and concrete.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Locations of towns where sampling was carried out (&#x2022;). Source: Adapted from <uri xlink:href="https://commons.wikimedia.org/wiki/File:Slovensko_hranice.svg">https://commons.wikimedia.org/wiki/File:Slovensko_hranice.svg</uri> (public domain).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-12-1352625-g001.tif"/>
</fig>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Sampling of coccinellid communities</title>
<p>At each location, sampling was carried out at monthly intervals from April to November (8 total sampling sessions per year) in two consecutive years, 2021 and 2022 (16 total sampling sessions for the entire research period at each site). In each month, beetles from all locations were collected within 5-day intervals. Coccinellids were beaten from lower branches (1&#x2013;3 m above the ground) with wooden sticks into a circular tray with a diameter of 1 m (<xref ref-type="bibr" rid="B44">Roy et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B45">Roy and Brown, 2018</xref>). All sampled trees were taller than 5 m and older than 15 years. At each sampling session, ten different Norway spruce trees and ten different blue spruce trees were sampled for ladybirds. A total of 10 branches were beaten per sampled tree. The samples were collected around the entire circumference of the tree crown. The maximal distance among the sampled trees was 100 m. Adult coccinellids were identified at the species level according to <xref ref-type="bibr" rid="B37">Nedv&#x11b;d (2020)</xref>. Coccinellid species were classified as &#x201c;conifer specialists&#x201d; according to their preference for coniferous trees following <xref ref-type="bibr" rid="B21">Koch (1989)</xref>, <xref ref-type="bibr" rid="B14">Holecov&#xe1; et&#xa0;al. (2018)</xref> and <xref ref-type="bibr" rid="B37">Nedv&#x11b;d (2020)</xref>. The representation of &#x201c;conifer specialists&#x201d; was calculated as the relative abundance of &#x201c;conifer specialists&#x201d; among all ladybirds.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Statistical analysis</title>
<p>Differences in ladybird communities on Norway spruce and blue spruce were examined separately at individual sites. Qualitative differences were determined by the strength of the correlation of the relative abundance of individual species of coccinellids (the quantity of individuals of a species expressed as a proportion of the total number of coccinellid individuals) on Norway spruce and blue spruce. The calculations were performed for (i) all species for the entire research period at individual sites, (ii) species of the tribe Coccinellini (aphidophagous species), (iii) species belonging to all other tribes (non-Coccinellini, species readily accepting food other than aphids) and (iv) the 7 most abundant species [<italic>Exochomus quadripustulatus</italic>, <italic>Aphidecta obliterata</italic>, <italic>Harmonia quadripunctata</italic> (Pontoppidan, 1763), <italic>Rhyzobius chrysomeloides</italic> (Herbst, 1792), <italic>Adalia conglomerata</italic> (Linnaeus, 1758), <italic>Coccinella septempunctata</italic>, <italic>Harmonia axyridis</italic>]. Quantitative differences were investigated in a randomised block pattern without replicates, where samples collected at one location and date formed blocks and the spruce species represented the treatment. If the distribution of the experimental data passed the normality test, we used the pairwise multiple comparison procedure with the Holm-Sidak method. If the distribution of the experimental data failed the test of normality or equal variances, we used the Friedman repeated measures analysis of variance on ranks. This method was used because the abundance of ladybirds changed significantly during the growing season, and we were interested in the overall difference in the abundance of coccinellids on Norway spruce and blue spruce. All analyses were performed using SigmaStat 3.5 (<xref ref-type="bibr" rid="B52">Systat Software Inc, 2006</xref>).</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Qualitative structure of the adult coccinellid communities</title>
<p>The community structure of coccinellids on both spruce species in all four towns was similar. The correlation of the relative abundance of coccinellid species on Norway spruce and blue spruce was high at all locations: Levice (R<sup>2</sup> = 0.991; P &#x2264; 0.001), Slia&#x10d; (R<sup>2</sup> = 0.920; P &#x2264; 0.001), &#x17d;ilina (R<sup>2</sup> = 0.903; P &#x2264; 0.001) and Zvolen (R<sup>2</sup> = 0.975; P &#x2264; 0.001). The correlation of the relative abundance of species across all sampling sites was high (R<sup>2</sup> = 0.986; P &#x2264; 0.001), which further indicated that both species host the same coccinellid communities (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Frequency of coccinellid species on blue spruce plotted against the frequency of coccinellid species on Norway spruce. Data for the total sample of coccinellids from the Levice, Slia&#x10d;, &#x17d;ilina and Zvolen locations (average values from frequencies at individual locations). The black line indicates the same species frequency on Norway spruce and blue spruce.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-12-1352625-g002.tif"/>
</fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Quantitative differences in the community composition of adult coccinellids</title>
<p>In contrast to the species composition of the coccinellid community, we found significant differences in the abundance of coccinellids on Norway spruce and blue spruce (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>, Appendix 1). At each site, there was a significantly greater abundance of ladybirds on Norway spruce than on blue spruce.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Comparison of the coccinellid abundance on Norway spruce (<italic>Picea abies</italic>) and Blue spruce (<italic>Picea pungens</italic>) in locations in Slovakia.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="bottom" align="left" rowspan="2">Locality</th>
<th valign="bottom" align="left" rowspan="2">Species</th>
<th valign="bottom" colspan="4" align="left">
<italic>Picea abies</italic>
</th>
<th valign="bottom" colspan="4" align="left">
<italic>Picea pungens</italic>
</th>
<th valign="bottom" align="center">Difference</th>
</tr>
<tr>
<th valign="middle" align="center">Nsp</th>
<th valign="middle" align="center">Ni</th>
<th valign="middle" align="center">Mean &#xb1; SE</th>
<th valign="middle" align="center">Median</th>
<th valign="middle" align="center">Nsp</th>
<th valign="middle" align="center">Ni</th>
<th valign="middle" align="center">Mean &#xb1; SE</th>
<th valign="middle" align="center">Median</th>
<th valign="middle" align="center">P</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="bottom" align="left">Levice</td>
<td valign="bottom" align="left">Coccinellidae</td>
<td valign="middle" align="center">22</td>
<td valign="middle" align="center">2322</td>
<td valign="middle" align="center">145.1 &#xb1; 20.04</td>
<td valign="middle" align="center">141.5</td>
<td valign="middle" align="center">17</td>
<td valign="middle" align="center">744</td>
<td valign="middle" align="center">46.5 &#xb1; 11.63</td>
<td valign="middle" align="center">37</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">Coccinellini</td>
<td valign="middle" align="center">12</td>
<td valign="middle" align="center">836</td>
<td valign="middle" align="center">52.3 &#xb1; 13.40</td>
<td valign="middle" align="center">33.5</td>
<td valign="middle" align="center">9</td>
<td valign="middle" align="center">212</td>
<td valign="middle" align="center">13.3 &#xb1; 3.42</td>
<td valign="middle" align="center">9.5</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">non-Coccinellini</td>
<td valign="middle" align="center">10</td>
<td valign="middle" align="center">1486</td>
<td valign="middle" align="center">92.9 &#xb1; 12.53</td>
<td valign="middle" align="center">98.5</td>
<td valign="middle" align="center">8</td>
<td valign="middle" align="center">532</td>
<td valign="middle" align="center">33.3 &#xb1; 9.98</td>
<td valign="middle" align="center">23</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Aphidecta obliterata</italic> (Linnaeus, 1758)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">209</td>
<td valign="middle" align="center">13.1 &#xb1; 5.43</td>
<td valign="middle" align="center">5.5</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">62</td>
<td valign="middle" align="center">3.9 &#xb1; 2.43</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0.004</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Coccinella septempunctata</italic> (Linnaeus, 1758)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">202</td>
<td valign="middle" align="center">12.6 &#xb1; 3.93</td>
<td valign="middle" align="center">4</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">61</td>
<td valign="middle" align="center">3.8 &#xb1; 1.19</td>
<td valign="middle" align="center">2.5</td>
<td valign="middle" align="center">0.077</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Exochomus quadripustulatus</italic> Linnaeus, 1758</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">1361</td>
<td valign="middle" align="center">85.1 &#xb1; 12.70</td>
<td valign="middle" align="center">76.5</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">470</td>
<td valign="middle" align="center">29.4 &#xb1; 9.56</td>
<td valign="middle" align="center">20.5</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Harmonia axyridis</italic> (Pallas, 1773)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">143</td>
<td valign="middle" align="center">8.9 &#xb1; 7.22</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">26</td>
<td valign="middle" align="center">1.6 &#xb1; 0.63</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">0.077</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Harmonia quadripunctata</italic> (Pontoppidan, 1763)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">217</td>
<td valign="middle" align="center">13.6 &#xb1; 3.67</td>
<td valign="middle" align="center">9.5</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">49</td>
<td valign="middle" align="center">3.1 &#xb1; 0.78</td>
<td valign="middle" align="center">2.5</td>
<td valign="middle" align="center">0.021</td>
</tr>
<tr>
<td valign="bottom" align="left">Slia&#x10d;</td>
<td valign="bottom" align="left">Coccinellidae</td>
<td valign="middle" align="center">28</td>
<td valign="middle" align="center">1443</td>
<td valign="middle" align="center">90.2 &#xb1; 24.63</td>
<td valign="middle" align="center">51</td>
<td valign="middle" align="center">22</td>
<td valign="middle" align="center">533</td>
<td valign="middle" align="center">33.3 &#xb1; 10.50</td>
<td valign="middle" align="center">22</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">Coccinellini</td>
<td valign="middle" align="center">17</td>
<td valign="middle" align="center">1066</td>
<td valign="middle" align="center">66.6 &#xb1; 24.02</td>
<td valign="middle" align="center">29</td>
<td valign="middle" align="center">14</td>
<td valign="middle" align="center">382</td>
<td valign="middle" align="center">23.9 &#xb1; 10.52</td>
<td valign="middle" align="center">10.5</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">non-Coccinellini</td>
<td valign="middle" align="center">11</td>
<td valign="middle" align="center">379</td>
<td valign="middle" align="center">23.7 &#xb1; 5.43</td>
<td valign="middle" align="center">13</td>
<td valign="middle" align="center">8</td>
<td valign="middle" align="center">151</td>
<td valign="middle" align="center">9.4 &#xb1; 1.94</td>
<td valign="middle" align="center">6.5</td>
<td valign="middle" align="center">0.004</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Adalia conglomerata</italic> (Linnaeus, 1758)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">264</td>
<td valign="middle" align="center">16.5 &#xb1; 4.97</td>
<td valign="middle" align="center">10</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">64</td>
<td valign="middle" align="center">4.0 &#xb1; 1.71</td>
<td valign="middle" align="center">1.5</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Aphidecta obliterata</italic> (Linnaeus, 1758)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">587</td>
<td valign="middle" align="center">36.7 &#xb1; 16.07</td>
<td valign="middle" align="center">11.5</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">192</td>
<td valign="middle" align="center">12.0 &#xb1; 5.95</td>
<td valign="middle" align="center">2</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Exochomus quadripustulatus</italic> Linnaeus, 1758</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">244</td>
<td valign="middle" align="center">15.3 &#xb1; 3.32</td>
<td valign="middle" align="center">12.5</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">103</td>
<td valign="middle" align="center">6.4 &#xb1; 1.25</td>
<td valign="middle" align="center">5</td>
<td valign="middle" align="center">0.007</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="middle" align="left">
<italic>Rhyzobius chrysomeloides</italic> (Herbst, 1792)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">100</td>
<td valign="middle" align="center">6.3 &#xb1; 2.13</td>
<td valign="middle" align="center">3</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">12</td>
<td valign="middle" align="center">0.8 &#xb1; 0.35</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0.021</td>
</tr>
<tr>
<td valign="bottom" align="left">&#x17d;ilina</td>
<td valign="bottom" align="left">Coccinellidae</td>
<td valign="middle" align="center">15</td>
<td valign="middle" align="center">1445</td>
<td valign="middle" align="center">90.3 &#xb1; 29.43</td>
<td valign="middle" align="center">53.5</td>
<td valign="middle" align="center">15</td>
<td valign="middle" align="center">579</td>
<td valign="middle" align="center">36.2 &#xb1; 6.09</td>
<td valign="middle" align="center">36</td>
<td valign="middle" align="center">0.005</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">Coccinellini</td>
<td valign="middle" align="center">9</td>
<td valign="middle" align="center">962</td>
<td valign="middle" align="center">60.1 &#xb1; 26.15</td>
<td valign="middle" align="center">35.5</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center">362</td>
<td valign="middle" align="center">22.6 &#xb1; 4.68</td>
<td valign="middle" align="center">20</td>
<td valign="middle" align="center">0.021</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">non-Coccinellini</td>
<td valign="middle" align="center">6</td>
<td valign="middle" align="center">483</td>
<td valign="middle" align="center">30.2 &#xb1; 7.20</td>
<td valign="middle" align="center">23.5</td>
<td valign="middle" align="center">8</td>
<td valign="middle" align="center">217</td>
<td valign="middle" align="center">13.7 &#xb1; 2.42</td>
<td valign="middle" align="center">13.5</td>
<td valign="middle" align="center">0.077</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Aphidecta obliterata</italic> (Linnaeus, 1758)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">824</td>
<td valign="middle" align="center">51.5 &#xb1; 24.89</td>
<td valign="middle" align="center">25</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">302</td>
<td valign="middle" align="center">18.9 &#xb1; 4.29</td>
<td valign="middle" align="center">14</td>
<td valign="middle" align="center">0.077</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Exochomus quadripustulatus</italic> Linnaeus, 1758</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">141</td>
<td valign="middle" align="center">8.8 &#xb1; 2.41</td>
<td valign="middle" align="center">6</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">68</td>
<td valign="middle" align="center">4.3 &#xb1; 0.86</td>
<td valign="middle" align="center">3.5</td>
<td valign="middle" align="center">0.454</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="middle" align="left">
<italic>Rhyzobius chrysomeloides</italic> (Herbst, 1792)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">133</td>
<td valign="middle" align="center">8.3 &#xb1; 7.55</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">125</td>
<td valign="middle" align="center">7.8 &#xb1; 6.91</td>
<td valign="middle" align="center">6</td>
<td valign="middle" align="center">0.757</td>
</tr>
<tr>
<td valign="bottom" align="left">Zvolen</td>
<td valign="bottom" align="left">Coccinellidae</td>
<td valign="middle" align="center">21</td>
<td valign="middle" align="center">1632</td>
<td valign="middle" align="center">102.0 &#xb1; 23.86</td>
<td valign="middle" align="center">49</td>
<td valign="middle" align="center">17</td>
<td valign="middle" align="center">513</td>
<td valign="middle" align="center">32.1 &#xb1; 5.23</td>
<td valign="middle" align="center">25</td>
<td valign="middle" align="center">0.004</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">Coccinellini</td>
<td valign="middle" align="center">11</td>
<td valign="middle" align="center">1065</td>
<td valign="middle" align="center">66.6 &#xb1; 18.80</td>
<td valign="middle" align="center">27.5</td>
<td valign="middle" align="center">10</td>
<td valign="middle" align="center">324</td>
<td valign="middle" align="center">20.3 &#xb1; 3.74</td>
<td valign="middle" align="center">13</td>
<td valign="middle" align="center">0.004</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">non-Coccinellini</td>
<td valign="middle" align="center">10</td>
<td valign="middle" align="center">567</td>
<td valign="middle" align="center">35.4 &#xb1; 7.79</td>
<td valign="middle" align="center">27</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center">189</td>
<td valign="middle" align="center">11.8 &#xb1; 2.59</td>
<td valign="middle" align="center">10</td>
<td valign="middle" align="center">0.004</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Adalia conglomerata</italic> (Linnaeus, 1758)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">142</td>
<td valign="middle" align="center">8.9 &#xb1; 2.42</td>
<td valign="middle" align="center">6</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">16</td>
<td valign="middle" align="center">1.0 &#xb1; 0.41</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">&#x2264;0.001</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Aphidecta obliterata</italic> (Linnaeus, 1758)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">714</td>
<td valign="middle" align="center">44.6 &#xb1; 14.73</td>
<td valign="middle" align="center">15.5</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">226</td>
<td valign="middle" align="center">14.1 &#xb1; 3.64</td>
<td valign="middle" align="center">9</td>
<td valign="middle" align="center">0.004</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Exochomus quadripustulatus</italic> Linnaeus, 1758</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">388</td>
<td valign="middle" align="center">24.3 &#xb1; 5.52</td>
<td valign="middle" align="center">16</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">108</td>
<td valign="middle" align="center">6.8 &#xb1; 1.76</td>
<td valign="middle" align="center">5.5</td>
<td valign="middle" align="center">0.002</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="bottom" align="left">
<italic>Harmonia quadripunctata</italic> (Pontoppidan, 1763)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">124</td>
<td valign="middle" align="center">7.8 &#xb1; 3.82</td>
<td valign="middle" align="center">1.5</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">34</td>
<td valign="middle" align="center">2.1 &#xb1; 0.72</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">0.210</td>
</tr>
<tr>
<td valign="bottom" align="left"/>
<td valign="middle" align="left">
<italic>Rhyzobius chrysomeloides</italic> (Herbst, 1792)</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">124</td>
<td valign="middle" align="center">7.8 &#xb1; 2.46</td>
<td valign="middle" align="center">4</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">47</td>
<td valign="middle" align="center">2.9 &#xb1; 0.78</td>
<td valign="middle" align="center">2</td>
<td valign="middle" align="center">0.21</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Coccinellidae &#x2013; total number of individuals of all species; Coccinellini &#x2013; a total number of individuals of the tribe Coccinellini; non-Coccinellini &#x2013; total number of individuals of other tribes, and total number of individuals of the most abundant species at localities.</p>
</fn>
<fn>
<p>Nsp, number of species; Ni, number of individuals; Mean &#xb1; SE, average number of individuals in 16 samples obtained during the study period &#xb1; standard error; Median, median number of individuals in 16 samples obtained during the study period; P, probability of difference between the number of individuals on <italic>P. abies</italic> and <italic>P. pungens</italic>.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>The abundance of coccinellids was significantly greater on Norway spruce at the whole-community level, as was that of the coccinellids of the tribe Coccinellini and of other tribes, with the exception of that at the &#x17d;ilina location. Some of the seven most abundant species (<italic>E. quadripustulatus</italic>, <italic>A. obliterata</italic>, <italic>H. quadripunctata</italic>, <italic>R. chrysomeloides</italic>, <italic>A. conglomerata</italic>, <italic>C. septempunctata</italic>, and <italic>H. axyridis</italic>) were significantly more abundant on Norway spruce than on blue spruce at individual locations (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). The total abundance of the coccinellid community (both years and all locations) was 2.88-fold greater on Norway spruce than on blue spruce.</p>
<p>Additionally, the total number of species found at individual sites on Norway spruce (mean 21.5 &#xb1; 2.66 species) was greater than that found on blue spruce (mean 17.8 &#xb1; 1.49 species). However, this difference was only marginally significant (F=8.133, P=0.065) due to the small number of locations sampled. For the whole research period at all locations, a total of 34 coccinellid species were recorded on Norway spruce, and 27 species were recorded on blue spruce. Among them, <italic>Exochomus quadripustulatus</italic> and <italic>Aphidecta obliterata</italic> were eudominant (&gt; 10%) on both spruce species. The invasive harlequin ladybird (<italic>Harmonia axyridis</italic>) was slightly more dominant on blue spruce (5.45%) than on Norway spruce (3.86%). Conifer specialist coccinellids strongly dominated communities on both spruce species [Norway spruce 87.66% (10 species), blue spruce 84.76% (9 species)].</p>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>The absence of qualitative differences in coccinellid communities between Norway spruce and blue spruce was mainly determined by the similar relative abundance of dominant species and the low relative abundance of rare species.</p>
<p>As a result, the relative abundances of coccinellid species on Norway spruce and blue spruce were similar across locations, although the dominant species across locations differed (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>), with the exception of <italic>Aphidecta obliterata</italic> and <italic>Exochomus quadripustulatus</italic>, which were abundant across all locations. Since the relative abundance of rare species was very low, it did not affect the correlation between the relative abundance of rare coccinellid species and that of either spruce species.</p>
<p>The species of spruce did not affect the relative abundance of coccinellid species in the total population that were established on the species. Additionally, Norway spruce and blue spruce have different species-specific properties that affect the attraction or repulsion of particular coccinellid species.</p>
<p>The invasive eurytopic coccinellid <italic>Harmonia axyridis</italic> (<xref ref-type="bibr" rid="B43">Roy et&#xa0;al., 2016</xref>) is known to dominate coccinellid communities on broadleaved trees (<xref ref-type="bibr" rid="B15">Honek et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B54">Vigl&#xe1;&#x161;ov&#xe1; et&#xa0;al., 2017</xref>). This ladybird species occurs on Norway spruce as well but usually in lower abundances (<xref ref-type="bibr" rid="B20">Kenis et&#xa0;al., 2020</xref>), which is in accordance with our results. Among conifers, <italic>Harmonia axyridis</italic> seems to prefer <italic>Scots pine</italic> (<italic>Pinus sylvestris</italic>) (<xref ref-type="bibr" rid="B14">Holecov&#xe1; et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B20">Kenis et&#xa0;al., 2020</xref>).</p>
<p>While no qualitative differences were found in the coccinellid communities on either spruce species, there were significant quantitative differences. All coccinellids (the community of all species), species of the tribe Coccinellini, species of other tribes (with the exception of the &#x17d;ilina location) and some of abundant species were significantly more abundant on Norway spruce than on blue spruce. We can speculate about the reasons for these differences in the abundance of coccinellids between both spruce species. Possible causes include trophic factors, such as differences in the abundance or species composition of the arthropod complex serving as the prey for coccinellids. The aphid communities on Norway spruce and blue spruce seem to consist of similar aphid species (see, for example, <xref ref-type="bibr" rid="B9">Fry&#x10d;, 2016</xref>; <xref ref-type="bibr" rid="B55">Wojciechowski et&#xa0;al., 2016</xref>). Trophic differences can be quantitative &#x2013; the abundance of aphids on blue spruce seems to be lower than that on Norway spruce (pers. observation TJ, PZ). Other groups of phytophagous insects, as possible alternative food sources for coccinellids (<xref ref-type="bibr" rid="B13">Hodek and Evans, 2012</xref>; <xref ref-type="bibr" rid="B34">Majerus, 2016</xref>), seem to occupy both spruce species similarly, such as the larvae of moths (<xref ref-type="bibr" rid="B31">Kulfan et&#xa0;al., 2010</xref>) and sawflies (<xref ref-type="bibr" rid="B26">Kula et&#xa0;al., 2016</xref>).</p>
<p>Another biotic difference is the quality of the host plant (<xref ref-type="bibr" rid="B10">Go&#xdf;ner and Ammer, 2006</xref>; <xref ref-type="bibr" rid="B3">Bezemer et&#xa0;al., 2014</xref>). Differences in plant architecture, structural heterogeneity and plant surface, e.g., foliage shape and density, may aid or hinder coccinellid movement and searching and foraging behaviour (<xref ref-type="bibr" rid="B47">Shah, 1982</xref>; <xref ref-type="bibr" rid="B11">Grevstad and Klepetka, 1992</xref>; <xref ref-type="bibr" rid="B40">Pervez and Yadav, 2018</xref>). Little is known about whether there are differences in needle chemicals between blue spruce and Norway spruce (<xref ref-type="bibr" rid="B50">Soukupov&#xe1; et&#xa0;al., 2001</xref>). A comparison of the needles of Norway spruce and blue spruce with regard to their resistance to emissions revealed similar concentrations of chlorophyll and carotenoids but showed differences in phenolic compound accumulation between the two spruce species (<xref ref-type="bibr" rid="B50">Soukupov&#xe1; et&#xa0;al., 2001</xref>). The effects of chemical differences in the needles of both spruce species on coccinellids and their prey are not known.</p>
<p>How would coccinellid communities change if Norway spruce trees completely disappeared? For example, we can consider the replacement of the spruce by deciduous trees in the D&#x11b;&#x10d;&#xed;nsk&#xfd; Sn&#x11b;&#x17e;n&#xed;k locality (Czech Republic, 50.79 N, 14.11E, 700 m a.s.l.), which has resulted in a shift in the composition of insect communities (<xref ref-type="bibr" rid="B29">Kula and Tryner, 2003a</xref>, <xref ref-type="bibr" rid="B30">b</xref>), including those of aphidophagous syrphids (Diptera) (<xref ref-type="bibr" rid="B25">Kula, 1997</xref>). A similar shift in the species composition of the local coccinellid fauna could also occur in the urban habitats in which this study was conducted.</p>
<p>Unlike the native Norway spruce, blue spruce is expected to be able to withstand unfavourable environmental conditions (<xref ref-type="bibr" rid="B8">Fechner, 1990</xref>; <xref ref-type="bibr" rid="B41">Qin et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B5">Bukharina et&#xa0;al., 2016</xref>). Our results suggest that in the absence of Norway spruce in urban environments, the coccinellid community inhabiting congeneric blue spruce will likely survive with no changes in composition, i.e., a decrease in species richness will not occur. In our study, the abundance of most coccinellid species was greater on Norway spruce than on blue spruce when coccinellids had the option to select Norway spruce and selection did not require migration to distant habitats. When such an option is not available, an increase in the abundance of coccinellids on blue spruce can be expected since the community of coccinellids adapted to conifers may find refuge there. However, confirmation of this assumption requires a more detailed study.</p>
<p>In conclusion, both native Norway spruce and introduced blue spruce host similar coccinellid communities in urban areas. Blue spruce was introduced to Slovakia more than 150 years ago (<xref ref-type="bibr" rid="B39">Pagan and Randu&#x161;ka, 1988</xref>), and in the time period since its introduction, it has recruited a coccinellid community with a species composition similar to that of the native Norway spruce. Closely related nonnative blue spruce seems to be a useful substitute host for spruce-associated insect fauna in the case of Norway spruce extinction in urban areas. A long-term study of coccinellid communities on spruces would be beneficial since the abundance of this insect is known to fluctuate interannually (this study; <xref ref-type="bibr" rid="B46">Selyemov&#xe1; et&#xa0;al., 2007</xref>).</p>
</sec>
<sec id="s5" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/supplementary material. Further inquiries can be directed to the corresponding author.</p>
</sec>
<sec id="s6" sec-type="ethics-statement">
<title>Ethics statement</title>
<p>The manuscript presents research on animals that do not require ethical approval for their study.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>TJ: Conceptualization, Data curation, Investigation, Methodology, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. LS: Conceptualization, Project administration, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. PZ: Conceptualization, Investigation, Methodology, Supervision, Writing &#x2013; original draft. MS: Funding acquisition, Investigation, Project administration, Writing &#x2013; review &amp; editing. ZM: Funding acquisition, Methodology, Project administration, Writing &#x2013; review &amp; editing. AH: Software, Supervision, Validation, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. JS: Formal analysis, Methodology, Software, Writing &#x2013; review &amp; editing. MH: Funding acquisition, Investigation, Project administration, Writing &#x2013; review &amp; editing. JK: Investigation, Methodology, Supervision, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing.</p>
</sec>
</body>
<back>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. This study was supported by a grant VEGA 2/0022/23 from the Scientific Grant Agency of Ministry of Education, Science, Research and Sport of the Slovak Republic (TJ, LS, JK, MS, MH), a grant for PhD students DoktoGrant APP0404 from the Slovak Academy of Sciences (TJ), a grant BAS-SAS-2022-02 (TJ, JK) and the Ministry of Agriculture of the Czech Republic, institutional support MZE-RO0423 (AH, ZM, and JS).</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We thank Milan Miku&#x161; for technical assistance with field research.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
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<app-group>
<app id="app1">
<title>Appendix 1</title>
<p>Total numbers of individuals of coccinellid species recorded at all locations in this study on Norway spruce (<italic>Picea abies</italic>).</p>
<p>
<italic>Adalia bipunctata</italic> (Linnaeus) 1; <italic>Adalia conglomerata</italic> (Linnaeus) 426; <italic>Adalia decempunctata</italic> (Linnaeus) 12; <italic>Anatis ocellata</italic> (Linnaeus) 9; <italic>Aphidecta obliterata</italic> (Linnaeus) 2334; <italic>Calvia decemguttata</italic> (Linnaeus) 3; <italic>Calvia quatuordecimguttata</italic> (Linnaeus) 2; <italic>Chilocorus bipustulatus</italic> Linnaeus 9; <italic>Clitostethus arcuatus</italic> (Rossi) 1; <italic>Coccinula quatuordecimpustulata</italic> (Linnaeus) 1; <italic>Coccinella quinquepunctata</italic> Linnaeus 1; <italic>Coccinella septempunctata</italic> Linnaeus 343; <italic>Exochomus quadripustulatus</italic> Linnaeus 2134; <italic>Halyzia sedecimguttata</italic> (Linnaeus) 1; <italic>Harmonia axyridis</italic> (Pallas) 264; <italic>Harmonia quadripunctata</italic> (Pontoppidan) 442; <italic>Hippodamia variegata</italic> (Goeze) 3; <italic>Myrrha octodecimguttata</italic> (Linnaeus) 2; <italic>Myzia oblongoguttata</italic> (Linnaeus) 24; <italic>Nephus quadrimaculatus</italic> (Herbst) 5; <italic>Oenopia conglobata</italic> (Linnaeus) 3; <italic>Platynaspis luteorubra</italic> (Goeze) 4; <italic>Propylea quatuordecimpunctata</italic> (Linnaeus) 11; <italic>Psyllobora vigintiduopunctata</italic> (Linnaeus) 3; <italic>Rhyzobius chrysomeloides</italic> (Herbst) 362; <italic>Scymnus abietis</italic> (Paykull) 258; <italic>Scymnus ferrugatus</italic> (Moll) 6; <italic>Scymnus frontalis</italic> (Fabricius) 11; <italic>Scymnus interruptus</italic> (Goeze) 24; <italic>Scymnus rubromaculatus</italic> (Goeze) 16; <italic>Scymnus subvillosus</italic> (Goeze) 1; <italic>Scymnus suturalis</italic> Thunberg 7; <italic>Stethorus pusillus</italic> (Herbst) 77; <italic>Vibidia duodecimguttata</italic> (Poda) 42.</p>
<p>Total numbers of individuals of coccinellid species recorded at all locations in this study on blue spruce (<italic>Picea pungens</italic>).</p>
<p>
<italic>Adalia conglomerata</italic> (Linnaeus) 87; <italic>Adalia decempunctata</italic> (Linnaeus) 11; <italic>Anatis ocellata</italic> (Linnaeus) 4; <italic>Aphidecta obliterata</italic> (Linnaeus) 782; <italic>Calvia quatuordecimguttata</italic> (Linnaeus) 2; <italic>Chilocorus bipustulatus</italic> Linnaeus 6; <italic>Coccinella septempunctata</italic> Linnaeus 113; <italic>Exochomus quadripustulatus</italic> Linnaeus 749; <italic>Harmonia axyridis</italic> (Pallas) 129; <italic>Harmonia quadripunctata</italic> (Pontoppidan) 120; <italic>Hippodamia variegata</italic> (Goeze) 1; <italic>Myrrha octodecimguttata</italic> (Linnaeus) 1; <italic>Myzia oblongoguttata</italic> (Linnaeus) 3; <italic>Oenopia conglobata</italic> (Linnaeus) 6; <italic>Platynaspis luteorubra</italic> (Goeze) 1; <italic>Propylea quatuordecimpunctata</italic> (Linnaeus) 6; <italic>Psyllobora vigintiduopunctata</italic> (Linnaeus) 3; <italic>Rhyzobius chrysomeloides</italic> (Herbst) 184; <italic>Scymnus abietis</italic> (Paykull) 78; <italic>Scymnus auritus</italic> Thunberg 1; <italic>Scymnus ferrugatus</italic> (Moll) 2; <italic>Scymnus frontalis</italic> (Fabricius) 5; <italic>Scymnus haemorrhoidalis</italic> Herbst 1; <italic>Scymnus interruptus</italic> (Goeze) 11; <italic>Scymnus rubromaculatus</italic> (Goeze) 8; <italic>Stethorus pusillus</italic> (Herbst) 43; <italic>Vibidia duodecimguttata</italic> (Poda) 12.</p>
</app>
</app-group>
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</article>