Links to Exploration step
Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Nestedness of habitat specialists within habitat generalists in a butterfly assemblage</title><author><name sortKey="Trivellini, Guido" sort="Trivellini, Guido" uniqKey="Trivellini G" first="Guido" last="Trivellini">Guido Trivellini</name><affiliation><nlm:aff id="icad12193-aff-0001"></nlm:aff></affiliation><affiliation><nlm:aff id="icad12193-aff-0002"></nlm:aff></affiliation></author><author><name sortKey="Polidori, Carlo" sort="Polidori, Carlo" uniqKey="Polidori C" first="Carlo" last="Polidori">Carlo Polidori</name><affiliation><nlm:aff id="icad12193-aff-0003"></nlm:aff></affiliation></author><author><name sortKey="Pasquaretta, Cristian" sort="Pasquaretta, Cristian" uniqKey="Pasquaretta C" first="Cristian" last="Pasquaretta">Cristian Pasquaretta</name><affiliation><nlm:aff id="icad12193-aff-0004"></nlm:aff></affiliation></author><author><name sortKey="Orsenigo, Simone" sort="Orsenigo, Simone" uniqKey="Orsenigo S" first="Simone" last="Orsenigo">Simone Orsenigo</name><affiliation><nlm:aff id="icad12193-aff-0005"></nlm:aff></affiliation></author><author><name sortKey="Bogliani, Giuseppe" sort="Bogliani, Giuseppe" uniqKey="Bogliani G" first="Giuseppe" last="Bogliani">Giuseppe Bogliani</name><affiliation><nlm:aff id="icad12193-aff-0001"></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">32336986</idno><idno type="pmc">7165506</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7165506</idno><idno type="RBID">PMC:7165506</idno><idno type="doi">10.1111/icad.12193</idno><date when="2016">2016</date><idno type="wicri:Area/Pmc/Corpus">000677</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000677</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Nestedness of habitat specialists within habitat generalists in a butterfly assemblage</title><author><name sortKey="Trivellini, Guido" sort="Trivellini, Guido" uniqKey="Trivellini G" first="Guido" last="Trivellini">Guido Trivellini</name><affiliation><nlm:aff id="icad12193-aff-0001"></nlm:aff></affiliation><affiliation><nlm:aff id="icad12193-aff-0002"></nlm:aff></affiliation></author><author><name sortKey="Polidori, Carlo" sort="Polidori, Carlo" uniqKey="Polidori C" first="Carlo" last="Polidori">Carlo Polidori</name><affiliation><nlm:aff id="icad12193-aff-0003"></nlm:aff></affiliation></author><author><name sortKey="Pasquaretta, Cristian" sort="Pasquaretta, Cristian" uniqKey="Pasquaretta C" first="Cristian" last="Pasquaretta">Cristian Pasquaretta</name><affiliation><nlm:aff id="icad12193-aff-0004"></nlm:aff></affiliation></author><author><name sortKey="Orsenigo, Simone" sort="Orsenigo, Simone" uniqKey="Orsenigo S" first="Simone" last="Orsenigo">Simone Orsenigo</name><affiliation><nlm:aff id="icad12193-aff-0005"></nlm:aff></affiliation></author><author><name sortKey="Bogliani, Giuseppe" sort="Bogliani, Giuseppe" uniqKey="Bogliani G" first="Giuseppe" last="Bogliani">Giuseppe Bogliani</name><affiliation><nlm:aff id="icad12193-aff-0001"></nlm:aff></affiliation></author></analytic><series><title level="j">Insect Conservation and Diversity</title><idno type="ISSN">1752-458X</idno><idno type="eISSN">1752-4598</idno><imprint><date when="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><title>Abstract</title><p><list list-type="order" id="icad12193-list-0001"><list-item><p>The habitat requirements of a species are the resources, conditions and space required for survival and reproduction. The habitat requirements of butterflies have been well studied, but the extent to which individuals within a species and between species utilise and share the habitat is poorly known.</p></list-item><list-item><p>In a butterfly assemblage in northern Italy, we found that adults from 30 species avoid deciduous high‐density forests and their ecotones, and they were positively related to open areas and their ecotones. Besides these common features, five groups of species can be discriminated in relation to a gradient from open area to forest, and species within groups were not equally specialised, as observed from a bipartite network analysis. In particular, some species appeared to be specialised and others appeared to be generalist, suggesting a nested pattern of resource use, rather than a clustered pattern in which each species uses a different subset of habitat types.</p></list-item><list-item><p>The degree of variation in specialisation among species varied with the number of species falling in each group. Thus, an increased number of species, and thus possibly competition, is more likely to promote the co‐occurrence of generalist and specialised species (nested patterns) rather than an increased niche segregation among species.</p></list-item><list-item><p>Ascertaining how species overlap their habitat use at a local scale can be relevant for conservation purposes, because specialised populations are potentially more susceptible to network distortions.</p></list-item></list></p></div></front><back><div1 type="bibliography"><listBibl><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Lin, D Y" uniqKey="Lin D">D.Y. Lin</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Vane Right, R I" uniqKey="Vane Right R">R.I. Vane‐Wright</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Stamp, N E" uniqKey="Stamp N">N.E. Stamp</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Insect Conserv Divers</journal-id><journal-id journal-id-type="iso-abbrev">Insect Conserv Divers</journal-id><journal-id journal-id-type="doi">10.1111/(ISSN)1752-4598</journal-id><journal-id journal-id-type="publisher-id">ICAD</journal-id><journal-title-group><journal-title>Insect Conservation and Diversity</journal-title></journal-title-group><issn pub-type="ppub">1752-458X</issn><issn pub-type="epub">1752-4598</issn><publisher><publisher-name>John Wiley and Sons Inc.</publisher-name><publisher-loc>Hoboken</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">32336986</article-id><article-id pub-id-type="pmc">7165506</article-id><article-id pub-id-type="doi">10.1111/icad.12193</article-id><article-id pub-id-type="publisher-id">ICAD12193</article-id><article-categories><subj-group subj-group-type="overline"><subject>Original Article</subject></subj-group><subj-group subj-group-type="heading"><subject>Original Articles</subject></subj-group></article-categories><title-group><article-title>Nestedness of habitat specialists within habitat generalists in a butterfly assemblage</article-title><alt-title alt-title-type="left-running-head">Guido Trivellini <italic>et al</italic>.</alt-title></title-group><contrib-group><contrib id="icad12193-cr-0001" contrib-type="author"><name><surname>Trivellini</surname><given-names>Guido</given-names></name><xref ref-type="aff" rid="icad12193-aff-0001"><sup>1</sup></xref><xref ref-type="aff" rid="icad12193-aff-0002"><sup>2</sup></xref></contrib><contrib id="icad12193-cr-0002" contrib-type="author"><name><surname>Polidori</surname><given-names>Carlo</given-names></name><xref ref-type="aff" rid="icad12193-aff-0003"><sup>3</sup></xref></contrib><contrib id="icad12193-cr-0003" contrib-type="author"><name><surname>Pasquaretta</surname><given-names>Cristian</given-names></name><xref ref-type="aff" rid="icad12193-aff-0004"><sup>4</sup></xref></contrib><contrib id="icad12193-cr-0004" contrib-type="author"><name><surname>Orsenigo</surname><given-names>Simone</given-names></name><xref ref-type="aff" rid="icad12193-aff-0005"><sup>5</sup></xref></contrib><contrib id="icad12193-cr-0005" contrib-type="author" corresp="yes"><name><surname>Bogliani</surname><given-names>Giuseppe</given-names></name><xref ref-type="aff" rid="icad12193-aff-0001"><sup>1</sup></xref><address><email>giuseppe.bogliani@unipv.it</email></address></contrib></contrib-group><contrib-group><contrib id="icad12193-cr-0006" contrib-type="editor"><name><surname>Didham</surname><given-names>Raphael</given-names></name></contrib><contrib id="icad12193-cr-0007" contrib-type="editor"><name><surname>DeVries</surname><given-names>Phil</given-names></name></contrib></contrib-group><aff id="icad12193-aff-0001"><label><sup>1</sup></label><named-content content-type="organisation-division">Department of Earth and Environmental Sciences</named-content><institution>University of Pavia</institution><city>Pavia</city><country country="IT">Italy</country></aff><aff id="icad12193-aff-0002"><label><sup>2</sup></label><named-content content-type="organisation-division">Eliante</named-content><institution>Non‐Profit Organisation for the Environment</institution><city>Milano</city><country country="IT">Italy</country></aff><aff id="icad12193-aff-0003"><label><sup>3</sup></label><named-content content-type="organisation-division">Instituto de Ciencias Ambientales (ICAM)</named-content><institution>Universidad de Castilla‐La Mancha</institution><city>Toledo</city><country country="ES">Spain</country></aff><aff id="icad12193-aff-0004"><label><sup>4</sup></label><named-content content-type="organisation-division">Department of Ecology, Physiology and Ethology</named-content><institution>IPHC</institution><institution>CNRS‐UDS</institution><city>Strasbourg Cedex</city><country country="FR">France</country></aff><aff id="icad12193-aff-0005"><label><sup>5</sup></label><named-content content-type="organisation-division">Dipartimento di Scienze Agrarie e Ambientali – Produzione, Territorio, Agroenergia</named-content><institution>Università degli Studi di Milano</institution><city>Milano</city><country country="IT">Italy</country></aff><author-notes><corresp id="correspondenceTo"><label>*</label>Correspondence: Giuseppe Bogliani, Department of Earth and Environmental Sciences, University of Pavia, Via Adolfo Ferrata 9, 27100 Pavia, Italy. E‐mail: <email>giuseppe.bogliani@unipv.it</email></corresp></author-notes><pub-date pub-type="epub"><day>25</day><month>9</month><year>2016</year></pub-date><pub-date pub-type="ppub"><month>11</month><year>2016</year></pub-date><volume>9</volume><issue>6</issue><issue-id pub-id-type="doi">10.1111/icad.2016.9.issue-6</issue-id><fpage>495</fpage><lpage>505</lpage><history><date date-type="accepted"><day>26</day><month>7</month><year>2016</year></date></history><permissions><pmc-comment> Copyright © 2016 The Royal Entomological Society </pmc-comment>
<copyright-statement content-type="article-copyright">© 2016 The Royal Entomological Society</copyright-statement><license><license-p>This article is being made freely available through PubMed Central as part of the COVID-19 public health emergency response. It can be used for unrestricted research re-use and analysis in any form or by any means with acknowledgement of the original source, for the duration of the public health emergency.</license-p></license></permissions><self-uri content-type="pdf" xlink:href="file:ICAD-9-495.pdf"></self-uri><abstract id="icad12193-abs-0001"><title>Abstract</title><p><list list-type="order" id="icad12193-list-0001"><list-item><p>The habitat requirements of a species are the resources, conditions and space required for survival and reproduction. The habitat requirements of butterflies have been well studied, but the extent to which individuals within a species and between species utilise and share the habitat is poorly known.</p></list-item><list-item><p>In a butterfly assemblage in northern Italy, we found that adults from 30 species avoid deciduous high‐density forests and their ecotones, and they were positively related to open areas and their ecotones. Besides these common features, five groups of species can be discriminated in relation to a gradient from open area to forest, and species within groups were not equally specialised, as observed from a bipartite network analysis. In particular, some species appeared to be specialised and others appeared to be generalist, suggesting a nested pattern of resource use, rather than a clustered pattern in which each species uses a different subset of habitat types.</p></list-item><list-item><p>The degree of variation in specialisation among species varied with the number of species falling in each group. Thus, an increased number of species, and thus possibly competition, is more likely to promote the co‐occurrence of generalist and specialised species (nested patterns) rather than an increased niche segregation among species.</p></list-item><list-item><p>Ascertaining how species overlap their habitat use at a local scale can be relevant for conservation purposes, because specialised populations are potentially more susceptible to network distortions.</p></list-item></list></p></abstract><kwd-group kwd-group-type="author-generated"><kwd id="icad12193-kwd-0001">Butterfly</kwd><kwd id="icad12193-kwd-0002">ecological network</kwd><kwd id="icad12193-kwd-0003">habitat requirements</kwd><kwd id="icad12193-kwd-0004">habitat specialisation</kwd><kwd id="icad12193-kwd-0005">niche segregation</kwd></kwd-group><funding-group><award-group id="funding-0001"><funding-source>Ticino Regional Park</funding-source></award-group><award-group id="funding-0002"><funding-source><institution-wrap><institution>Universidad de Castilla‐La Mancha </institution><institution-id institution-id-type="open-funder-registry">10.13039/501100007480</institution-id></institution-wrap></funding-source></award-group><award-group id="funding-0003"><funding-source><institution-wrap><institution>European Social Fund (ESF) </institution><institution-id institution-id-type="open-funder-registry">10.13039/501100004895</institution-id></institution-wrap></funding-source></award-group></funding-group><counts><fig-count count="4"></fig-count><table-count count="2"></table-count><page-count count="11"></page-count><word-count count="7208"></word-count></counts><custom-meta-group><custom-meta><meta-name>source-schema-version-number</meta-name><meta-value>2.0</meta-value></custom-meta><custom-meta><meta-name>cover-date</meta-name><meta-value>November 2016</meta-value></custom-meta><custom-meta><meta-name>details-of-publishers-convertor</meta-name><meta-value>Converter:WILEY_ML3GV2_TO_JATSPMC version:5.8.0 mode:remove_FC converted:15.04.2020</meta-value></custom-meta></custom-meta-group></article-meta></front><body id="icad12193-body-0001"><sec id="icad12193-sec-0001"><title>Introduction</title><p>Butterflies (Lepidoptera) are a well‐studied insect group both taxonomically and ecologically (e.g. Gilbert, <xref rid="icad12193-bib-0027" ref-type="ref">1984</xref>; Kocher & Williams, <xref rid="icad12193-bib-0037" ref-type="ref">2000</xref>; Ghazoul, <xref rid="icad12193-bib-0026" ref-type="ref">2002</xref>; Stefanescu <italic>et al</italic>., <xref rid="icad12193-bib-0063" ref-type="ref">2004</xref>, <xref rid="icad12193-bib-0062" ref-type="ref">2011</xref>), particularly because they interact with plants both as larval herbivores and adult pollinators (Borges <italic>et al</italic>., <xref rid="icad12193-bib-0012" ref-type="ref">2003</xref>; Hardy <italic>et al</italic>., <xref rid="icad12193-bib-0030" ref-type="ref">2007</xref>) and have different habitat requirements even at each life stage (New <italic>et al</italic>., <xref rid="icad12193-bib-0047" ref-type="ref">1995</xref>; Samways & Lu, <xref rid="icad12193-bib-0056" ref-type="ref">2007</xref>; Dennis, <xref rid="icad12193-bib-0018" ref-type="ref">2010</xref>). In turn, because these interactions are important mechanisms of biological diversification (Leather, <xref rid="icad12193-bib-0038" ref-type="ref">1990</xref>; Basset, <xref rid="icad12193-bib-0005" ref-type="ref">1991a</xref>,<xref rid="icad12193-bib-0006" ref-type="ref">b</xref>; Slansky, <xref rid="icad12193-bib-0060" ref-type="ref">1993</xref>), the abundance and distribution of butterflies within a particular area can give insights into the environment's quality (Kocher & Williams, <xref rid="icad12193-bib-0037" ref-type="ref">2000</xref>).</p><p>These largely specialised butterfly–habitat associations are influenced by many physical and biological factors (Leps & Spitzer, <xref rid="icad12193-bib-0039" ref-type="ref">1990</xref>; Spitzer <italic>et al</italic>., <xref rid="icad12193-bib-0061" ref-type="ref">1993</xref>), including temperature, humidity, light levels, rainfall patterns, local microclimate conditions, grassland type and host–plant species richness and distribution (Hill <italic>et al</italic>., <xref rid="icad12193-bib-0031" ref-type="ref">2001</xref>; Meyer & Sisk, <xref rid="icad12193-bib-0043" ref-type="ref">2001</xref>; Collinge <italic>et al</italic>., <xref rid="icad12193-bib-0017" ref-type="ref">2003</xref>; Menéndez <italic>et al</italic>., <xref rid="icad12193-bib-0042" ref-type="ref">2007</xref>). All these variables define the <italic>resource‐based habitat</italic> or <italic>habitat requirements</italic> of each butterfly species, i.e., the habitat characterised by resources and conditions required by organisms and their access to them (Dennis, <xref rid="icad12193-bib-0018" ref-type="ref">2010</xref>). Furthermore, competition among species, presence of invasive species and avoidance strategies to escape from natural enemies (enemy‐free space) also affect species occurrence in a given area (Murphy, <xref rid="icad12193-bib-0046" ref-type="ref">2004</xref>; Dennis, <xref rid="icad12193-bib-0018" ref-type="ref">2010</xref>; Wagner & Van Driesche, <xref rid="icad12193-bib-0066" ref-type="ref">2010</xref>; Hanula & Horn, <xref rid="icad12193-bib-0029" ref-type="ref">2011</xref>).</p><p>While habitat requirements have been assessed for many butterfly species in detail, the degree to which species within an assemblage segregate their habitat use (i.e. how they are specialised in a subset of habitat types) has been less well studied.</p><p>The concepts of habitat <italic>requirements</italic> and habitat <italic>specialisation</italic> are strongly linked to the classical differences between the <italic>pure</italic> ecological niche based on the <italic>physiological optimum</italic> (Austin, <xref rid="icad12193-bib-0002" ref-type="ref">1980</xref>, <xref rid="icad12193-bib-0003" ref-type="ref">1985</xref>) (niche requirements) and the true and realised ecological niche (Hutchinson, <xref rid="icad12193-bib-0032" ref-type="ref">1957</xref>; Pulliam, <xref rid="icad12193-bib-0053" ref-type="ref">2000</xref>) (niche specialisation). Thus, while habitat requirements can be seen as a property of a species, habitat specialisation is a variable property that can vary with many interacting factors such as the presence of inter‐specific competitors, the population density and thus the intra‐specific competition and the body size of the individuals, among others (Chase & Leibold, <xref rid="icad12193-bib-0016" ref-type="ref">2003</xref>; Dennis <italic>et al</italic>., <xref rid="icad12193-bib-0020" ref-type="ref">2003</xref>, <xref rid="icad12193-bib-0019" ref-type="ref">2011</xref>; Rosenfeld, <xref rid="icad12193-bib-0055" ref-type="ref">2003</xref>).</p><p>Consider, for example, two butterfly species occurring in the same area that use similar habitat types, e.g., open grass areas and ecotones, and thus require the same habitat conditions. The two species may either use both habitat types equally (generalism) or segregate the use of the two habitat types (specialisation). Species may in fact overlap to different degrees the relative use of each type of habitat (Julliard <italic>et al</italic>., <xref rid="icad12193-bib-0034" ref-type="ref">2006</xref>; Poisot <italic>et al</italic>., <xref rid="icad12193-bib-0048" ref-type="ref">2011</xref>), a pattern that may occur either because of an evolutionary trade‐off between the ability of species to exploit a range of resources and their capability of using each one (Futuyma & Moreno, <xref rid="icad12193-bib-0025" ref-type="ref">1988</xref>), or because of factors such as inter‐specific competition (Schroder & Rosenzweig, <xref rid="icad12193-bib-0057" ref-type="ref">1975</xref>). It is thus important to analyse the two concepts (habitat requirements and habitat specialisation) simultaneously in a given species assemblage.</p><p>Here, we analysed the relationship between habitat requirements and habitat specialisation in an assemblage of butterflies in a riverine natural park in northern Italy. The aim was to test whether butterfly species with similar habitat requirements overlap their habitat use or, alternatively, they segregate their habitat use despite having similar habitat requirements. To do this, we first defined the requirements, i.e., which habitat types positively or negatively correlate with species abundances, and then used a network‐based approach (Blüthgen <italic>et al</italic>., <xref rid="icad12193-bib-0010" ref-type="ref">2006</xref>) to analyse the degree of habitat use overlap within groups of species with similar habitat requirements.</p></sec><sec id="icad12193-sec-0002"><title>Materials and methods</title><sec id="icad12193-sec-0003"><title>Study area</title><p>The study was conducted in the Ticino Regional Park (Lombardy, northern Italy, approximately from 45°44′0″N to 45°10′0″N and from 8°38′0″E to 9°16′0″E, with an elevation of 50–200 m a.s.l.) in summer 2008–2009. The study area is characterised by a continental climate with cold winters and hot summers and covers about 91 000 hectares. The park represents one of the most important green hot spots of the Po river Plain, which is otherwise very anthropised.</p><p>Within the park, four sites were chosen for the sampling programme. The two southern sites (coded as 1 and 4) (Fig. <xref rid="icad12193-fig-0001" ref-type="fig">1</xref>) were generally characterised by a thick riparian oak forest with some agricultural areas (poplar plantations, transects 1 and 2). Site 1 in particular showed a high degree of habitat diversity, including a flooded forest with <italic>Alnus glutinosa</italic>,<italic> Salix alba</italic> and <italic>Populus alba</italic>. The two northern sites (coded as 2 and 3) (Fig. <xref rid="icad12193-fig-0001" ref-type="fig">1</xref>) include areas that have previously been intensively used as military areas, and they can be considered partially anthropised areas. Although some natural parts still persist, the presence of alien and invasive plant species is relevant. These two sites were mainly characterised by open areas with shrubs (<italic>Calluna vulgaris</italic>,<italic> Crataegus monogyna</italic>,<italic> Rubus caesius</italic>,<italic> Cytisus scoparius</italic>) and by a low density of trees of exotic species, such as <italic>Robinia pseudoacacia</italic>,<italic> Ailanthus altissima</italic> and <italic>Prunus serotina</italic>.</p><fig fig-type="Figure" xml:lang="en" id="icad12193-fig-0001" orientation="portrait" position="float"><label>Figure 1</label><caption><p>Study area and sampling. (a) Location of the four plots (numbered 1–4) where species were sampled; (b) a plot in which the three transects performed are shown; (c) details of one transect showing the points recorded with the <styled-content style="fixed-case" toggle="no">GPS</styled-content> for a single species of butterfly throughout the whole study.</p></caption><graphic id="nlm-graphic-1" xlink:href="ICAD-9-495-g001"></graphic></fig></sec><sec id="icad12193-sec-0004"><title>Species sampling procedure</title><p>Sampling was carried out from the end of June to the first days of September for 9 weeks/year. Sites 1 and 2 were sampled in 2008, while sites 3 and 4 were sampled in 2009. Surveys were conducted between 09:30 and 17:00 h. Every site contained three 500 m line transects (<italic>n</italic> = 12 plots), in which we followed established guidelines for butterfly monitoring to minimise observer bias (Pollard & Yates, <xref rid="icad12193-bib-0050" ref-type="ref">1993</xref>). Observations were carried out during systematic walking surveys (75 min per plot visit – standardised search time) only in favourable temperature conditions (temperature >17 °C) and in calm to light winds. Every transect walk was repeated twice a week, for a total of 216 transects and 108 walked kilometres.</p><p>In contrast to previous research on butterflies, this study was based on a precise georeference of every sampled individual leading to a database of butterfly positions characterised by a large sample (<italic>n</italic> = 2924) and a good level of accuracy (Breed & Severns, <xref rid="icad12193-bib-0014" ref-type="ref">2015</xref>). All the points were recorded by a GPS (Garmin 60 CSx) whose standard error is considered to be between 4 and 7 m in good weather conditions. We considered the average GPS error to be two to seven times lower than the minimum detectable linear dimension of the available land cover, which is more accurate (minimum cartographic unit: 0.16 ha, scale 1:10 000) than land cover data used in other studies of butterfly species distributions (minimum cartographic unit: 2.25 ha, scale 1:50 000, Gutiérrez <italic>et al</italic>., <xref rid="icad12193-bib-0028" ref-type="ref">2010</xref>).</p></sec><sec id="icad12193-sec-0005"><title>GIS analyses and definition of habitat types</title><p>All the recorded points were imported and overlaid under a GIS environment (ESRI Arc View 3.2a) with the database of land cover classes (hereafter: habitat classes) describing the whole Lombardy region, thereby including the 12 plots (DUSAF, <italic>Destinazione Uso dei Suoli Agricoli e Forestali</italic>, ERSAF <xref rid="icad12193-bib-0023" ref-type="ref">2008</xref>). In the sampled areas, eight habitats were recognised together with a further seven ecotones between these habitats for a total of 15 habitat classes (see Table <xref rid="icad12193-tbl-0001" ref-type="table">1</xref>). The detail level is equal to an information scale of 1:10 000. For each land use feature with a polygonal configuration and a measurable surface, the minimum dimensional threshold of representability corresponds to 0.16 ha, and the smallest detectable linear dimension of the polygon is 20 m (<ext-link ext-link-type="uri" xlink:href="http://www.ersaf.lombardia.it/upload/ersaf/gestionedocumentale/WEB_Libro_suolo_EN_784_11577.pdf">http://www.ersaf.lombardia.it/upload/ersaf/gestionedocumentale/WEB_Libro_suolo_EN_784_11577.pdf</ext-link>).</p><table-wrap id="icad12193-tbl-0001" xml:lang="en" orientation="portrait" position="float"><label>Table 1</label><caption><p>Habitats in our study area in which butterfly species were sampled</p></caption><table frame="hsides" rules="groups"><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><thead valign="top"><tr style="border-bottom:solid 1px #000000"><th align="left" valign="top" rowspan="1" colspan="1">DUSAF code</th><th align="left" valign="top" rowspan="1" colspan="1">Habitat description</th><th align="left" valign="top" rowspan="1" colspan="1">Dominant plant species</th><th align="left" valign="top" rowspan="1" colspan="1">Phytosociological classification</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">31111</td><td align="left" rowspan="1" colspan="1">Coppice deciduous forests at high and medium density</td><td align="left" rowspan="1" colspan="1"><italic>Quercus robur</italic>,<italic> Carpinus betulus</italic></td><td align="left" rowspan="1" colspan="1"><italic>Carpinion betuli</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">31121</td><td align="left" rowspan="1" colspan="1">Coppice deciduous forests at low density</td><td align="left" rowspan="1" colspan="1"><italic>Quercus robur</italic>,<italic> Carpinus betulus</italic></td><td align="left" rowspan="1" colspan="1"><italic>Carpinion betuli</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">3113</td><td align="left" rowspan="1" colspan="1">Riparian forest</td><td align="left" rowspan="1" colspan="1"><italic>Alnus glutinosa</italic>,<italic> Salix alba</italic>,<italic> Populus alba</italic></td><td align="left" rowspan="1" colspan="1"><italic>Salicion albae</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">331</td><td align="left" rowspan="1" colspan="1">Beaches, dunes and stony riverbeds</td><td align="left" rowspan="1" colspan="1"><italic>Bidens frondosus</italic>,<italic> Persicaria hydropiper</italic>,<italic> Ranunculus sceleratus</italic></td><td align="left" rowspan="1" colspan="1"><italic>Bidention tripartitae</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">324</td><td align="left" rowspan="1" colspan="1">Shrubland or grassland with scattered trees</td><td align="left" rowspan="1" colspan="1"><italic>Calluna vulgaris</italic>,<italic> Cytisus scoparius</italic>,<italic> Molinia caerulea</italic></td><td align="left" rowspan="1" colspan="1"><italic>Calluno‐Genistion pilosae</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">2311</td><td align="left" rowspan="1" colspan="1">Permanent grassland in the absence of shrubs and trees</td><td align="left" rowspan="1" colspan="1"><italic>Arrhenatherum elatius</italic>,<italic> Dactylis glomerata</italic>,<italic> Holcus lanatus</italic>,<italic> Poa trivialis</italic>,<italic> Rumex acetosa</italic>,<italic> Achillea millefolium</italic></td><td align="left" rowspan="1" colspan="1"><italic>Arrhenatherion elatioris</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">2241</td><td align="left" rowspan="1" colspan="1">Poplar plantations</td><td align="left" rowspan="1" colspan="1"><italic>Populus</italic> spp.</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">1412</td><td align="left" rowspan="1" colspan="1">Uncultivated green areas</td><td align="left" rowspan="1" colspan="1"><italic>Artemisia vulgaris</italic>,<italic> Elytrigia repens</italic>,<italic> Melilotus officinalis</italic>,<italic> Solidago gigantea</italic>,<italic> Urtica dioica</italic></td><td align="left" rowspan="1" colspan="1"><italic>Dauco carotae – Melilotion albi</italic></td></tr></tbody></table><table-wrap-foot id="icad12193-ntgp-0001"><fn id="icad12193-note-0001"><p>A further seven habitat classes, i.e., ecotones between pairs of these habitats (1412_31111, 2311_31121, 2311_3113, 31111_31121, 31111_324, 31121_3113, 3113_331), were also recorded.</p></fn></table-wrap-foot><permissions><copyright-holder>John Wiley & Sons, Ltd</copyright-holder><license><license-p>This article is being made freely available through PubMed Central as part of the COVID-19 public health emergency response. It can be used for unrestricted research re-use and analysis in any form or by any means with acknowledgement of the original source, for the duration of the public health emergency.</license-p></license></permissions></table-wrap><p>All twelve 500 m transects were first divided into 300 sectors (<italic>n</italic> = 25 in each transect), each characterised by a length of 20 m and a width of 50 m, in order to consider the influence of habitat classes contiguous to the point of the supposed selection operated by every individual (Fig. <xref rid="icad12193-fig-0001" ref-type="fig">1</xref>). A double GIS spatial join application associated the field‐sampled points with the sectors previously associated with habitat classes (Fig. <xref rid="icad12193-fig-0001" ref-type="fig">1</xref>). When more than one habitat class fell into the same sector, we considered it as a separate habitat class composed and identified by the two source–habitat classes. We then counted the abundance of sampled individuals of each species in each sector and clustered the sectors according to the habitat class they showed, thereby calculating abundances for each habitat type.</p></sec><sec id="icad12193-sec-0006"><title>Habitat requirements of species</title><p>Data were prepared for analysis with the R package ‘vegan’ (version 2.0–5, The R Development Core Team <xref rid="icad12193-bib-0054" ref-type="ref">2011</xref>) to associate the different habitat preferences with each species and assess habitat requirements. The matrix had species as rows and line transects as columns. We calculated Morisita dissimilarity indices among species using the ‘vegdist’ function. Morisita is an index of patchiness, and it is used to calculate overlap among samples (Morisita, <xref rid="icad12193-bib-0045" ref-type="ref">1959</xref>). The dissimilarity matrix obtained was clustered specifying the ‘complete linkage’ method in the ‘hclust’ function in the ‘vegan’ package. We applied the ‘cascadeKM’ function to the standardised square root indices and applied the Calinski–Harabasz criterion (Milligan & Cooper, <xref rid="icad12193-bib-0044" ref-type="ref">1985</xref>) to select the best partition inside the dendrogram obtained by the cluster, thereby dividing the whole species group into subgroups with a similar use of habitat types.</p><p>In order to understand whether environmental characteristics influenced the abundance of individuals, we applied a generalised linear mixed effects model (GLMM) for count data (Atkins <italic>et al</italic>., <xref rid="icad12193-bib-0001" ref-type="ref">2012</xref>) for each of the previously selected groups. We ran zero‐inflated GLMMs due to the huge amount of zeros in our data set. GLMMs were run by applying the AD Model Builder procedure using the ‘glmmADMB’ package in R (Skaug <italic>et al</italic>., <xref rid="icad12193-bib-0059" ref-type="ref">2013</xref>), specifying the Poisson distribution inside the model (Bolker <italic>et al</italic>., <xref rid="icad12193-bib-0011" ref-type="ref">2012</xref>). Before starting the analysis, we applied a conservative method to calculate significance for each factor included in the model based on the evaluation of the standard deviation calculated for each estimate multiplied by 2. Consequently, estimates were significant only if the interval enclosed by two times their standard deviations did not cross 0. According to Breslow (<xref rid="icad12193-bib-0015" ref-type="ref">2004</xref>), generalised linear mixed models may yield biased estimates of the variance component when the mean number of counts per treatment combination is less than five, so we removed from the analysis all the species whose number of counted individuals was below this value. Land cover class classification and the year of the sample were inserted in the model as fixed effects and the plot as a random effect.</p></sec><sec id="icad12193-sec-0007"><title>Habitat–species network</title><p>Even though network analysis has been previously applied to feeding interactions (Bosch <italic>et al</italic>., <xref rid="icad12193-bib-0013" ref-type="ref">2009</xref>; Poulin, <xref rid="icad12193-bib-0051" ref-type="ref">2010</xref>; Junker <italic>et al</italic>., <xref rid="icad12193-bib-0035" ref-type="ref">2012</xref>; Polidori <italic>et al</italic>., <xref rid="icad12193-bib-0049" ref-type="ref">2013</xref>), it is also adequate for species–habitat webs within given areas when habitats are well characterised and thus represent discrete nodes of the webs. Ecologists continuously produce detailed classifications of habitat types based on sophisticated analyses of their components (e.g. soil, vegetation structure, climate), so that these classifications can be used as a basis for species–habitat webs.</p><p>Network indices based on information theory (Shannon entropy) were used to characterise our species–habitat webs in each area (2008 and 2009) and site (two in 2008 and two in 2009) (Dormann <italic>et al</italic>., <xref rid="icad12193-bib-0021" ref-type="ref">2009</xref>). The nodes of each network are composed of butterfly species versus their habitat types, and link weights are the number of individuals collected in each habitat type.</p><p>Because we were interested in the degree of habitat segregation between species, we considered the degree of complementarity (or exclusiveness) of interactions, defined as the standardised index <italic>H</italic><sub>2</sub>′ (Blüthgen <italic>et al</italic>., <xref rid="icad12193-bib-0010" ref-type="ref">2006</xref>). This index characterises the specialisation and habitat partitioning (niche overlap) in the butterfly assemblage. We first calculated the two‐dimensional Shannon index as:<disp-formula id="icad12193-disp-0001"><mml:math id="nlm-math-1"><mml:mrow><mml:msub><mml:mi>H</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mo>−</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>I</mml:mi></mml:munderover><mml:msup><mml:mrow><mml:munder><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:munder></mml:mrow><mml:mi>J</mml:mi></mml:msup><mml:mfrac><mml:msub><mml:mi>a</mml:mi><mml:mi mathvariant="normal">ij</mml:mi></mml:msub><mml:mi>m</mml:mi></mml:mfrac><mml:mo>·</mml:mo><mml:mi mathvariant="normal">ln</mml:mi><mml:mfrac><mml:msub><mml:mi>a</mml:mi><mml:mi mathvariant="normal">ij</mml:mi></mml:msub><mml:mi>m</mml:mi></mml:mfrac></mml:mrow></mml:math></disp-formula>where <italic>i</italic> is a butterfly species, <italic>I</italic> is the total number of individuals collected, <italic>j</italic> is one of <italic>J</italic> total habitat types, <italic>a</italic><sub>ij</sub> represents the number of records of the habitat type <italic>j</italic> frequented by the butterfly species <italic>i</italic> and <italic>m</italic> is the total number of habitat records associated with all butterfly species in a network.</p><p>We then standardised <italic>H</italic><sub>2</sub> against its minimum (<italic>H</italic><sub>2min</sub>) and maximum (<italic>H</italic><sub>2max</sub>) possible value for the given habitat type frequencies (i.e. marginal totals in the <italic>I </italic>× <italic>J</italic> matrix fixed), as follows:<disp-formula id="icad12193-disp-0002"><mml:math id="nlm-math-2"><mml:mrow><mml:msubsup><mml:mi>H</mml:mi><mml:mrow><mml:mn>2</mml:mn></mml:mrow><mml:mo>′</mml:mo></mml:msubsup><mml:mo>=</mml:mo><mml:mfrac><mml:mrow><mml:msub><mml:mi>H</mml:mi><mml:mrow><mml:mn>2</mml:mn><mml:mi mathvariant="normal">max</mml:mi></mml:mrow></mml:msub><mml:mo>−</mml:mo><mml:msub><mml:mi>H</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>H</mml:mi><mml:mrow><mml:mn>2</mml:mn><mml:mi mathvariant="normal">max</mml:mi></mml:mrow></mml:msub><mml:mo>−</mml:mo><mml:msub><mml:mi>H</mml:mi><mml:mrow><mml:mn>2</mml:mn><mml:mi mathvariant="normal">min</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:mfrac></mml:mrow></mml:math></disp-formula></p><p>Consequently, <italic>H</italic><sub>2</sub>′ ranges from 0 for the most generalised to 1 for the most specialised case.</p><p>Each value of <italic>H</italic><sub>2</sub>′ was tested against a null model of random associations (<italic>H</italic><sub>2ran</sub>). A number of random permutations of the matrix were performed using Patefield's <italic>r </italic>× <italic>c</italic> randomisation algorithm (Blüthgen <italic>et al</italic>., <xref rid="icad12193-bib-0010" ref-type="ref">2006</xref>) for which probability is derived (Fonseca & Ganade, <xref rid="icad12193-bib-0024" ref-type="ref">1996</xref>; Manly, <xref rid="icad12193-bib-0041" ref-type="ref">1997</xref>).</p><p>As a species‐level measure of <italic>partner diversity</italic>, we used the Kullback–Leibler distance (Blüthgen <italic>et al</italic>., <xref rid="icad12193-bib-0010" ref-type="ref">2006</xref>), which not only considers the diversity of partners but also their respective availability, thereby comparing the distribution of the interactions with each butterfly species (<italic>p</italic>′<italic>j</italic>) to the overall habitat type availability (<italic>qj</italic>); the index is defined as:<disp-formula id="icad12193-disp-0003"><mml:math id="nlm-math-3"><mml:mrow><mml:msub><mml:mi>d</mml:mi><mml:mi mathvariant="normal">i</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:munderover><mml:mo movablelimits="false">∑</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>c</mml:mi></mml:munderover><mml:mrow><mml:mo stretchy="false">(</mml:mo><mml:msubsup><mml:mi>p</mml:mi><mml:mi mathvariant="normal">ij</mml:mi><mml:mo>′</mml:mo></mml:msubsup></mml:mrow><mml:mo>·</mml:mo><mml:mi mathvariant="normal">Ln</mml:mi><mml:mfrac><mml:msubsup><mml:mi>p</mml:mi><mml:mi mathvariant="normal">ij</mml:mi><mml:mo>′</mml:mo></mml:msubsup><mml:msub><mml:mi>q</mml:mi><mml:mi mathvariant="normal">j</mml:mi></mml:msub></mml:mfrac><mml:mrow><mml:mo stretchy="false">)</mml:mo></mml:mrow></mml:mrow></mml:math></disp-formula>and then normalised as:<disp-formula id="icad12193-disp-0004"><mml:math id="nlm-math-4"><mml:mrow><mml:msubsup><mml:mi>d</mml:mi><mml:mi mathvariant="normal">i</mml:mi><mml:mo>′</mml:mo></mml:msubsup><mml:mo>=</mml:mo><mml:mfrac><mml:mrow><mml:msub><mml:mi>d</mml:mi><mml:mi mathvariant="normal">i</mml:mi></mml:msub><mml:mo>−</mml:mo><mml:msub><mml:mi>d</mml:mi><mml:mi mathvariant="normal">min</mml:mi></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>d</mml:mi><mml:mi mathvariant="normal">max</mml:mi></mml:msub><mml:mo>−</mml:mo><mml:msub><mml:mi>d</mml:mi><mml:mi mathvariant="normal">min</mml:mi></mml:msub></mml:mrow></mml:mfrac></mml:mrow></mml:math></disp-formula></p><p>The mean values of <italic>d</italic><sub>i</sub>′ between groups of species were compared using <sc>anova</sc>. Linear relationships between indices and other parameters were tested with Spearman's rank correlation test.</p></sec></sec><sec id="icad12193-sec-0008"><title>Results</title><sec id="icad12193-sec-0009"><title>Overall habitat requirements of species</title><p>Overall, 30 species of butterflies were collected. The Calinski‐Harabasz partition applied to the dissimilarity matrix (Fig. S1) suggested five subgroups of species based on their habitat requirements (Table <xref rid="icad12193-tbl-0002" ref-type="table">2</xref>, Fig. <xref rid="icad12193-fig-0002" ref-type="fig">2</xref>).</p><table-wrap id="icad12193-tbl-0002" xml:lang="en" orientation="portrait" position="float"><label>Table 2</label><caption><p>Species sampled, number of individuals collected (<italic>N</italic>), group assigned by the ‘cascadeKM’ analysis of the dendrogram based on the number of observations in the habitat classes and specialisation index (<italic>d</italic><sub>i</sub>′) for the species</p></caption><table frame="hsides" rules="groups"><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><col style="border-right:solid 1px #000000" span="1"></col><thead valign="top"><tr style="border-bottom:solid 1px #000000"><th align="left" valign="top" rowspan="1" colspan="1">Code</th><th align="left" valign="top" rowspan="1" colspan="1">Family</th><th align="left" valign="top" rowspan="1" colspan="1">Species</th><th align="char" valign="top" rowspan="1" colspan="1"><italic>N</italic></th><th align="char" valign="top" rowspan="1" colspan="1">Group</th><th align="char" valign="top" rowspan="1" colspan="1"><italic>d</italic><sub>i</sub>′</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Apatura ilia</italic> (Denis & Schiffermüller)</td><td align="char" char="." rowspan="1" colspan="1">6</td><td align="char" char="." rowspan="1" colspan="1">1</td><td align="char" char="." rowspan="1" colspan="1">0.242</td></tr><tr><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Argynnis paphia</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">18</td><td align="char" char="." rowspan="1" colspan="1">2</td><td align="char" char="." rowspan="1" colspan="1">0.200</td></tr><tr><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">Pieridae</td><td align="left" rowspan="1" colspan="1"><italic>Pieris mannii</italic> Mayer</td><td align="char" char="." rowspan="1" colspan="1">59</td><td align="char" char="." rowspan="1" colspan="1">4</td><td align="char" char="." rowspan="1" colspan="1">0.112</td></tr><tr><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">Pieridae</td><td align="left" rowspan="1" colspan="1"><italic>Pieris napi</italic> Linnaeus</td><td align="char" char="." rowspan="1" colspan="1">90</td><td align="char" char="." rowspan="1" colspan="1">4</td><td align="char" char="." rowspan="1" colspan="1">0.244</td></tr><tr><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">Pieridae</td><td align="left" rowspan="1" colspan="1"><italic>Pieris rapae</italic> Linnaeus</td><td align="char" char="." rowspan="1" colspan="1">60</td><td align="char" char="." rowspan="1" colspan="1">3</td><td align="char" char="." rowspan="1" colspan="1">0.166</td></tr><tr><td align="left" rowspan="1" colspan="1">6</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Brenthis daphne</italic> Bergsträsser </td><td align="char" char="." rowspan="1" colspan="1">89</td><td align="char" char="." rowspan="1" colspan="1">2</td><td align="char" char="." rowspan="1" colspan="1">0.044</td></tr><tr><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">Hesperiidae</td><td align="left" rowspan="1" colspan="1"><italic>Carcharodus alceae</italic> (Esper)</td><td align="char" char="." rowspan="1" colspan="1">3</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">Lycaenidae</td><td align="left" rowspan="1" colspan="1"><italic>Celastrina argiolus</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">70</td><td align="char" char="." rowspan="1" colspan="1">1</td><td align="char" char="." rowspan="1" colspan="1">0.166</td></tr><tr><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Coenonympha pamphilus</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">53</td><td align="char" char="." rowspan="1" colspan="1">2</td><td align="char" char="." rowspan="1" colspan="1">0.077</td></tr><tr><td align="left" rowspan="1" colspan="1">10</td><td align="left" rowspan="1" colspan="1">Pieridae</td><td align="left" rowspan="1" colspan="1"><italic>Colias crocea</italic> (Geoffroy)</td><td align="char" char="." rowspan="1" colspan="1">20</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.171</td></tr><tr><td align="left" rowspan="1" colspan="1">11</td><td align="left" rowspan="1" colspan="1">Lycaenidae</td><td align="left" rowspan="1" colspan="1"><italic>Cupido</italic> (<italic>Everes</italic>) <italic>argiades</italic> (Pallas)</td><td align="char" char="." rowspan="1" colspan="1">52</td><td align="char" char="." rowspan="1" colspan="1">3</td><td align="char" char="." rowspan="1" colspan="1">0.369</td></tr><tr><td align="left" rowspan="1" colspan="1">12</td><td align="left" rowspan="1" colspan="1">Pieridae</td><td align="left" rowspan="1" colspan="1"><italic>Gonepteryx rhamni</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">31</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.112</td></tr><tr><td align="left" rowspan="1" colspan="1">13</td><td align="left" rowspan="1" colspan="1">Hesperiidae</td><td align="left" rowspan="1" colspan="1"><italic>Hesperia comma</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">2</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1">14</td><td align="left" rowspan="1" colspan="1">Hesperiidae</td><td align="left" rowspan="1" colspan="1"><italic>Heteropterus morpheus</italic> (Pallas)</td><td align="char" char="." rowspan="1" colspan="1">62</td><td align="char" char="." rowspan="1" colspan="1">3</td><td align="char" char="." rowspan="1" colspan="1">0.168</td></tr><tr><td align="left" rowspan="1" colspan="1">15</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Aglais io</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">82</td><td align="char" char="." rowspan="1" colspan="1">3</td><td align="char" char="." rowspan="1" colspan="1">0.387</td></tr><tr><td align="left" rowspan="1" colspan="1">16</td><td align="left" rowspan="1" colspan="1">Papilionidae</td><td align="left" rowspan="1" colspan="1"><italic>Iphiclides podalirius</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">224</td><td align="char" char="." rowspan="1" colspan="1">1</td><td align="char" char="." rowspan="1" colspan="1">0.281</td></tr><tr><td align="left" rowspan="1" colspan="1">17</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Issoria lathonia</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">54</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.130</td></tr><tr><td align="left" rowspan="1" colspan="1">18</td><td align="left" rowspan="1" colspan="1">Lycaenidae</td><td align="left" rowspan="1" colspan="1"><italic>Lycaena phlaeas</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">26</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.197</td></tr><tr><td align="left" rowspan="1" colspan="1">19</td><td align="left" rowspan="1" colspan="1">Lycaenidae</td><td align="left" rowspan="1" colspan="1"><italic>Lycaena tityrus</italic> (Poda)</td><td align="char" char="." rowspan="1" colspan="1">45</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.198</td></tr><tr><td align="left" rowspan="1" colspan="1">20</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Maniola jurtina</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">166</td><td align="char" char="." rowspan="1" colspan="1">2</td><td align="char" char="." rowspan="1" colspan="1">0.155</td></tr><tr><td align="left" rowspan="1" colspan="1">21</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Melanargia galathaea</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">68</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.162</td></tr><tr><td align="left" rowspan="1" colspan="1">22</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Minois dryas</italic> (Scopoli)</td><td align="char" char="." rowspan="1" colspan="1">895</td><td align="char" char="." rowspan="1" colspan="1">2</td><td align="char" char="." rowspan="1" colspan="1">0.215</td></tr><tr><td align="left" rowspan="1" colspan="1">23</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Hipparchia statilinus</italic> (Hufnagel)</td><td align="char" char="." rowspan="1" colspan="1">192</td><td align="char" char="." rowspan="1" colspan="1">1</td><td align="char" char="." rowspan="1" colspan="1">0.603</td></tr><tr><td align="left" rowspan="1" colspan="1">24</td><td align="left" rowspan="1" colspan="1">Hesperiidae</td><td align="left" rowspan="1" colspan="1"><italic>Ochlodes sylvanus</italic> (Bremer & Grey)</td><td align="char" char="." rowspan="1" colspan="1">60</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.135</td></tr><tr><td align="left" rowspan="1" colspan="1">25</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Pararge aegeria</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">165</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.525</td></tr><tr><td align="left" rowspan="1" colspan="1">26</td><td align="left" rowspan="1" colspan="1">Lycaenidae</td><td align="left" rowspan="1" colspan="1"><italic>Plebejus argus</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">13</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.050</td></tr><tr><td align="left" rowspan="1" colspan="1">27</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Polygonia c‐album</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">52</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.239</td></tr><tr><td align="left" rowspan="1" colspan="1">28</td><td align="left" rowspan="1" colspan="1">Lycaenidae</td><td align="left" rowspan="1" colspan="1"><italic>Polyommatus icarus</italic> (Rottemburg)</td><td align="char" char="." rowspan="1" colspan="1">10</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.200</td></tr><tr><td align="left" rowspan="1" colspan="1">29</td><td align="left" rowspan="1" colspan="1">Hesperiidae</td><td align="left" rowspan="1" colspan="1"><italic>Pyrgus malvae</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">9</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.131</td></tr><tr><td align="left" rowspan="1" colspan="1">30</td><td align="left" rowspan="1" colspan="1">Nymphalidae</td><td align="left" rowspan="1" colspan="1"><italic>Vanessa atalanta</italic> (Linnaeus)</td><td align="char" char="." rowspan="1" colspan="1">46</td><td align="char" char="." rowspan="1" colspan="1">5</td><td align="char" char="." rowspan="1" colspan="1">0.217</td></tr></tbody></table><table-wrap-foot id="icad12193-ntgp-0002"><fn id="icad12193-note-0002"><p><italic>d</italic><sub>i</sub>′ was not calculated in cases of small sample size (denoted by ‘–’) (see text for details).</p></fn></table-wrap-foot><permissions><copyright-holder>John Wiley & Sons, Ltd</copyright-holder><license><license-p>This article is being made freely available through PubMed Central as part of the COVID-19 public health emergency response. It can be used for unrestricted research re-use and analysis in any form or by any means with acknowledgement of the original source, for the duration of the public health emergency.</license-p></license></permissions></table-wrap><fig fig-type="Figure" xml:lang="en" id="icad12193-fig-0002" orientation="portrait" position="float"><label>Figure 2</label><caption><p>Cluster dendrogram (above) obtained by applying Morisita dissimilarity indices based on the dissimilarity index matrix. The five group partitions determined by the Calinski criterion are shown by dotted lines. Below, there are the values of habitat specialisation (<italic>d</italic><sub>i</sub>′) for each species included in the five groups (codes for species above the bars, see Table <xref rid="icad12193-tbl-0002" ref-type="table">2</xref> for species names). Note that for a few species (e.g. eight), the sample size was too small to perform the network analysis (see text for details).</p></caption><graphic id="nlm-graphic-3" xlink:href="ICAD-9-495-g002"></graphic></fig><p>Following the GLMM results (Fig. <xref rid="icad12193-fig-0003" ref-type="fig">3</xref>), in the first group, we found a positive influence on the abundance of open areas (324, 1412) and a negative influence of natural dense deciduous forests (31111) and poplar plantations (2241) on the abundance. Ecotones between open and forest areas were also positively associated with abundance (31111_324, 1412_31111). Similarly, in the second group, we found a positive influence on the abundance of open heath land (324) and a negative influence on the abundance of poplar plantations and dense deciduous forests (31111). Many types of ecotones (31111_324, 2311_3113, 2311_31121) and riparian vegetation (3113) were also positively associated with the abundances of these species. Negative relationships between the abundance and open heath land (324) and its ecotone and natural dense deciduous forests (31111_324) were found in the third group, while the abundances of the species belonging to the fourth group were negatively associated with open anthropised heath areas (1412) and positively associated with the ecotone between open green areas and sparse deciduous forests (2311_31121). In the fifth group, positive influences on the abundance of sparse deciduous forests (31121), open green areas (2311) and poplar plantations (2241) were found. In addition, species of this group were positively associated with some ecotones, particularly those including riparian vegetation (31121_3113, 2311_3113). The year of sampling also affected the abundance of the species of groups 1, 2 and 3 (Fig. <xref rid="icad12193-fig-0003" ref-type="fig">3</xref>).</p><fig fig-type="Figure" xml:lang="en" id="icad12193-fig-0003" orientation="portrait" position="float"><label>Figure 3</label><caption><p>Estimated parameters derived from <styled-content style="fixed-case" toggle="no">GLMM</styled-content>. Significant influences of the parameters on the species abundance are shown by grey highlighting. The estimates (round black points) and their standard deviations multiplied by 2 (bold black lines enclosed by vertical segments) are provided. Standard deviation segments were cut when their values crossed −3 or +3 in each graph. Continuous bold lines without vertical segments are used when the model was not able to estimate reliable parameters. Significant influences of the parameters on the species abundance are shown by grey highlighting. Digits refer to habitat and ecotone codes as in Table <xref rid="icad12193-tbl-0001" ref-type="table">1</xref>.</p></caption><graphic id="nlm-graphic-5" xlink:href="ICAD-9-495-g003"></graphic></fig></sec><sec id="icad12193-sec-0010"><title>Habitat–species network</title><p>Overall, the relationships between species and habitats were not random in the studied butterfly assemblage (<italic>H</italic><sub>2</sub>′ = 0.302, <italic>P</italic> < 0.0001). Among species, habitat specialisation (<italic>d</italic><sub>i</sub>′) varied from 0.043 (denoting a generalist species) to 0.60 (denoting a highly specialised species) with an average of 0.21 ± 0.127, showing important variability.</p><p>The mean <italic>d</italic><sub>i</sub>′ calculated for each of the five groups identified by the Calinski criterion differed marginally (<italic>F</italic> = 2.9, df = 27, <italic>P</italic> = 0.04), and was highest in group 1 (species strongly positively associated with open habitats) and more similar among the other four groups (Fig. <xref rid="icad12193-fig-0004" ref-type="fig">4</xref>). The weakness of this difference was clearly due to the large variance within groups, i.e., species with similar habitat requirements were not all equally specialised (Table <xref rid="icad12193-tbl-0002" ref-type="table">2</xref>, Fig. <xref rid="icad12193-fig-0002" ref-type="fig">2</xref>). In particular, within all groups but one (group 2), it seemed that one or two species were much more specialised than the others. In group 2, this subdivision was more difficult but the difference between the highest <italic>d</italic><sub>i</sub>′ (species 1 and 22) and the lowest <italic>d</italic><sub>i</sub>′ (species 6 and 9) was still large. The within‐group variability in <italic>d</italic><sub>i</sub>′, when expressed with its coefficient of variation (CV), increased with the number of species included in the group (Spearman's correlation test, ρ<italic> </italic>= 0.97, <italic>n</italic> = 5, <italic>P</italic> = 0.017).</p><fig fig-type="Figure" xml:lang="en" id="icad12193-fig-0004" orientation="portrait" position="float"><label>Figure 4</label><caption><p>Box‐and‐whisker diagrams showing medians (horizontal lines within boxes), means (+), 1° and 3° quartile (top and bottom lines of the boxes) and maximum and minimum values (○) of habitat specialisation (<italic>d</italic><sub>i</sub>′) of the species included in the five groups determined by the Calinski criterion. Ends of the whiskers represent the lowest datum still within 1.5× interquartile range of the lower quartile and the highest datum still within 1.5× interquartile range of the upper quartile.</p></caption><graphic id="nlm-graphic-7" xlink:href="ICAD-9-495-g004"></graphic></fig></sec></sec><sec id="icad12193-sec-0011"><title>Discussion</title><p>We showed for the first time that high precision of hand‐held GPS georeferenced data can give detailed insights into how butterflies occupy possible suitable habitats based on their ecological needs, as previously shown for other animal groups (Lindeque & Lindeque, <xref rid="icad12193-bib-0040" ref-type="ref">1991</xref>; Kissling <italic>et al</italic>., <xref rid="icad12193-bib-0036" ref-type="ref">2014</xref>; Breed & Severns, <xref rid="icad12193-bib-0014" ref-type="ref">2015</xref>).</p><p>We found a coherent picture and a general rule by which butterfly species seemed to occupy suitable habitats based on their species‐specific ecological needs. In particular, species grouping seemed to be based on a continuum index of lightness, ranging from group 1 generally being influenced by open areas to group 5 being strongly related to closed‐canopy forests. If we look at the estimates associated with forest habitats and open habitats in our model (Fig. <xref rid="icad12193-fig-0003" ref-type="fig">3</xref>), we can see that they changed sign while proceeding from group 1 to group 5. In particular, forests are negatively related to the abundance of the sampled individuals in the first group, but they increase their positive influence while moving to group 5; the opposite occurs for the above‐mentioned general open dry areas. Furthermore, all the species, regardless of the group identity, seemed to avoid natural dense deciduous forests and associated ecotones. These are closed and shaded habitats, and luminosity was shown to be very important for butterflies, determining minimum and maximum flight conditions (Douwes, <xref rid="icad12193-bib-0022" ref-type="ref">1976</xref>) as well as flight duration (Shreeve, <xref rid="icad12193-bib-0058" ref-type="ref">1984</xref>). For example, in a restoration experiment of ponderosa pine forests in North America, butterfly species richness and abundance were up to three times greater in restoration treatment units where host plant and nectar plant species richness were similar to control units, but insolation (light intensity) was significantly greater (Waltz & Covington, <xref rid="icad12193-bib-0067" ref-type="ref">2004</xref>).</p><p>This differential importance of habitat light intensity in structuring our studied butterfly assemblage is confirmed by a comparison of our results with previous information on the habitat characteristics for the studied species, which revealed a general congruence. All the three species of the first group are reported to inhabit bushy places, and dry and hot areas, and are usually associated with forest ecotones and clearing margins (Table S1). Species of the second group showed a negative relationship with the presence of forest areas, and a positive association with open natural heath areas. Furthermore, species of group 2 seemed to be more influenced by ecotones than species of group 1, with grassland representing the key habitat type. We also found a positive influence of riparian vegetation on individual abundance, probably because most of the individuals belonging to group 2 were sampled in a site characterised by a single line of riparian vegetation and large open areas, thereby representing once again a sort of structural ecotone inside an open area. According to literature, many species of group 2 are defined as ecotone species (Table S1). Groups 3 and 4, on the other hand, which were negatively associated with open heath land and its ecotones, confirmed the decreasing importance of open areas while moving from group 1 to group 5. Indeed, species of group 5 were clearly associated with poplar plantations and sparse deciduous forests. Many species in group 5 were previously reported as being somehow linked to a forest environment (Table S1), but some species are reported to be associated with very different habitats (from open grassland to forests) (Table S1). Additionally, the significant effect of the year in some groups showed a temporal shift in species abundance, suggesting population fluctuations depending on change in abiotic or biotic conditions.</p><p>The degree of specialisation was not similar among species within groups in our studied assemblage. For example, <italic>Hipparchia statilinus</italic> was much more specialised than the other species of group 1, while <italic>Minois dryas</italic> frequented many more coppice deciduous forests and their ecotones than the other species of group 2. Two species of group 3 (<italic>Everes argiades</italic> and <italic>Aglais io</italic>), which in literature are reported as generalist species in many habitats including flowery bushy places, grassy banks, forest clearings, flowering meadows, forest edges and sheltered rocky gullies (Table S1), were very specialised in our study. Even the two closely related species (both from the genus <italic>Pieris</italic>) comprising group 4 differed greatly in specialisation.</p><p>Because the index of specialisation <italic>d</italic><sub>i</sub>′ increases with the level of specialisation, the positive correlation between the variance of <italic>d</italic>′ and the number of species across groups means that an increased number of species are more likely to promote the co‐occurrence of generalist and specialised species (nested patterns) rather than an increased niche segregation among species (a clustered pattern). Thus, possible increased competition (i.e. a higher number of species sharing the same habitat requirements) would not increase habitat segregation, while a few highly specialised species would use a subset of habitats exploited by more generalist species. This consideration is consistent with previous network‐based studies, which showed nestedness common in both mutualistic (e.g. plant–pollinator) networks (Bascompte <italic>et al</italic>., <xref rid="icad12193-bib-0004" ref-type="ref">2003</xref>) and agonistic (e.g. host–parasite) networks (Joppa <italic>et al</italic>., <xref rid="icad12193-bib-0033" ref-type="ref">2010</xref>) (although compartmentalisation is also observed in agonistic webs, e.g. Prado & Lewinsohn, <xref rid="icad12193-bib-0052" ref-type="ref">2004</xref>). Bastolla <italic>et al</italic>. (<xref rid="icad12193-bib-0007" ref-type="ref">2009</xref>) showed that the process by which a new species enters a community will likely lead to a nested pattern of interactions, because the new species tries to minimise its competitive load, an action that forces it to interact with the most generalist species. This could also be applied to habitats: When a new species arrives in a given area, it will first select sites with certain characteristics (requirements) and then reduce the overlap with competing species, largely using only a subset of the habitats used by generalist species.</p></sec><sec id="icad12193-sec-0012"><title>Conclusion</title><p>Our results stress the importance of considering both the habitat requirements of species and the specialisation in habitat use by each species when analysing butterfly assemblages. Indeed, many species studied here, previously known as generalists, were found to be locally specialised in our analysis. This is particularly important in the light of butterfly conservation, because the more specialised species may also be those with higher susceptibility to network distortions and thus with higher fragility to co‐extinction (Blüthgen, <xref rid="icad12193-bib-0008" ref-type="ref">2010</xref>). This would provide elements for deciding protection priorities for the species at local scales.</p><p>Although none of the sampled species are listed as threatened in the European Red List of Lepidoptera (with the exception of <italic>Hipparchia statilinus</italic>, which is listed as Near Threatened, Van Swaay <italic>et al</italic>., <xref rid="icad12193-bib-0064" ref-type="ref">2010</xref>), five species (<italic>Coenonympha pamphilus</italic>,<italic> Lycaena phlaeas</italic>,<italic> Maniola jurtina</italic>,<italic> Ochlodes sylvanus</italic> and <italic>Polyommatus icarus</italic>) are reported as being declined in Europe since 1990 (Van Swaay <italic>et al</italic>., <xref rid="icad12193-bib-0065" ref-type="ref">2013</xref>). These species may become locally vulnerable through persistent increased habitat specialisation associated with habitat disturbance; therefore, their ecology should be finely monitored in order to adopt adequate conservation plans.</p></sec><sec sec-type="supplementary-material"><title>Supporting information</title><supplementary-material content-type="local-data"><caption><p><bold>Fig. S1.</bold> Calinski criterion applied to the dissimilarity matrix. The number of possible group partitions is shown on the <italic>y</italic>‐axis and the values obtained by the Calinski–Harabasz algorithm are shown on the <italic>x</italic>‐axis.</p></caption><media xlink:href="ICAD-9-495-s001.tiff"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data"><caption><p><bold>Table S1.</bold> Previous published information on the habitat requirements of the studied butterfly species.</p></caption><media xlink:href="ICAD-9-495-s002.doc"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material></sec></body><back><ack id="icad12193-sec-0013"><title>Acknowledgements</title><p>This work could not have been carried out without the help of Stefano Basso and Ester Manzini, students who helped greatly with the fieldwork. Thanks are also due to Marina Trentin for the support she provided during and after the fieldwork. This work was made possible by funding provided by Ticino Regional Park. CP was funded by a postdoctoral contract funded by the Universidad de Castilla‐La Mancha and the European Social Fund (ESF). Kelsey Horvath kindly revised the English editing.</p></ack><ref-list content-type="cited-references" id="icad12193-bibl-0001"><title>References</title><ref id="icad12193-bib-0001"><mixed-citation publication-type="journal" id="icad12193-cit-0001"><string-name><surname>Atkins</surname>, <given-names>D.C.</given-names></string-name>, <string-name><surname>Baldwin</surname>, <given-names>S.A.</given-names></string-name>, <string-name><surname>Zheng</surname>, <given-names>C.</given-names></string-name>, <string-name><surname>Gallop</surname>, <given-names>R.J.</given-names></string-name> & <string-name><surname>Neighbors</surname>, <given-names>C.A.</given-names></string-name> (<year>2012</year>) <article-title>Tutorial on Count Regression and Zero‐Altered Count Models for Longitudinal Substance Use Data</article-title>. <source xml:lang="en">Psychology of Addictive Behaviors</source>, <volume>27</volume>, <fpage>166</fpage>–<lpage>177</lpage>.<pub-id pub-id-type="pmid">22905895</pub-id></mixed-citation></ref><ref id="icad12193-bib-0002"><mixed-citation publication-type="journal" id="icad12193-cit-0002"><string-name><surname>Austin</surname>, <given-names>M.P.</given-names></string-name> (<year>1980</year>) <article-title>Searching for a model for use in vegetation analysis</article-title>. <source xml:lang="en">Vegetation</source>, <volume>42</volume>, <fpage>11</fpage>–<lpage>21</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0003"><mixed-citation publication-type="journal" id="icad12193-cit-0003"><string-name><surname>Austin</surname>, <given-names>M.P.</given-names></string-name> (<year>1985</year>) <article-title>Continuum concept, ordination methods and niche theory</article-title>. <source xml:lang="en">Annual Review of Ecology and Systematic</source>, <volume>16</volume>, <fpage>39</fpage>–<lpage>61</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0004"><mixed-citation publication-type="journal" id="icad12193-cit-0004"><string-name><surname>Bascompte</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>Jordano</surname>, <given-names>P.</given-names></string-name>, <string-name><surname>Melia′n</surname>, <given-names>C.J.</given-names></string-name> & <string-name><surname>Olesen</surname>, <given-names>J.M.</given-names></string-name> (<year>2003</year>) <article-title>The nested assembly of plant–animal mutualistic networks</article-title>. <source xml:lang="en">Proceedings of Natural Academy of Science USA</source>, <volume>100</volume>, <fpage>9383</fpage>–<lpage>9387</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0005"><mixed-citation publication-type="journal" id="icad12193-cit-0005"><string-name><surname>Basset</surname>, <given-names>Y.</given-names></string-name> (<year>1991a</year>) <article-title>The seasonality of arboreal arthropods foraging within an Australian rainforest tree</article-title>. <source xml:lang="en">Ecological Entomology</source>, <volume>16</volume>, <fpage>265</fpage>–<lpage>278</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0006"><mixed-citation publication-type="journal" id="icad12193-cit-0006"><string-name><surname>Basset</surname>, <given-names>Y.</given-names></string-name> (<year>1991b</year>) <article-title>Leaf production of an over storey rainforest tree and its effects on the temporal distribution of associated insect herbivores</article-title>. <source xml:lang="en">Oecologia</source>, <volume>88</volume>, <fpage>211</fpage>–<lpage>219</lpage>.<pub-id pub-id-type="pmid">28312134</pub-id></mixed-citation></ref><ref id="icad12193-bib-0007"><mixed-citation publication-type="journal" id="icad12193-cit-0007"><string-name><surname>Bastolla</surname>, <given-names>U.</given-names></string-name>, <string-name><surname>Fortuna</surname>, <given-names>M.</given-names></string-name>, <string-name><surname>Pascual‐Garcìa</surname>, <given-names>A.</given-names></string-name>, <string-name><surname>Ferrera</surname>, <given-names>A.</given-names></string-name>, <string-name><surname>Luque</surname>, <given-names>B.</given-names></string-name> & <string-name><surname>Bascompte</surname>, <given-names>J.</given-names></string-name> (<year>2009</year>) <article-title>The architecture of mutualistic networks minimizes competition and increases biodiversity</article-title>. <source xml:lang="en">Nature</source>, <volume>458</volume>, <fpage>1018</fpage>–<lpage>1020</lpage>.<pub-id pub-id-type="pmid">19396144</pub-id></mixed-citation></ref><ref id="icad12193-bib-0008"><mixed-citation publication-type="journal" id="icad12193-cit-0008"><string-name><surname>Blüthgen</surname>, <given-names>N.</given-names></string-name> (<year>2010</year>) <article-title>Why network analysis is often disconnected from community ecology: a critique and an ecologist's guide</article-title>. <source xml:lang="en">Basic and Applied Ecology</source>, <volume>11</volume>, <fpage>185</fpage>–<lpage>195</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0010"><mixed-citation publication-type="journal" id="icad12193-cit-0010"><string-name><surname>Blüthgen</surname>, <given-names>N.</given-names></string-name>, <string-name><surname>Menzel</surname>, <given-names>F.</given-names></string-name> & <string-name><surname>Blüthgen</surname>, <given-names>N.</given-names></string-name> (<year>2006</year>) <article-title>Measuring specialization in species interaction networks</article-title>. <source xml:lang="en">BMC Ecology</source>, <volume>6</volume>, <fpage>12</fpage>.<pub-id pub-id-type="pmid">16965615</pub-id></mixed-citation></ref><ref id="icad12193-bib-0011"><mixed-citation publication-type="miscellaneous" id="icad12193-cit-0011"><string-name><surname>Bolker</surname>, <given-names>B.</given-names></string-name>, <string-name><surname>Skaug</surname>, <given-names>H.</given-names></string-name>, <string-name><surname>Magnusson</surname>, <given-names>A.</given-names></string-name> & <string-name><surname>Nielsen</surname>, <given-names>A.</given-names></string-name> (<year>2012</year>) <article-title>Getting started with the glmmADMB package</article-title>. <<ext-link ext-link-type="uri" xlink:href="http://glmmadmb.r-forge.r-project.org/glmmADMB.pdf">http://glmmadmb.r-forge.r-project.org/glmmADMB.pdf</ext-link>>.</mixed-citation></ref><ref id="icad12193-bib-0012"><mixed-citation publication-type="journal" id="icad12193-cit-0012"><string-name><surname>Borges</surname>, <given-names>R.M.</given-names></string-name>, <string-name><surname>Gowda</surname>, <given-names>V.</given-names></string-name> & <string-name><surname>Zacharias</surname>, <given-names>M.</given-names></string-name> (<year>2003</year>) <article-title>Butterfly pollination and high contrast visual signals in a low‐density distylous plant</article-title>. <source xml:lang="en">Oecologia</source>, <volume>136</volume>, <fpage>571</fpage>–<lpage>573</lpage>.<pub-id pub-id-type="pmid">12845517</pub-id></mixed-citation></ref><ref id="icad12193-bib-0013"><mixed-citation publication-type="journal" id="icad12193-cit-0013"><string-name><surname>Bosch</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>González</surname>, <given-names>A.M.</given-names></string-name>, <string-name><surname>Rodrigo</surname>, <given-names>A.</given-names></string-name> & <string-name><surname>Navarro</surname>, <given-names>D.</given-names></string-name> (<year>2009</year>) <article-title>Plant‐pollinator networks: adding the pollinator's perspective</article-title>. <source xml:lang="en">Ecology Letters</source>, <volume>12</volume>, <fpage>409</fpage>–<lpage>419</lpage>.<pub-id pub-id-type="pmid">19379135</pub-id></mixed-citation></ref><ref id="icad12193-bib-0014"><mixed-citation publication-type="journal" id="icad12193-cit-0014"><string-name><surname>Breed</surname>, <given-names>G.A.</given-names></string-name> & <string-name><surname>Severns</surname>, <given-names>P.M.</given-names></string-name> (<year>2015</year>) <article-title>Low relative error in consumer‐grade GPS units make them ideal for measuring small‐scale animal movement patterns</article-title>. <source xml:lang="en">PeerJ</source>, <volume>3</volume>, <fpage>e1205</fpage>; doi:<pub-id pub-id-type="doi">10.7717/peerj.1205</pub-id>.<pub-id pub-id-type="pmid">26312190</pub-id></mixed-citation></ref><ref id="icad12193-bib-0015"><mixed-citation publication-type="book" id="icad12193-cit-0015"><string-name><surname>Breslow</surname>, <given-names>N.E.</given-names></string-name> (<year>2004</year>) <chapter-title>Whither PQL?</chapter-title><source xml:lang="en">Proceedings of the Second Seattle Symposium in Biostatistics: analysis of Correlated Data</source> (ed. by <person-group person-group-type="editor"><name name-style="western"><surname>Lin</surname><given-names>D.Y.</given-names></name></person-group> and <person-group person-group-type="editor"><name name-style="western"><surname>Heagerty</surname><given-names>P.J.</given-names></name></person-group>), pp. <fpage>1</fpage>–<lpage>22</lpage>. <publisher-name>Springer</publisher-name>, <publisher-loc>New York City, New York</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0016"><mixed-citation publication-type="book" id="icad12193-cit-0016"><string-name><surname>Chase</surname>, <given-names>J.M.</given-names></string-name> & <string-name><surname>Leibold</surname>, <given-names>M.A.</given-names></string-name> (<year>2003</year>) <source xml:lang="en">Ecological niches: linking classical and contemporary approaches</source>. <publisher-name>University of Chicago Press</publisher-name>, <publisher-loc>Chicago, Illinois</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0017"><mixed-citation publication-type="journal" id="icad12193-cit-0017"><string-name><surname>Collinge</surname>, <given-names>S.K.</given-names></string-name>, <string-name><surname>Prudic</surname>, <given-names>K.L.</given-names></string-name> & <string-name><surname>Oliver</surname>, <given-names>J.C.</given-names></string-name> (<year>2003</year>) <article-title>Effects of local habitat characteristics and landscape context on grassland butterfly diversity</article-title>. <source xml:lang="en">Conservation Biology</source>, <volume>17</volume>, <fpage>178</fpage>–<lpage>187</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0018"><mixed-citation publication-type="book" id="icad12193-cit-0018"><string-name><surname>Dennis</surname>, <given-names>R.L.H.</given-names></string-name> (<year>2010</year>) <source xml:lang="en">A Resource‐Based Habitat View for Conservation. Butterflies in the British Landscape</source>. <publisher-name>Wiley‐Blackwell</publisher-name>, <publisher-loc>Oxford, UK</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0019"><mixed-citation publication-type="journal" id="icad12193-cit-0019"><string-name><surname>Dennis</surname>, <given-names>R.L.H.</given-names></string-name>, <string-name><surname>Dapporto</surname>, <given-names>L.</given-names></string-name>, <string-name><surname>Fattorini</surname>, <given-names>S.</given-names></string-name> & <string-name><surname>Cook</surname>, <given-names>L.M.</given-names></string-name> (<year>2011</year>) <article-title>The generalism‐specialism debate: the role of generalists in the life and death of species</article-title>. <source xml:lang="en">Biological Journal of the Linnean Society</source>, <volume>104</volume>, <fpage>725</fpage>–<lpage>737</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0020"><mixed-citation publication-type="journal" id="icad12193-cit-0020"><string-name><surname>Dennis</surname>, <given-names>R.L.H.</given-names></string-name>, <string-name><surname>Shreeve</surname>, <given-names>T.G.</given-names></string-name> & <string-name><surname>Van Dyck</surname>, <given-names>H.</given-names></string-name> (<year>2003</year>) <article-title>Towards a functional resource‐based concept for habitat: a butterfly biology viewpoint</article-title>. <source xml:lang="en">Oikos</source>, <volume>102</volume>, <fpage>417</fpage>–<lpage>426</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0021"><mixed-citation publication-type="journal" id="icad12193-cit-0021"><string-name><surname>Dormann</surname>, <given-names>C.F.</given-names></string-name>, <string-name><surname>Fründ</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>Blüthgen</surname>, <given-names>N.</given-names></string-name> & <string-name><surname>Gruber</surname>, <given-names>B.</given-names></string-name> (<year>2009</year>) <article-title>Indices, graphs and null models: analysing bipartite ecological networks</article-title>. <source xml:lang="en">The Open Ecology Journal</source>, <volume>2</volume>, <fpage>7</fpage>–<lpage>24</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0022"><mixed-citation publication-type="journal" id="icad12193-cit-0022"><string-name><surname>Douwes</surname>, <given-names>P.</given-names></string-name> (<year>1976</year>) <article-title>Activity in Heodes virgaureae (Lep., Lycaenidae) in relation to air temperature, solar radiation, and time of day</article-title>. <source xml:lang="en">Oecologia</source>, <volume>22</volume>, <fpage>287</fpage>–<lpage>298</lpage>.<pub-id pub-id-type="pmid">28308692</pub-id></mixed-citation></ref><ref id="icad12193-bib-0023"><mixed-citation publication-type="book" id="icad12193-cit-0023"><collab collab-type="authors">ERSAF</collab>. (<year>2008</year>) <source xml:lang="en">Destinazione d'Uso dei Suoli Agricoli e Forestali</source>. <publisher-name>Ente Regionale per i Servizi all'Agricoltura e alle Foreste della Lombardia</publisher-name>, <publisher-loc>Milano, Italy</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0024"><mixed-citation publication-type="journal" id="icad12193-cit-0024"><string-name><surname>Fonseca</surname>, <given-names>C.R.</given-names></string-name> & <string-name><surname>Ganade</surname>, <given-names>G.</given-names></string-name> (<year>1996</year>) <article-title>Asymmetries, compartments and null 485 interactions in an Amazonian ant‐plant community</article-title>. <source xml:lang="en">Journal of Animal Ecology</source>, <volume>65</volume>, <fpage>339</fpage>–<lpage>347</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0025"><mixed-citation publication-type="journal" id="icad12193-cit-0025"><string-name><surname>Futuyma</surname>, <given-names>D.J.</given-names></string-name> & <string-name><surname>Moreno</surname>, <given-names>G.</given-names></string-name> (<year>1988</year>) <article-title>The evolution of ecological specialisation</article-title>. <source xml:lang="en">Annual Review of Ecology and Systematic</source>, <volume>19</volume>, <fpage>207</fpage>–<lpage>233</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0026"><mixed-citation publication-type="journal" id="icad12193-cit-0026"><string-name><surname>Ghazoul</surname>, <given-names>J.</given-names></string-name> (<year>2002</year>) <article-title>Impact of logging on the richness and diversity of forest butterflies in a tropical dry forest in Thailand</article-title>. <source xml:lang="en">Biodiversity and Conservation</source>, <volume>11</volume>, <fpage>521</fpage>–<lpage>541</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0027"><mixed-citation publication-type="book" id="icad12193-cit-0027"><string-name><surname>Gilbert</surname>, <given-names>L.E.</given-names></string-name> (<year>1984</year>) <chapter-title>The Biology of Butterfly Communities</chapter-title><source xml:lang="en">The Biology of Butterflies</source> (ed. by <person-group person-group-type="editor"><name name-style="western"><surname>Vane‐Wright</surname><given-names>R.I.</given-names></name></person-group> and <person-group person-group-type="editor"><name name-style="western"><surname>Ackery</surname><given-names>P.R.</given-names></name></person-group>), Vol. 11, pp. <fpage>41</fpage>–<lpage>54</lpage>. Symposium of the Royal Entomological Society of London. <publisher-name>Academic Press</publisher-name>, <publisher-loc>London, UK</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0028"><mixed-citation publication-type="journal" id="icad12193-cit-0028"><string-name><surname>Gutiérrez</surname>, <given-names>J.I.</given-names></string-name>, <string-name><surname>Gutiérrez</surname>, <given-names>D.</given-names></string-name> & <string-name><surname>Wilson</surname>, <given-names>R.J.</given-names></string-name> (<year>2010</year>) <article-title>The contributions of topoclimate and land cover to species distributions and abundance: fine‐resolution tests for a mountain butterfly fauna</article-title>. <source xml:lang="en">Global Ecology and Biogeography</source>, <volume>19</volume>, <fpage>159</fpage>–<lpage>173</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0029"><mixed-citation publication-type="journal" id="icad12193-cit-0029"><string-name><surname>Hanula</surname>, <given-names>J.L.</given-names></string-name> & <string-name><surname>Horn</surname>, <given-names>S.</given-names></string-name> (<year>2011</year>) <article-title>Removing an exotic shrub from riparian forest increases butterfly abundance and diversity</article-title>. <source xml:lang="en">Forest Ecology and Management</source>, <volume>262</volume>, <fpage>674</fpage>–<lpage>680</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0030"><mixed-citation publication-type="journal" id="icad12193-cit-0030"><string-name><surname>Hardy</surname>, <given-names>P.B.</given-names></string-name>, <string-name><surname>Sparks</surname>, <given-names>T.H.</given-names></string-name>, <string-name><surname>Isaac</surname>, <given-names>N.J.B.</given-names></string-name> & <string-name><surname>Dennis</surname>, <given-names>R.L.H.</given-names></string-name> (<year>2007</year>) <article-title>Specialism for larval and adult consumer resources among British butterflies: implications for conservation</article-title>. <source xml:lang="en">Biological Conservation</source>, <volume>138</volume>, <fpage>440</fpage>–<lpage>452</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0031"><mixed-citation publication-type="journal" id="icad12193-cit-0031"><string-name><surname>Hill</surname>, <given-names>J.K.</given-names></string-name>, <string-name><surname>Hamer</surname>, <given-names>K.C.</given-names></string-name>, <string-name><surname>Tangah</surname>, <given-names>J.</given-names></string-name> & <string-name><surname>Dawood</surname>, <given-names>M.</given-names></string-name> (<year>2001</year>) <article-title>Ecology of Tropical butterflies in rainforest gaps</article-title>. <source xml:lang="en">Oecologia</source>, <volume>128</volume>, <fpage>294</fpage>–<lpage>302</lpage>.<pub-id pub-id-type="pmid">28547478</pub-id></mixed-citation></ref><ref id="icad12193-bib-0032"><mixed-citation publication-type="journal" id="icad12193-cit-0032"><string-name><surname>Hutchinson</surname>, <given-names>G.E.</given-names></string-name> (<year>1957</year>) <article-title>Concluding remarks</article-title>. <source xml:lang="en">Cold Spring Harbor Symposia on Quantitative Biology</source>, <volume>22</volume>, <fpage>415</fpage>–<lpage>427</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0033"><mixed-citation publication-type="journal" id="icad12193-cit-0033"><string-name><surname>Joppa</surname>, <given-names>L.N.</given-names></string-name>, <string-name><surname>Montoya</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>Sole</surname>, <given-names>R.</given-names></string-name>, <string-name><surname>Sanderson</surname>, <given-names>J.</given-names></string-name> & <string-name><surname>Pimm</surname>, <given-names>S.L.</given-names></string-name> (<year>2010</year>) <article-title>On nestedness in ecological networks</article-title>. <source xml:lang="en">Evolutionary and Ecology Research</source>, <volume>12</volume>, <fpage>35</fpage>–<lpage>46</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0034"><mixed-citation publication-type="journal" id="icad12193-cit-0034"><string-name><surname>Julliard</surname>, <given-names>R.</given-names></string-name>, <string-name><surname>Clavel</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>Devictor</surname>, <given-names>V.</given-names></string-name>, <string-name><surname>Jiguet</surname>, <given-names>F.</given-names></string-name> & <string-name><surname>Couvet</surname>, <given-names>D.</given-names></string-name> (<year>2006</year>) <article-title>Spatial segregation of specialists and generalists in bird communities</article-title>. <source xml:lang="en">Ecology Letters</source>, <volume>9</volume>, <fpage>1237</fpage>–<lpage>1244</lpage>.<pub-id pub-id-type="pmid">17040326</pub-id></mixed-citation></ref><ref id="icad12193-bib-0035"><mixed-citation publication-type="journal" id="icad12193-cit-0035"><string-name><surname>Junker</surname>, <given-names>R.R.</given-names></string-name>, <string-name><surname>Blüthgen</surname>, <given-names>N.</given-names></string-name>, <string-name><surname>Brehm</surname>, <given-names>T.</given-names></string-name>, <string-name><surname>Binkenstein</surname>, <given-names>J.</given-names></string-name> & <string-name><surname>Paulus</surname>, <given-names>J.</given-names></string-name> (<year>2012</year>) <article-title>Specialization on traits as basis for the niche‐breadth of flower visitors and as structuring mechanism of ecological networks</article-title>. <source xml:lang="en">Functional Ecology</source>, <volume>27</volume>, <fpage>329</fpage>–<lpage>341</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0036"><mixed-citation publication-type="journal" id="icad12193-cit-0036"><string-name><surname>Kissling</surname>, <given-names>D.W.</given-names></string-name>, <string-name><surname>Pattemore</surname>, <given-names>D.E.</given-names></string-name> & <string-name><surname>Hagen</surname>, <given-names>M.</given-names></string-name> (<year>2014</year>) <article-title>Challenges and prospects in the telemetry of insects</article-title>. <source xml:lang="en">Biological Reviews</source>, <volume>89</volume>, <fpage>511</fpage>–<lpage>530</lpage>.<pub-id pub-id-type="pmid">24106908</pub-id></mixed-citation></ref><ref id="icad12193-bib-0037"><mixed-citation publication-type="journal" id="icad12193-cit-0037"><string-name><surname>Kocher</surname>, <given-names>S.D.</given-names></string-name> & <string-name><surname>Williams</surname>, <given-names>E.H.</given-names></string-name> (<year>2000</year>) <article-title>The diversity and abundance of North American butterflies, vary with habitat disturbance and geography</article-title>. <source xml:lang="en">Journal of Biogeography</source>, <volume>27</volume>, <fpage>785</fpage>–<lpage>794</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0038"><mixed-citation publication-type="journal" id="icad12193-cit-0038"><string-name><surname>Leather</surname>, <given-names>S.R.</given-names></string-name> (<year>1990</year>) <article-title>The analysis of species‐area relationships, with particular reference to macrolepidoptera on Rosaceae: how important is insect data‐set quality?</article-title><source xml:lang="en">Entomologist</source>, <volume>109</volume>, <fpage>8</fpage>–<lpage>16</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0039"><mixed-citation publication-type="journal" id="icad12193-cit-0039"><string-name><surname>Leps</surname>, <given-names>J.</given-names></string-name> & <string-name><surname>Spitzer</surname>, <given-names>K.</given-names></string-name> (<year>1990</year>) <article-title>Ecological determinants of butterfly communities (Lepidoptera, Papilionoidea) in the Tam Dao Mountains, Vietnam</article-title>. <source xml:lang="en">Acta Entomology Bohemoslov</source>, <volume>87</volume>, <fpage>182</fpage>–<lpage>194</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0040"><mixed-citation publication-type="journal" id="icad12193-cit-0040"><string-name><surname>Lindeque</surname>, <given-names>M.</given-names></string-name> & <string-name><surname>Lindeque</surname>, <given-names>J.M.</given-names></string-name> (<year>1991</year>) <article-title>Satellite tracking of elephants in north‐western Namibia</article-title>. <source xml:lang="en">African Journal of Ecology</source>, <volume>29</volume>, <fpage>196</fpage>–<lpage>206</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0041"><mixed-citation publication-type="book" id="icad12193-cit-0041"><string-name><surname>Manly</surname>, <given-names>B.</given-names></string-name> (<year>1997</year>) <source xml:lang="en">Randomization bootstrap and Monte Carlo methods in biology</source>. <publisher-name>Chapman and Hall</publisher-name>, <publisher-loc>London, UK</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0042"><mixed-citation publication-type="journal" id="icad12193-cit-0042"><string-name><surname>Menéndez</surname>, <given-names>R.</given-names></string-name>, <string-name><surname>González‐Megías</surname>, <given-names>A.</given-names></string-name>, <string-name><surname>Collingham</surname>, <given-names>Y.</given-names></string-name>, <string-name><surname>Fox</surname>, <given-names>R.</given-names></string-name>, <string-name><surname>Roy</surname>, <given-names>D.B.</given-names></string-name>, <string-name><surname>Ohlemüller</surname>, <given-names>R.</given-names></string-name> & <string-name><surname>Thomas</surname>, <given-names>C.D.</given-names></string-name> (<year>2007</year>) <article-title>Direct and indirect effects of climate and habitat factors on butterfly diversity</article-title>. <source xml:lang="en">Ecology</source>, <volume>88</volume>, <fpage>605</fpage>–<lpage>611</lpage>.<pub-id pub-id-type="pmid">17503588</pub-id></mixed-citation></ref><ref id="icad12193-bib-0043"><mixed-citation publication-type="journal" id="icad12193-cit-0043"><string-name><surname>Meyer</surname>, <given-names>C.L.</given-names></string-name> & <string-name><surname>Sisk</surname>, <given-names>T.D.</given-names></string-name> (<year>2001</year>) <article-title>Butterfly response to microclimatic conditions following ponderosa pine restoration</article-title>. <source xml:lang="en">Restoration Ecology</source>, <volume>9</volume>, <fpage>453</fpage>–<lpage>461</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0044"><mixed-citation publication-type="journal" id="icad12193-cit-0044"><string-name><surname>Milligan</surname>, <given-names>G.W.</given-names></string-name> & <string-name><surname>Cooper</surname>, <given-names>M.C.</given-names></string-name> (<year>1985</year>) <article-title>An examination of procedures for determining the number of clusters in a data set</article-title>. <source xml:lang="en">Psychometrika</source>, <volume>50</volume>, <fpage>159</fpage>–<lpage>179</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0045"><mixed-citation publication-type="journal" id="icad12193-cit-0045"><string-name><surname>Morisita</surname>, <given-names>M.</given-names></string-name> (<year>1959</year>) <article-title>Measuring interspecific association and similarity between communities. Memoirs of the faculty of Kyushu University</article-title>. <source xml:lang="en">Series E. Biology</source>, <volume>3</volume>, <fpage>36</fpage>–<lpage>80</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0046"><mixed-citation publication-type="journal" id="icad12193-cit-0046"><string-name><surname>Murphy</surname>, <given-names>S.M.</given-names></string-name> (<year>2004</year>) <article-title>Enemy‐free space maintains swallowtail butterfly host shift</article-title>. <source xml:lang="en">Proceedings of the National Academy of Science of USA</source>, <volume>101</volume>, <fpage>18048</fpage>–<lpage>18052</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0047"><mixed-citation publication-type="journal" id="icad12193-cit-0047"><string-name><surname>New</surname>, <given-names>T.R.</given-names></string-name>, <string-name><surname>Pyle</surname>, <given-names>R.M.</given-names></string-name>, <string-name><surname>Thomas</surname>, <given-names>J.A.</given-names></string-name>, <string-name><surname>Thomas</surname>, <given-names>C.D.</given-names></string-name> & <string-name><surname>Hammond</surname>, <given-names>P.C.</given-names></string-name> (<year>1995</year>) <article-title>Butterfly conservation management</article-title>. <source xml:lang="en">Annual Review Entomology</source>, <volume>40</volume>, <fpage>57</fpage>–<lpage>83</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0048"><mixed-citation publication-type="journal" id="icad12193-cit-0048"><string-name><surname>Poisot</surname>, <given-names>T.</given-names></string-name>, <string-name><surname>Thrall</surname>, <given-names>P.H.</given-names></string-name> & <string-name><surname>Hochberg</surname>, <given-names>M.E.</given-names></string-name> (<year>2011</year>) <article-title>Trophic network structure emerges through antagonistic coevolution in temporally varying environments</article-title>. <source xml:lang="en">Proceedings of the Royal Society B‐Biological Sciences</source>, <volume>279</volume>, <fpage>299</fpage>–<lpage>308</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0049"><mixed-citation publication-type="journal" id="icad12193-cit-0049"><string-name><surname>Polidori</surname>, <given-names>C.</given-names></string-name>, <string-name><surname>Santoro</surname>, <given-names>D.</given-names></string-name> & <string-name><surname>Blüthgen</surname>, <given-names>N.</given-names></string-name> (<year>2013</year>) <article-title>Does prey mobility affect niche width and individual specialization in hunting wasps? A network‐based analysis</article-title>. <source xml:lang="en">Oikos</source>, <volume>122</volume>, <fpage>385</fpage>–<lpage>394</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0050"><mixed-citation publication-type="book" id="icad12193-cit-0050"><string-name><surname>Pollard</surname>, <given-names>E.</given-names></string-name> & <string-name><surname>Yates</surname>, <given-names>T.J.</given-names></string-name> (<year>1993</year>) <source xml:lang="en">Monitoring Butterflies for Ecology and Conservation</source>. <publisher-name>Chapman & Hall</publisher-name>, <publisher-loc>London, UK</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0051"><mixed-citation publication-type="journal" id="icad12193-cit-0051"><string-name><surname>Poulin</surname>, <given-names>R.</given-names></string-name> (<year>2010</year>) <article-title>Network analysis shining light on parasite ecology and diversity</article-title>. <source xml:lang="en">Trends in Parasitology</source>, <volume>26</volume>, <fpage>492</fpage>–<lpage>498</lpage>.<pub-id pub-id-type="pmid">20561821</pub-id></mixed-citation></ref><ref id="icad12193-bib-0052"><mixed-citation publication-type="journal" id="icad12193-cit-0052"><string-name><surname>Prado</surname>, <given-names>P.I.</given-names></string-name> & <string-name><surname>Lewinsohn</surname>, <given-names>T.M.</given-names></string-name> (<year>2004</year>) <article-title>Compartments in insect‐plant associations and their consequences for community structure</article-title>. <source xml:lang="en">Journal of Animal Ecology</source>, <volume>73</volume>, <fpage>1168</fpage>–<lpage>1178</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0053"><mixed-citation publication-type="journal" id="icad12193-cit-0053"><string-name><surname>Pulliam</surname>, <given-names>H.R.</given-names></string-name> (<year>2000</year>) <article-title>On the relationship between niche and distribution</article-title>. <source xml:lang="en">Ecology Letters</source>, <volume>3</volume>, <fpage>349</fpage>–<lpage>361</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0054"><mixed-citation publication-type="book" id="icad12193-cit-0054"><collab collab-type="authors">R Development Core Team</collab>(<year>2011</year>) <source xml:lang="en">R: A Language and Environment for Statistical Computing</source>. <publisher-name>R Foundation for Statistical Computing</publisher-name>, <publisher-loc>Vienna, Austria</publisher-loc> ISBN 3‐900051‐07‐0. < <ext-link ext-link-type="uri" xlink:href="http://www.R-project.org">http://www.R-project.org</ext-link> >.</mixed-citation></ref><ref id="icad12193-bib-0055"><mixed-citation publication-type="journal" id="icad12193-cit-0055"><string-name><surname>Rosenfeld</surname>, <given-names>J.</given-names></string-name> (<year>2003</year>) <article-title>Assessing the habitat requirements of stream fishes: an overview and evaluation of different approaches</article-title>. <source xml:lang="en">Transaction of American Fisheries Society</source>, <volume>132</volume>, <fpage>953</fpage>–<lpage>968</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0056"><mixed-citation publication-type="journal" id="icad12193-cit-0056"><string-name><surname>Samways</surname>, <given-names>M.J.</given-names></string-name> & <string-name><surname>Lu</surname>, <given-names>S.S.</given-names></string-name> (<year>2007</year>) <article-title>Key traits in a threatened butterfly and its common sibling: implications for conservation</article-title>. <source xml:lang="en">Biodiversity and Conservation</source>, <volume>16</volume>, <fpage>4095</fpage>–<lpage>4107</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0057"><mixed-citation publication-type="journal" id="icad12193-cit-0057"><string-name><surname>Schroder</surname>, <given-names>G.D.</given-names></string-name> & <string-name><surname>Rosenzweig</surname>, <given-names>M.L.</given-names></string-name> (<year>1975</year>) <article-title>Perturbation analysis of competition and overlap in habitat utilization between <italic>Dipodomys ordii</italic> and <italic>Dipodomys merriami</italic></article-title>. <source xml:lang="en">Oecologia</source>, <volume>19</volume>, <fpage>9</fpage>–<lpage>28</lpage>.<pub-id pub-id-type="pmid">28308827</pub-id></mixed-citation></ref><ref id="icad12193-bib-0058"><mixed-citation publication-type="journal" id="icad12193-cit-0058"><string-name><surname>Shreeve</surname>, <given-names>T.G.</given-names></string-name> (<year>1984</year>) <article-title>Habitat selection, mate location, and microclimatic constraints on the activity of the speckled wood butterfly <italic>Pararge aegeria</italic></article-title>. <source xml:lang="en">Oikos</source>, <volume>42</volume>, <fpage>371</fpage>–<lpage>377</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0059"><mixed-citation publication-type="miscellaneous" id="icad12193-cit-0059"><string-name><surname>Skaug</surname>, <given-names>H.J.</given-names></string-name>, <string-name><surname>Fournier</surname>, <given-names>D.A.</given-names></string-name>, <string-name><surname>Nielsen</surname>, <given-names>A.</given-names></string-name>, <string-name><surname>Magnusson</surname>, <given-names>A.</given-names></string-name> & <string-name><surname>Bolker</surname>, <given-names>B.M.</given-names></string-name> (<year>2013</year>) <article-title>glmmADMB: generalized linear mixed models using AD Model Builder</article-title>. R package version 0.7.7. <<ext-link ext-link-type="uri" xlink:href="http://r-forge.r-project.org/projects/glmmadmb/">http://r-forge.r-project.org/projects/glmmadmb/</ext-link>>.</mixed-citation></ref><ref id="icad12193-bib-0060"><mixed-citation publication-type="book" id="icad12193-cit-0060"><string-name><surname>Slansky</surname>, <given-names>F.</given-names></string-name> (<year>1993</year>) <chapter-title>Nutritional ecology: the fundamental quests for nutrients</chapter-title><source xml:lang="en">Caterpillars – Ecological and Evolutionary Constraints on Foraging</source> (ed. by <person-group person-group-type="editor"><name name-style="western"><surname>Stamp</surname><given-names>N.E.</given-names></name></person-group> and <person-group person-group-type="editor"><name name-style="western"><surname>Casey</surname><given-names>T.E.</given-names></name></person-group>), pp. <fpage>29</fpage>–<lpage>91</lpage>. <publisher-name>Chapman & Hall</publisher-name>, <publisher-loc>New York City, New York</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0061"><mixed-citation publication-type="journal" id="icad12193-cit-0061"><string-name><surname>Spitzer</surname>, <given-names>K.</given-names></string-name>, <string-name><surname>Novotny</surname>, <given-names>V.</given-names></string-name>, <string-name><surname>Tonner</surname>, <given-names>M.</given-names></string-name> & <string-name><surname>Leps</surname>, <given-names>J.</given-names></string-name> (<year>1993</year>) <article-title>Habitat preferences, distribution and seasonality of the butterflies (Lepidoptera, Papilionoidea) in a mountain tropical rain forest, Vietnam</article-title>. <source xml:lang="en">Journal of Biogeography</source>, <volume>20</volume>, <fpage>109</fpage>–<lpage>121</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0062"><mixed-citation publication-type="journal" id="icad12193-cit-0062"><string-name><surname>Stefanescu</surname>, <given-names>C.</given-names></string-name>, <string-name><surname>Carnicer</surname>, <given-names>J.</given-names></string-name> & <string-name><surname>Peñuelas</surname>, <given-names>J.</given-names></string-name> (<year>2011</year>) <article-title>Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change</article-title>. <source xml:lang="en">Ecography</source>, <volume>34</volume>, <fpage>353</fpage>–<lpage>363</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0063"><mixed-citation publication-type="journal" id="icad12193-cit-0063"><string-name><surname>Stefanescu</surname>, <given-names>C.</given-names></string-name>, <string-name><surname>Herrando</surname>, <given-names>S.</given-names></string-name> & <string-name><surname>Paramo</surname>, <given-names>F.</given-names></string-name> (<year>2004</year>) <article-title>Butterfly species richness in the north‐west mediterranean basin: the role of natural and human‐induced factors</article-title>. <source xml:lang="en">Journal of Biogeography</source>, <volume>31</volume>, <fpage>905</fpage>–<lpage>915</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0064"><mixed-citation publication-type="book" id="icad12193-cit-0064"><string-name><surname>Van Swaay</surname>, <given-names>C.</given-names></string-name>, <string-name><surname>Cuttelod</surname>, <given-names>A.</given-names></string-name>, <string-name><surname>Collins</surname>, <given-names>S.</given-names></string-name>, <string-name><surname>Maes</surname>, <given-names>D.</given-names></string-name>, <string-name><surname>López Munguira</surname>, <given-names>M.</given-names></string-name>, <string-name><surname>Šašić</surname>, <given-names>M.</given-names></string-name>, <string-name><surname>Settele</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>Verovnik</surname>, <given-names>R.</given-names></string-name>, <string-name><surname>Verstrael</surname>, <given-names>T.</given-names></string-name>, <string-name><surname>Warren</surname>, <given-names>M.</given-names></string-name>, <string-name><surname>Wiemers</surname>, <given-names>M.</given-names></string-name> & <string-name><surname>Wynhoff</surname>, <given-names>I.</given-names></string-name> (<year>2010</year>) <source xml:lang="en">European Red List of Butterflies</source>. <publisher-name>Publications Office of the European Union</publisher-name>, <publisher-loc>Luxembourg</publisher-loc>, 47 <<ext-link ext-link-type="uri" xlink:href="http://ec.europa.eu/environment/nature/conservation/species/redlist/downloads/European_butterflies.pdf">http://ec.europa.eu/environment/nature/conservation/species/redlist/downloads/European_butterflies.pdf</ext-link>>.</mixed-citation></ref><ref id="icad12193-bib-0065"><mixed-citation publication-type="book" id="icad12193-cit-0065"><string-name><surname>Van Swaay</surname>, <given-names>C.A.M.</given-names></string-name>, <string-name><surname>Van Strien</surname>, <given-names>A.J.</given-names></string-name>, <string-name><surname>Harpke</surname>, <given-names>A.</given-names></string-name>, <string-name><surname>Fontaine</surname>, <given-names>B.</given-names></string-name>, <string-name><surname>Stefanescu</surname>, <given-names>C.</given-names></string-name>, <string-name><surname>Roy</surname>, <given-names>D.</given-names></string-name>, <string-name><surname>Maes</surname>, <given-names>D.</given-names></string-name>, <string-name><surname>Kühn</surname>, <given-names>E.</given-names></string-name>, <string-name><surname>Õunap</surname>, <given-names>E.</given-names></string-name>, <string-name><surname>Regan</surname>, <given-names>E.</given-names></string-name>, <string-name><surname>Švitra</surname>, <given-names>G.</given-names></string-name>, <string-name><surname>Heliölä</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>Settele</surname>, <given-names>J.</given-names></string-name>, <string-name><surname>Pettersson</surname>, <given-names>L.B.</given-names></string-name>, <string-name><surname>Botham</surname>, <given-names>M.</given-names></string-name>, <string-name><surname>Musche</surname>, <given-names>M.</given-names></string-name>, <string-name><surname>Titeux</surname>, <given-names>N.</given-names></string-name>, <string-name><surname>Cornish</surname>, <given-names>N.</given-names></string-name>, <string-name><surname>Leopold</surname>, <given-names>P.</given-names></string-name>, <string-name><surname>Julliard</surname>, <given-names>R.</given-names></string-name>, <string-name><surname>Verovnik</surname>, <given-names>R.</given-names></string-name>, <string-name><surname>Öberg</surname>, <given-names>S.</given-names></string-name>, <string-name><surname>Popov</surname>, <given-names>S.</given-names></string-name>, <string-name><surname>Collins</surname>, <given-names>S.</given-names></string-name>, <string-name><surname>Goloshchapova</surname>, <given-names>S.</given-names></string-name>, <string-name><surname>Roth</surname>, <given-names>T.</given-names></string-name>, <string-name><surname>Brereton</surname>, <given-names>T.</given-names></string-name> & <string-name><surname>Warren</surname>, <given-names>M.S.</given-names></string-name> (<year>2013</year>) <article-title>The European Grassland Butterfly Indicator: 1990–2011</article-title>. EEA Technical Report, European Environment Agency. <publisher-name>Publications Office of the European Union</publisher-name>, <publisher-loc>Luxembourg</publisher-loc>.</mixed-citation></ref><ref id="icad12193-bib-0066"><mixed-citation publication-type="journal" id="icad12193-cit-0066"><string-name><surname>Wagner</surname>, <given-names>D.L.</given-names></string-name> & <string-name><surname>Van Driesche</surname>, <given-names>R.G.</given-names></string-name> (<year>2010</year>) <article-title>Threats posed to rare or endangered insects by invasion of nonnative species</article-title>. <source xml:lang="en">Annual Review of Entomology</source>, <volume>55</volume>, <fpage>547</fpage>–<lpage>568</lpage>.</mixed-citation></ref><ref id="icad12193-bib-0067"><mixed-citation publication-type="journal" id="icad12193-cit-0067"><string-name><surname>Waltz</surname>, <given-names>A.E.M.</given-names></string-name> & <string-name><surname>Covington</surname>, <given-names>W.W.</given-names></string-name> (<year>2004</year>) <article-title>Ecological restoration treatments increase butterfly richness and abundance: mechanisms of response</article-title>. <source xml:lang="en">Restoration Ecology</source>, <volume>12</volume>, <fpage>85</fpage>–<lpage>96</lpage>.</mixed-citation></ref></ref-list></back></pmc></record>Pour manipuler ce document sous Unix (Dilib)
EXPLOR_STEP=$WICRI_ROOT/Sante/explor/StressCovidV1/Data/Pmc/Corpus
HfdSelect -h $EXPLOR_STEP/biblio.hfd -nk 0006779 | SxmlIndent | more
Ou
HfdSelect -h $EXPLOR_AREA/Data/Pmc/Corpus/biblio.hfd -nk 0006779 | SxmlIndent | more
Pour mettre un lien sur cette page dans le réseau Wicri
{{Explor lien
|wiki= Sante
|area= StressCovidV1
|flux= Pmc
|étape= Corpus
|type= RBID
|clé=
|texte=
}}
| This area was generated with Dilib version V0.6.33. |  |