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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">CLIMBER: Climatic niche characteristics of the butterflies in Europe</title>
<author><name sortKey="Schweiger, Oliver" sort="Schweiger, Oliver" uniqKey="Schweiger O" first="Oliver" last="Schweiger">Oliver Schweiger</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Harpke, Alexander" sort="Harpke, Alexander" uniqKey="Harpke A" first="Alexander" last="Harpke">Alexander Harpke</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wiemers, Martin" sort="Wiemers, Martin" uniqKey="Wiemers M" first="Martin" last="Wiemers">Martin Wiemers</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Settele, Josef" sort="Settele, Josef" uniqKey="Settele J" first="Josef" last="Settele">Josef Settele</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">iDiv, German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, 04103 Leipzig, Germany</nlm:aff>
</affiliation>
</author>
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<idno type="pmid">24478578</idno>
<idno type="pmc">3904140</idno>
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<idno type="RBID">PMC:3904140</idno>
<idno type="doi">10.3897/zookeys.367.6185</idno>
<date when="2014">2014</date>
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<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">CLIMBER: Climatic niche characteristics of the butterflies in Europe</title>
<author><name sortKey="Schweiger, Oliver" sort="Schweiger, Oliver" uniqKey="Schweiger O" first="Oliver" last="Schweiger">Oliver Schweiger</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Harpke, Alexander" sort="Harpke, Alexander" uniqKey="Harpke A" first="Alexander" last="Harpke">Alexander Harpke</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wiemers, Martin" sort="Wiemers, Martin" uniqKey="Wiemers M" first="Martin" last="Wiemers">Martin Wiemers</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Settele, Josef" sort="Settele, Josef" uniqKey="Settele J" first="Josef" last="Settele">Josef Settele</name>
<affiliation><nlm:aff id="A1">Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">iDiv, German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, 04103 Leipzig, Germany</nlm:aff>
</affiliation>
</author>
</analytic>
<series><title level="j">ZooKeys</title>
<idno type="ISSN">1313-2989</idno>
<idno type="eISSN">1313-2970</idno>
<imprint><date when="2014">2014</date>
</imprint>
</series>
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<front><div type="abstract" xml:lang="en"><label>Abstract</label>
<p>Detailed information on species’ ecological niche characteristics that can be related to declines and extinctions is indispensable for a better understanding of the relationship between the occurrence and performance of wild species and their environment and, moreover, for an improved assessment of the impacts of global change. Knowledge on species characteristics such as habitat requirements is already available in the ecological literature for butterflies, but information about their climatic requirements is still lacking. Here we present a unique dataset on the climatic niche characteristics of 397 European butterflies representing 91% of the European species (see Appendix). These characteristics were obtained by combining detailed information on butterfly distributions in Europe (which also led to the ‘Distribution Atlas of Butterflies in Europe’) and the corresponding climatic conditions. The presented dataset comprises information for the position and breadth of the following climatic niche characteristics: mean annual temperature, range in annual temperature, growing degree days, annual precipitation sum, range in annual precipitation and soil water content. The climatic niche position is indicated by the median and mean value for each climate variable across a species’ range, accompanied by the 95% confidence interval for the mean and the number of grid cells used for calculations. Climatic niche breadth is indicated by the standard deviation and the minimum and maximum values for each climatic variable across a species’ range. Database compilation was based on high quality standards and the data are ready to use for a broad range of applications.</p>
<pmc-comment>PageBreak</pmc-comment>
<p>It is already evident that the information provided in this dataset is of great relevance for basic and applied ecology. Based on the species temperature index (STI, i.e. the mean temperature value per species), the community temperature index (CTI, i.e. the average STI value across the species in a community) was recently adopted as an indicator of climate change impact on biodiversity by the pan-European framework supporting the Convention on Biological Diversity (Streamlining European Biodiversity Indicators 2010) and has already been used in several scientific publications. The application potential of this database ranges from theoretical aspects such as assessments of past niche evolution or analyses of trait interdependencies to the very applied aspects of measuring, monitoring and projecting historical, ongoing and potential future responses to climate change using butterflies as an indicator.</p>
</div>
</front>
<back><div1 type="bibliography"><listBibl></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">Zookeys</journal-id>
<journal-id journal-id-type="iso-abbrev">Zookeys</journal-id>
<journal-id journal-id-type="publisher-id">ZooKeys</journal-id>
<journal-title-group><journal-title>ZooKeys</journal-title>
</journal-title-group>
<issn pub-type="ppub">1313-2989</issn>
<issn pub-type="epub">1313-2970</issn>
<publisher><publisher-name>Pensoft Publishers</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">24478578</article-id>
<article-id pub-id-type="pmc">3904140</article-id>
<article-id pub-id-type="doi">10.3897/zookeys.367.6185</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Article</subject>
</subj-group>
</article-categories>
<title-group><article-title>CLIMBER: Climatic niche characteristics of the butterflies in Europe</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Schweiger</surname>
<given-names>Oliver</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Harpke</surname>
<given-names>Alexander</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wiemers</surname>
<given-names>Martin</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Settele</surname>
<given-names>Josef</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
</contrib-group>
<aff id="A1"><label>1</label>
Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</aff>
<aff id="A2"><label>2</label>
iDiv, German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, 04103 Leipzig, Germany</aff>
<author-notes><corresp>Corresponding author: Oliver Schweiger (<email xlink:type="simple">oliver.schweiger@ufz.de</email>
)</corresp>
<fn fn-type="edited-by"><p>Academic editor: V. Chavan</p>
</fn>
</author-notes>
<pub-date pub-type="collection"><year>2014</year>
</pub-date>
<pub-date pub-type="epub"><day>6</day>
<month>1</month>
<year>2014</year>
</pub-date>
<issue>367</issue>
<fpage>65</fpage>
<lpage>84</lpage>
<history><date date-type="received"><day>2</day>
<month>9</month>
<year>2013</year>
</date>
<date date-type="accepted"><day>28</day>
<month>11</month>
<year>2013</year>
</date>
</history>
<permissions><copyright-statement>Oliver Schweiger, Alexander Harpke, Martin Wiemers, Josef Settele</copyright-statement>
<license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0"><license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<abstract><label>Abstract</label>
<p>Detailed information on species’ ecological niche characteristics that can be related to declines and extinctions is indispensable for a better understanding of the relationship between the occurrence and performance of wild species and their environment and, moreover, for an improved assessment of the impacts of global change. Knowledge on species characteristics such as habitat requirements is already available in the ecological literature for butterflies, but information about their climatic requirements is still lacking. Here we present a unique dataset on the climatic niche characteristics of 397 European butterflies representing 91% of the European species (see Appendix). These characteristics were obtained by combining detailed information on butterfly distributions in Europe (which also led to the ‘Distribution Atlas of Butterflies in Europe’) and the corresponding climatic conditions. The presented dataset comprises information for the position and breadth of the following climatic niche characteristics: mean annual temperature, range in annual temperature, growing degree days, annual precipitation sum, range in annual precipitation and soil water content. The climatic niche position is indicated by the median and mean value for each climate variable across a species’ range, accompanied by the 95% confidence interval for the mean and the number of grid cells used for calculations. Climatic niche breadth is indicated by the standard deviation and the minimum and maximum values for each climatic variable across a species’ range. Database compilation was based on high quality standards and the data are ready to use for a broad range of applications.</p>
<pmc-comment>PageBreak</pmc-comment>
<p>It is already evident that the information provided in this dataset is of great relevance for basic and applied ecology. Based on the species temperature index (STI, i.e. the mean temperature value per species), the community temperature index (CTI, i.e. the average STI value across the species in a community) was recently adopted as an indicator of climate change impact on biodiversity by the pan-European framework supporting the Convention on Biological Diversity (Streamlining European Biodiversity Indicators 2010) and has already been used in several scientific publications. The application potential of this database ranges from theoretical aspects such as assessments of past niche evolution or analyses of trait interdependencies to the very applied aspects of measuring, monitoring and projecting historical, ongoing and potential future responses to climate change using butterflies as an indicator.</p>
</abstract>
<kwd-group><label>Keywords</label>
<kwd>Climate change</kwd>
<kwd>climate warming</kwd>
<kwd>CTI</kwd>
<kwd>global change</kwd>
<kwd>global warming</kwd>
<kwd>modelling</kwd>
<kwd>risk</kwd>
<kwd>trend</kwd>
<kwd>STI</kwd>
<kwd>Europe</kwd>
<kwd>butterflies</kwd>
<kwd>Lepidoptera</kwd>
<kwd>Papilionidae</kwd>
<kwd>Pieridae</kwd>
<kwd>Lycaenidae</kwd>
<kwd>Riodinidae</kwd>
<kwd>Nymphalidae</kwd>
<kwd>Hesperiidae</kwd>
</kwd-group>
</article-meta>
<notes><sec sec-type="Citation"><title>Citation</title>
<p>Schweiger O, Harpke A, Wiemers M, Settele J (2014) CLIMBER: Climatic niche characteristics of the butterflies in Europe. ZooKeys 367: 65–84. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.3897/zookeys.367.6185">10.3897/zookeys.367.6185</ext-link>
Resource ID: GBIF key: <ext-link ext-link-type="uri" xlink:href="http://www.gbif.org/dataset/e2bcea8c-dfea-475e-a4ae-af282b4ea1c5">http://www.gbif.org/dataset/e2bcea8c-dfea-475e-a4ae-af282b4ea1c5</ext-link>
</p>
</sec>
<sec sec-type="Resource Citation"><title>Resource Citation:</title>
<p>Helmholtz Centre for Environmental Research - UFZ (2013). CLIMBER: Climatic niche characteristics of the butterflies in Europe. 397 records, Online at <ext-link ext-link-type="uri" xlink:href="http://ipt.pensoft.net/ipt/resource.do?r=climber">http://ipt.pensoft.net/ipt/resource.do?r=climber</ext-link>
, version 1.3 (released on 3/12/2013), Resource ID: GBIF key: <ext-link ext-link-type="uri" xlink:href="http://www.gbif.org/dataset/e2bcea8c-dfea-475e-a4ae-af282b4ea1c5">http://www.gbif.org/dataset/e2bcea8c-dfea-475e-a4ae-af282b4ea1c5</ext-link>
, Data Paper ID: doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.3897/zookeys.367.6185">10.3897/zookeys.367.6185</ext-link>
</p>
</sec>
</notes>
</front>
<body><sec><title>Introduction</title>
<p>Global change seriously threatens biodiversity at all organisational levels ranging from genetic diversity, performance and occurrence of single species, taxonomic, phylogenetic and functional diversity of communities and species assemblages to properties of whole ecosystems including the provision of ecosystem services for human well-being (<xref rid="B9" ref-type="bibr">Lavergne et al. 2010</xref>
; <xref rid="B11" ref-type="bibr">Parmesan 2006</xref>
; <xref rid="B13" ref-type="bibr">Potts et al. 2010</xref>
; <xref rid="B16" ref-type="bibr">Schröter et al. 2005</xref>
). But species are not equally at risk when facing global change (e.g. <xref rid="B40" ref-type="bibr">Settele et al. 2008</xref>
). In the context of climate change, several species-specific ecological characteristics have been identified to determine vulnerability, including diets, habitat requirements, ecological specialisation and plasticity and the ecological characteristics of interacting species (<xref rid="B4" ref-type="bibr">Heikkinen et al. 2010</xref>
; <xref rid="B14" ref-type="bibr">Pöyry et al. 2009</xref>
; <xref rid="B48" ref-type="bibr">Schweiger et al. 2012</xref>
; <xref rid="B21" ref-type="bibr">Visser 2008</xref>
; <xref rid="B22" ref-type="bibr">Warren et al. 2001</xref>
). Thus, good knowledge of the ecological characteristics relevant for the reaction of species and communities to particular drivers of global change is needed, which can then be utilised as powerful indicators for conservation planning and action.</p>
<p>One of the most important ecological characteristics to assess how species react to climate change obviously is the climatic niche. While knowledge on particular species characteristics such as habitat requirements is already available for some species groups, crucial publicly available information about climatic requirements is still lacking for the majority of the species. Here we present a unique dataset on climatic niche characteristics of 397 (91%) butterfly species in Europe, which have been shown to be particularly sensitive to changing climates (<xref rid="B6" ref-type="bibr">Hill et al. 2002</xref>
; <xref rid="B40" ref-type="bibr">Settele et al. 2008</xref>
; <xref rid="B22" ref-type="bibr">Warren et al. 2001</xref>
). Based on projections of future suitable climatic conditions, <xref rid="B40" ref-type="bibr">Settele et al. (2008)</xref>
showed that under the assumption of unlimited dispersal 7% of the European butterflies are at an extremely high or very high risk (i.e. a loss of more than 95% and 85%, respectively of their current range size until 2080), 6% are at high risk (>70% loss) and 18% are at risk (>50% loss; <xref ref-type="fig" rid="F1">Fig. 1</xref>
). However, the more realistic assumption of no dispersal (in the given amount of time) projected 33% of the butterflies to be at an extremely high or very high risk, 26% to be at high risk and 19% to be at risk (<xref ref-type="fig" rid="F1">Fig. 1</xref>
).</p>
<fig id="F1" orientation="portrait" position="float"><label>Figure 1.</label>
<caption><p>Proportion of species (%) with different climatic risk status after <xref rid="B40" ref-type="bibr">Settele at al. (2008)</xref>
assuming full dispersal (<bold>a</bold>
) and no dispersal capacity (<bold>b</bold>
).</p>
</caption>
<graphic xlink:href="ZooKeys-367-065-g001"></graphic>
</fig>
<pmc-comment>PageBreak</pmc-comment>
<p>Based on detailed data on the distribution of European butterflies, which also led to the ‘Distribution Atlas of European Butterflies’ (<xref rid="B33" ref-type="bibr">Kudrna 2002</xref>
), the ‘Climatic Risk Atlas of European Butterflies (<xref rid="B40" ref-type="bibr">Settele et al. 2008</xref>
) and the ‘Distribution Atlas of Butterflies in Europe’ (<xref rid="B34" ref-type="bibr">Kudrna et al. 2011</xref>
), we extracted measures of climatic conditions (indicating niche breadth and position) within the distributional range of each species. As a consequence of this approach, users of this dataset should be aware that the provided measures refer to the realised climatic niche and not to the fundamental niche (sensu <xref rid="B7" ref-type="bibr">Hutchinson 1957</xref>
; but see discussion in <xref rid="B1" ref-type="bibr">Araújo and Guisan 2006</xref>
). The extracted measures reflect two primary properties of climate, energy and water, which are known to affect butterfly species performance and distributions as a consequence of physiological limitations (<xref rid="B3" ref-type="bibr">Buckley et al. 2011</xref>
; <xref rid="B15" ref-type="bibr">Roy et al. 2001</xref>
). Most of these measures are quite independent from each other and cover different aspects of the climatic niche (<xref ref-type="fig" rid="F2">Fig. 2</xref>
).</p>
<fig id="F2" orientation="portrait" position="float"><label>Figure 2.</label>
<caption><p>Results from a principal component analysis of the species-specific mean values of six different climate variables. Mean values per species have been calculated based on the observed records per 50 km × 50 km CGRS grid cell across a species’ European distribution. PC1 explained 58% and PC2 32% of the variability. Niche characteristics according to annual temperature (temp) and growing degree days until August (gdd) are highly correlated. Also, the two measures of water availability, annual precipitation (pre) and soil water content (swc) show some similarity, while the indicators of annual range in precipitation (pre.range) and temperature (temp.range) are negatively correlated. In spite of these similarities, aspects of energy, water and their annual variability can be assessed independently with a choice of at least three of the indicators.</p>
</caption>
<graphic xlink:href="ZooKeys-367-065-g002"></graphic>
</fig>
<p>By combining a comprehensive database on the distribution of European butterflies with publicly available climatic data in combination with a constantly high level of quality control at crucial steps of the data generation, CLIMBER represents a unique and ready-to-use dataset for a broad variety of potential applications. Analysis of phylogenetic signals in the climatic niche characteristics, for instance, can be used to assess past niche evolution which can lead to projections of potential future risks in the face of rapid climate change (for a comparable analysis for birds see <xref rid="B8" ref-type="bibr">Lavergne et al. 2013</xref>
). Also, analyses relating climatic niche properties to other species traits can be helpful to assess interdependencies of different ecologi<pmc-comment>PageBreak</pmc-comment>
cal characteristics, as has been done recently for birds and their temperature and habitat preferences (<xref rid="B2" ref-type="bibr">Barnagaud et al. 2012</xref>
). So far the most powerful application of climatic niche characteristics provided in this dataset comes from the ‘species temperature index’ (STI). The STI is simply the mean temperature value per species across its range. Based on the STI, the ‘community temperature index’ (CTI) has been suggested as a powerful and robust tool to measure the response of local communities to temperature change (<xref rid="B26" ref-type="bibr">Devictor et al. 2008</xref>
; <xref rid="B46" ref-type="bibr">Devictor et al. 2012a</xref>
; <xref rid="B47" ref-type="bibr">Devictor et al. 2012b</xref>
). The CTI is calculated as the average STI value across the species or specimens in a community and has been used to analyse the temporal response to climate warming of local bird and butterfly communities across Europe. One striking result of this study was the detection of time lag effects in the community response to climate warming and that these lag effects differed between the two species groups (<xref rid="B46" ref-type="bibr">Devictor et al. 2012a</xref>
).</p>
<p>STI values for European butterflies can be of great value for governmental and non-governmental conservation organisations (<xref rid="B49" ref-type="bibr">Van Swaay et al. 2010</xref>
; <xref rid="B50" ref-type="bibr">Van Swaay et al. 2008</xref>
). Based on the STI, the CTI was recently adopted as an indicator of climate <pmc-comment>PageBreak</pmc-comment>
change impact on biodiversity by the pan-European framework supporting the Convention on Biological Diversity (Streamlining European Biodiversity Indicators 2010; <ext-link ext-link-type="uri" xlink:href="http://ec.europa.eu/environment/nature/knowledge/eu2010_indicators">http://ec.europa.eu/environment/nature/knowledge/eu2010_indicators</ext-link>
). Thus, STI and corresponding CTI values can perfectly complement and enrich the analysis of all kind of butterfly monitoring schemes. To address the fact that temperature is not the only changing climatic factor or aspect of the climatic niche, we think that the additionally provided climatic niche characteristics concerning water availability and annual climatic variability can help to enrich the landscape of target-specific analyses and indicators (<xref ref-type="fig" rid="F2">Fig. 2</xref>
). By providing public access to this dataset, we hope to contribute to improvements of the scientific understanding of how climate change affects species and communities and to improve monitoring and conservation actions for climate change mitigation.</p>
</sec>
<sec><title>Metadata</title>
<p>For the description of the metadata we followed the standards suggested by <xref rid="B10" ref-type="bibr">Michener et al. (1997)</xref>
in a slightly modified way.</p>
</sec>
<sec><title>Title</title>
<p>CLIMBER: Climatic niche characteristics of the butterflies in Europe</p>
</sec>
<sec><title>Contributors</title>
<sec sec-type="Dataset owner"><title>Dataset owner</title>
<p>Oliver Schweiger, Alexander Harpke, Martin Wiemers, Josef Settele</p>
<p>Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Strasse 4, 06120 Halle, Germany</p>
</sec>
<sec sec-type="Contact person"><title>Contact person</title>
<p>Oliver Schweiger</p>
<p>Affiliation: Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology</p>
<p>Address: Theodor-Lieser-Strasse 4, 06120 Halle, Germany</p>
<p>Phone: +49 345 558 5306</p>
<p>Email: oliver.schweiger@ufz.de</p>
<pmc-comment>PageBreak</pmc-comment>
</sec>
</sec>
<sec><title>Geographic, temporal and taxonomic coverage</title>
<sec sec-type="Geographic coverage and spatial resolution"><title>Geographic coverage and spatial resolution</title>
<p>Climatic niche characteristics are provided for all butterfly species occurring within a European window of 11°W, 32°E longitude and 34°N, 72°N latitude (<xref ref-type="fig" rid="F3">Fig. 3</xref>
). Resolution of butterfly distribution and corresponding climate data used to calculate climatic niche characteristics corresponds to the 50 km × 50 km Common European Chorological Grid Reference System (CGRS; <ext-link ext-link-type="uri" xlink:href="http://www.eea.europa.eu/data-and-maps/data/common-european-chorological-grid-reference-system-cgrs">http://www.eea.europa.eu/data-and-maps/data/common-european-chorological-grid-reference-system-cgrs</ext-link>
).</p>
<fig id="F3" orientation="portrait" position="float"><label>Figure 3.</label>
<caption><p>Geographic coverage used for the calculation of the climatic species characteristics. Purple dots indicate 50 km × 50 km CGRS grid cells with available species records.</p>
</caption>
<graphic xlink:href="ZooKeys-367-065-g003"></graphic>
</fig>
<p>The geographic window excludes data from the Atlantic islands under European administration (the Azores, Madeira and Canary Islands) as well as Cyprus and Iceland. Due to low levels of recording, data from Belarus, Ukraine, Moldova, and Russia were also excluded. Additionally, no climate data were available for two species with extremely local distributions on the Pontine Islands and the Greek island of Nissiros. These restrictions led to the exclusion of 38 of the European butterfly species listed in <xref rid="B34" ref-type="bibr">Kudrna et al. (2011)</xref>
, but confined to these regions (<xref ref-type="table" rid="T1">Table 1</xref>
).</p>
<table-wrap id="T1" orientation="portrait" position="float"><label>Table 1.</label>
<caption><p>Species occurring in Europe and listed in <xref rid="B34" ref-type="bibr">Kudrna et al. (2011)</xref>
but not considered for the assignment of climatic niche characteristics in this database.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Species</th>
<th rowspan="1" colspan="1">European range</th>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Azanus ubaldus</named-content>
</italic>
(Stoll, 1782)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Catopsilia florella</named-content>
</italic>
(Fabricius, 1775)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Chazara persephone</named-content>
</italic>
(Hübner, [1805])</td>
<td rowspan="1" colspan="1">Ukraine</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Chilades galba</named-content>
</italic>
(Lederer, 1855)</td>
<td rowspan="1" colspan="1">Cyprus</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Cigaritis acamas</named-content>
</italic>
(Klug, 1834)</td>
<td rowspan="1" colspan="1">Cyprus</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Cyclyrius webbianus</named-content>
</italic>
(Brulle, 1839)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Euchloe eversi</named-content>
</italic>
Stamm, 1963</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Euchloe grancanariensis</named-content>
</italic>
Acosta, 2008</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Euchloe hesperidum</named-content>
</italic>
Rothschild, 1913</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Glaucopsyche paphos</named-content>
</italic>
Chapman, 1920</td>
<td rowspan="1" colspan="1">Cyprus</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Gonepteryx cleobule</named-content>
</italic>
(Hübner, 1825)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Gonepteryx eversi</named-content>
</italic>
Rehnelt, 1974</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Gonepteryx maderensis</named-content>
</italic>
Felder, 1863</td>
<td rowspan="1" colspan="1">Madeira</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Gonepteryx palmae</named-content>
</italic>
Stamm, 1963</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia azorina</named-content>
</italic>
(Strecker, 1899)</td>
<td rowspan="1" colspan="1">Azores</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia bacchus</named-content>
</italic>
Higgins, 1967</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia cypriensis</named-content>
</italic>
(Holik, 1949)</td>
<td rowspan="1" colspan="1">Cyprus</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia gomera</named-content>
</italic>
Higgins, 1967</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia maderensis</named-content>
</italic>
(Bethune-Baker, 1891)</td>
<td rowspan="1" colspan="1">Madeira</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia sbordonii</named-content>
</italic>
Kudrna, 1984</td>
<td rowspan="1" colspan="1">Pontine Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia tamadabae</named-content>
</italic>
Owen & Smith, 1992</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia tilosi</named-content>
</italic>
(Manil, 1984)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hipparchia wyssii</named-content>
</italic>
(Christ, 1889)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Hypolimnas misippus</named-content>
</italic>
(Linnaeus, 1764)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Maniola cypricola</named-content>
</italic>
(Graves, 1928)</td>
<td rowspan="1" colspan="1">Cyprus</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Maniola halicarnassus</named-content>
</italic>
Thomas, 1990</td>
<td rowspan="1" colspan="1">Nissiros Island</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Neolycaena rhymnus</named-content>
</italic>
(Eversmann, 1832)</td>
<td rowspan="1" colspan="1">Ukraine</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pararge xiphia</named-content>
</italic>
(Fabricius, 1775)</td>
<td rowspan="1" colspan="1">Madeira</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pararge xiphioides</named-content>
</italic>
Staudinger, 1871</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pieris cheiranthi</named-content>
</italic>
(Hübner, 1808)</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pieris wollastoni</named-content>
</italic>
Butler, 1866</td>
<td rowspan="1" colspan="1">Madeira</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Polyommatus corydonius</named-content>
</italic>
(Herrich-Schäffer, 1852)</td>
<td rowspan="1" colspan="1">Ukraine</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Polyommatus damocles</named-content>
</italic>
(Herrich-Schäffer, 1844)</td>
<td rowspan="1" colspan="1">Ukraine</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Polyommatus damone</named-content>
</italic>
(Eversmann, 1841)</td>
<td rowspan="1" colspan="1">Ukraine</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pseudochazara euxina</named-content>
</italic>
(Kusnezov, 1909)</td>
<td rowspan="1" colspan="1">Ukraine</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Thymelicus christi</named-content>
</italic>
Rebel, 1894</td>
<td rowspan="1" colspan="1">Canary Islands</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Tomares callimachus</named-content>
</italic>
(Eversmann, 1848)</td>
<td rowspan="1" colspan="1">Ukraine</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Vanessa vulcania</named-content>
</italic>
(Godart, 1819)</td>
<td rowspan="1" colspan="1">Canary Islands & Madeira</td>
</tr>
</tbody>
</table>
</table-wrap>
<pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
</sec>
<sec sec-type="Temporal reference period"><title>Temporal reference period</title>
<p>Only butterfly distribution data from the period of 1981 to 2000 were considered due to low sampling intensity in earlier periods (<xref ref-type="fig" rid="F4">Fig. 4</xref>
) and to minimize errors due to ongoing range shifts as a response to recent climate change.</p>
<fig id="F4" orientation="portrait" position="float"><label>Figure 4.</label>
<caption><p>Temporal availability of records and corresponding sampling intensity. Only the period of 1981–2000 has been considered in CLIMBER.</p>
</caption>
<graphic xlink:href="ZooKeys-367-065-g004"></graphic>
</fig>
</sec>
</sec>
<sec><title>Taxonomy</title>
<sec sec-type="Taxonomic ranks"><title>Taxonomic ranks</title>
<p><bold>Phylum:</bold>
<named-content content-type="taxon-name">Arthropoda</named-content>
</p>
<p><bold>Subphylum:</bold>
<named-content content-type="taxon-name">Hexapoda</named-content>
</p>
<p><bold>Class:</bold>
<named-content content-type="taxon-name">Insecta</named-content>
</p>
<p><bold>Order:</bold>
<named-content content-type="taxon-name">Lepidoptera</named-content>
</p>
<p><bold>Superfamily:</bold>
<named-content content-type="taxon-name">Papilionoidea</named-content>
(sensu <xref rid="B37" ref-type="bibr">Regier et al. 2013</xref>
; <xref rid="B44" ref-type="bibr">Wahlberg et al. 2013</xref>
)</p>
<p><bold>Families:</bold>
<named-content content-type="taxon-name">Hesperiidae</named-content>
, <named-content content-type="taxon-name">Lycaenidae</named-content>
, <named-content content-type="taxon-name">Nymphalidae</named-content>
, <named-content content-type="taxon-name">Papilionidae</named-content>
, <named-content content-type="taxon-name">Pieridae</named-content>
, <named-content content-type="taxon-name">Riodinidae</named-content>
</p>
<p><bold>Common name:</bold>
butterflies</p>
</sec>
</sec>
<sec><title>Taxonomic coverage</title>
<p>The taxonomic coverage spans all butterfly species within the selected geographic window (397 species) and represents 91% of all European species (<xref ref-type="fig" rid="F5">Fig. 5</xref>
). Thirty-eight <pmc-comment>PageBreak</pmc-comment>
species from less well sampled Eastern European countries, Atlantic and small Mediterranean islands have not been considered (<xref ref-type="fig" rid="F5">Fig. 5a</xref>
). The taxonomy of European butterfly species follows <xref rid="B34" ref-type="bibr">Kudrna et al. (2011)</xref>
. Erroneous use of brackets around authors’ names was corrected in 15 cases (cf. <xref rid="B43" ref-type="bibr">Tshikolovets 2011</xref>
; <xref ref-type="table" rid="T2">Table 2</xref>
).</p>
<fig id="F5" orientation="portrait" position="float"><label>Figure 5.</label>
<caption><p>Taxonomic coverage according to the entire European butterfly fauna (<bold>a</bold>
) and families (<bold>b</bold>
). Values indicate number of species.</p>
</caption>
<graphic xlink:href="ZooKeys-367-065-g005"></graphic>
</fig>
<table-wrap id="T2" orientation="portrait" position="float"><label>Table 2.</label>
<caption><p>Corrected species names (cf. <xref rid="B43" ref-type="bibr">Tshikolovets 2011</xref>
) in comparison to<xref rid="B34" ref-type="bibr">Kudrna et al. (2011)</xref>
.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Corrected species names</th>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Anthocharis damone</named-content>
</italic>
Boisduval, 1836</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Apatura metis</named-content>
</italic>
Freyer, 1829</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Argynnis elisa</named-content>
</italic>
Godart, 1823</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Aricia morronensis</named-content>
</italic>
Ribbe, 1910</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Cacyreus marshalli</named-content>
</italic>
Butler, 1898</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Colias aurorina</named-content>
</italic>
Herrich-Schäffer, 1850</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Erebia ottomana</named-content>
</italic>
Herrich-Schäffer, 1847</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Maniola chia</named-content>
</italic>
Thomson, 1987</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Maniola halicarnassus</named-content>
</italic>
Thomson, 1990</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Melitaea asteria</named-content>
</italic>
Freyer, 1828</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Melitaea varia</named-content>
</italic>
Meyer-Dür, 1851</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pararge xiphioides</named-content>
</italic>
Staudinger, 1871</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plebejus trappi</named-content>
</italic>
(Verity, 1927)</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pseudochazara amymone</named-content>
</italic>
Brown, 1976</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Pseudochazara orestes</named-content>
</italic>
Prins & Poorten, 1981</td>
</tr>
</tbody>
</table>
</table-wrap>
<pmc-comment>PageBreak</pmc-comment>
<p><italic><named-content content-type="taxon-name">Aricia artaxerxes</named-content>
</italic>
(Fabricius, 1793) and <italic><named-content content-type="taxon-name">Aricia montensis</named-content>
</italic>
Verity, 1928 are treated in CLIMBER as distinct species with parapatric distributions (see <xref rid="B38" ref-type="bibr">Sanudo-Restrepo et al. 2013</xref>
). The latter species is confined to the Iberian Peninsula and North Africa.</p>
<p>For the local Macedonian endemic <italic><named-content content-type="taxon-name">Pseudochazara amymone</named-content>
</italic>
Brown, 1976 no data were available for the considered time period. After its first discovery in Greece in 1975, the species was not reliably recorded again until its recent rediscovery in Southern Albania (<xref rid="B27" ref-type="bibr">Eckweiler 2012</xref>
). According to <xref rid="B27" ref-type="bibr">Eckweiler (2012)</xref>
, <italic><named-content content-type="taxon-name">Pseudochazara amymone</named-content>
</italic>
should be considered a subspecies of <italic><named-content content-type="taxon-name">Pseudochazara mamurra</named-content>
</italic>
(Herrich-Schäffer, [1846]), which is widespread in the Middle East.</p>
<p>The following species in our database actually comprise records of more than one species, most of which were recognized only recently, and are difficult or impossible to distinguish without genitalia examination or molecular methods.</p>
<list list-type="bullet"><list-item><p><italic><named-content content-type="taxon-name">Carcharodus alceae</named-content>
</italic>
(Esper, 1780) probably contains data of the sibling species <italic><named-content content-type="taxon-name">Carcharodus tripolinus</named-content>
</italic>
(Verity, 1925) from the Southern Iberian Peninsula, differing only in genitalia characters.</p>
</list-item>
<list-item><p><italic><named-content content-type="taxon-name">Leptidea sinapis</named-content>
</italic>
(Linnaeus, 1758) is a complex of three sibling species, and includes data of <italic><named-content content-type="taxon-name">Leptidea juvernica</named-content>
</italic>
Williams, 1946, and <italic><named-content content-type="taxon-name">Leptidea reali</named-content>
</italic>
Reissinger, 1990 (<xref rid="B24" ref-type="bibr">Dincă et al. 2011b</xref>
; <xref rid="B25" ref-type="bibr">Dincă et al. 2013</xref>
). Whereas <italic><named-content content-type="taxon-name">Leptidea sinapis</named-content>
</italic>
can be separated by their genitalia, the other two taxa can only be separated from each other by molecular characters. <italic><named-content content-type="taxon-name">Leptidea reali</named-content>
</italic>
seems to replace <italic><named-content content-type="taxon-name">Leptidea juvernica</named-content>
</italic>
in SW Europe, and both occur largely in sympatry with <italic><named-content content-type="taxon-name">Leptidea sinapis</named-content>
</italic>
.</p>
</list-item>
<list-item><p><italic><named-content content-type="taxon-name">Lycaena tityrus</named-content>
</italic>
(Poda, 1761) includes data of <italic><named-content content-type="taxon-name">Lycaena bleusei</named-content>
</italic>
Oberthür, 1884 from Central Spain and Central Portugal, which appears to be a distinct species according to unpublished molecular data.</p>
</list-item>
<list-item><p><italic><named-content content-type="taxon-name">Melitaea athalia</named-content>
</italic>
(Rottemburg, 1775) includes the Southwest European <italic><named-content content-type="taxon-name">Melitaea nevadensis</named-content>
</italic>
Oberthür, 1904 (syn. <italic>celadussa</italic>
Fruhstorfer, 1910) which might only be a subspecies of the former. Molecular data are inconclusive regarding the taxonomic status of these parapatric taxa.</p>
</list-item>
<list-item><p><italic><named-content content-type="taxon-name">Melitaea phoebe</named-content>
</italic>
(Goeze, 1779) recently turned out to be a complex of at least two largely sympatric species with distinctive larval colouration, and our data probably include records of <italic><named-content content-type="taxon-name">Melitaea ornata</named-content>
</italic>
Christoph, 1893 (syn. <italic>telona</italic>
Fruhstorfer, 1908 and <italic>emipunica</italic>
Verity, 1919) (see <xref rid="B41" ref-type="bibr">Toth et al. 2013</xref>
; <xref rid="B42" ref-type="bibr">Toth and Varga 2011</xref>
; <xref rid="B43" ref-type="bibr">Tshikolovets 2011</xref>
).</p>
</list-item>
<list-item><p><italic><named-content content-type="taxon-name">Polyommatus icarus</named-content>
</italic>
(Rottemburg, 1775) includes data of <italic><named-content content-type="taxon-name">Polyommatus celina</named-content>
</italic>
(Austaut, 1879), which was recognized as a distinct species from North Africa and the Canary Islands by molecular methods (<xref rid="B45" ref-type="bibr">Wiemers et al. 2010</xref>
), but also occurs in Southern Spain, and appears to replace <italic><named-content content-type="taxon-name">Polyommatus icarus</named-content>
</italic>
in the Balearic Islands, Sardinia, and Sicily (<xref rid="B23" ref-type="bibr">Dincă et al. 2011a</xref>
).</p>
</list-item>
<list-item><p><italic><named-content content-type="taxon-name">Pontia daplidice</named-content>
</italic>
(Linnaeus, 1758) includes the data of the sibling species <italic><named-content content-type="taxon-name">Pontia edusa</named-content>
</italic>
(Fabricius, 1777), a parapatric taxon, which can only be distinguished by molecular methods (<xref rid="B29" ref-type="bibr">Geiger and Scholl 1982</xref>
; <xref rid="B32" ref-type="bibr">John et al. 2013</xref>
; Wiemers unpubl.).</p>
</list-item>
</list>
<pmc-comment>PageBreak</pmc-comment>
</sec>
<sec sec-type="methods"><title>Methods</title>
<sec sec-type="Butterfly distribution data"><title>Butterfly distribution data</title>
<p>Climatic niche characteristics of the butterflies in Europe are based on their European distribution. Butterfly distributions were available from about 7000 georeferenced localities and about 200,000 database records. These records were stored in a database and constituted also the basis for ‘The Distribution Atlas of European Butterflies’ (<xref rid="B33" ref-type="bibr">Kudrna 2002</xref>
) and, as an updated version, for the ‘Distribution Atlas of Butterflies in Europe’ (<xref rid="B34" ref-type="bibr">Kudrna et al. 2011</xref>
; <xref ref-type="fig" rid="F6">Fig. 6</xref>
). The data are owned by the Helmholtz Centre for Environmental Research (and thus by the originators of CLIMBER). To avoid problems of occasional undersampling and imprecise geo-reference of some locations at the local scale, we re-sampled the localities to 1720 CGRS grid cells at a 50 km × 50 km resolution. Distribution data refer to the period of 1981–2000 and cover the abovementioned European window of 11°W, 32°E longitude and 34°N, 72°N latitude. We also provide an estimation of species range sizes by the number of grid cells used for calculating the climatic species characteristics.</p>
<fig id="F6" orientation="portrait" position="float"><label>Figure 6.</label>
<caption><p>Work flow and data sources for the generation of CLIMBER. Butterfly distribution data are based on a database which combines information from local recorders and private, regional and national databases. Thereof, species distributional maps have been developed. Together with maps of original and derived climate variables, based on interpolated data from local weather stations, species distribution-climate relationships have been assessed in a GIS. Based on these relationships several statistics describing the climatic characteristics of 397 European butterfly species have been developed and stored in CLIMBER. Several steps of quality control ensure a high level of data accuracy. CRU; Climate Research Unit, University of East Anglia (<ext-link ext-link-type="uri" xlink:href="http://www.cru.uea.ac.uk/">http://www.cru.uea.ac.uk/</ext-link>
). ALARM; EU, FP6 project ‘Assessing Large Scale Risks for Biodiversity with Tested Methods’ (<ext-link ext-link-type="uri" xlink:href="http://www.alarmproject.net/climate/climate/">http://www.alarmproject.net/climate/climate/</ext-link>
).</p>
</caption>
<graphic xlink:href="ZooKeys-367-065-g006"></graphic>
</fig>
<pmc-comment>PageBreak</pmc-comment>
</sec>
<sec sec-type="Climate data"><title>Climate data</title>
<p>We used monthly, interpolated climate data (publicly available at <ext-link ext-link-type="uri" xlink:href="http://www.alarmproject.net/climate/climate">http://www.alarmproject.net/climate/climate</ext-link>
), originally provided via the ALARM project (<xref rid="B17" ref-type="bibr">Settele et al. 2012</xref>
; <xref rid="B39" ref-type="bibr">Settele et al. 2005</xref>
; <xref rid="B19" ref-type="bibr">Spangenberg et al. 2012</xref>
) at a 10 arcmin grid resolution (<xref rid="B35" ref-type="bibr">Mitchell et al. 2004</xref>
; <xref rid="B36" ref-type="bibr">New et al. 2000</xref>
) and aggregated them to the CGRS grid (<xref ref-type="fig" rid="F6">Fig. 6</xref>
). For a detailed description of the climate data see <xref rid="B28" ref-type="bibr">Fronzek et al. (2012)</xref>
. The following basic climatic variables were used to assess aspects of the climatic niche: mean annual temperature (°C), range of annual temperature (°C), annual precipitation sum (mm), range of annual precipitation (mm), accumulated growing degree days with a base temperature of 5°C until February, April, June and August and soil water content for the upper horizon (0.5 m). Different time periods for calculating accumulated growing degree days enable the consideration of different phenologies and phenological aspects in the analysis of the climatic species characteristics. We do not provide growing degree days for periods ending later than August because these values are highly correlated with mean annual temperature in any case. Soil water content originated from the dynamic vegetation model LPJ-GUESS (<xref rid="B30" ref-type="bibr">Hickler et al. 2009</xref>
; <xref rid="B31" ref-type="bibr">Hickler et al. 2004</xref>
) which provides a process-based representation of the water balance in terrestrial ecosystems. According to the time period of the butterfly distribution data, we used averaged values for the period 1971–2000 for the climate data.</p>
</sec>
<sec sec-type="Calculation of the climatic niche characteristics"><title>Calculation of the climatic niche characteristics</title>
<p>Climatic niche characteristics were calculated per butterfly species according to the climatic conditions across their respective ranges, i.e. across all grid cells in which a particular species occurs (see <xref rid="B46" ref-type="bibr">Devictor et al. 2012a</xref>
; <xref rid="B48" ref-type="bibr">Schweiger et al. 2012</xref>
; <xref rid="B49" ref-type="bibr">Van Swaay et al. 2010</xref>
; <xref rid="B50" ref-type="bibr">Van Swaay et al. 2008</xref>
; <xref ref-type="fig" rid="F6">Fig. 6</xref>
). The dataset comprises information for the position and breadth of the climatic niche. Niche position is indicated by the median and mean value for each climate variable across a species’ range, accompanied by the 95% confidence interval for the mean. Niche breadth is indicated by the standard deviation and the minimum and maximum values for each climatic variable across a species’ range.</p>
</sec>
<sec sec-type="Data verification"><title>Data verification</title>
<p>Several steps of quality control ensure a high level of data accuracy (<xref ref-type="fig" rid="F6">Fig. 6</xref>
). During the step of compiling butterfly records for Europe, taxonomic experts addressed problems of potential misidentification, synonymy and the taxonomic concept. Once the species distribution maps had been produced, internal and external control ensured the elimination of obviously wrong records (species outside their natural range). Climate data are based on original climate variables from the Climate Research Unit (CRU) of <pmc-comment>PageBreak</pmc-comment>
the University of East Anglia and derived climate variables generated by the ALARM project. Both, CRU and ALARM ensured a high level of internal and external quality control. Data quality for the calculation of the climatic niche characteristics for each butterfly species is high (about 200,000 records for butterfly distribution; well recognised and commonly accepted climate data). Additionally, we provide the number of grid cells which have been used to calculate the climatic species characteristics and the standard deviation to assess uncertainty of the measures.</p>
</sec>
</sec>
<sec><title>Data status and accessibility</title>
<sec sec-type="Status"><title>Status</title>
<p>Data set version: v1.3</p>
<p>Latest update: 18.10.2013.</p>
<p>Metadata status: Metadata are complete and stored with the data.</p>
</sec>
<sec sec-type="Accessibility"><title>Accessibility</title>
<p><bold>Copyright restrictions:</bold>
None.</p>
<p><bold>Proprietary restrictions:</bold>
This dataset is freely available for non-commercial scientific use.</p>
<p><bold>Citation:</bold>
Data users must cite this Data Paper properly in any publication that results from an analysis using the provided data as a whole or in parts as: Schweiger O, Harpke A, Wiemers M, Settele J (2013). CLIMBER: Climatic niche characteristics of the butterflies in Europe. ZooKeys 367: 65–84. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.3897/zookeys.367.6185">10.3897/zookeys.367.6185</ext-link>
</p>
<p><bold>In addition to the Data Paper the resource should be cited as:</bold>
Helmholtz Centre for Environmental Research - UFZ (2013). CLIMBER: Climatic niche characteristics</p>
<p>of the butterflies in Europe. 397 records, Online at <ext-link ext-link-type="uri" xlink:href="http://ipt.pensoft.net/ipt/resource.do?r=climber">http://ipt.pensoft.net/ipt/resource.do?r=climber</ext-link>
, version 1.3 (released on 3/12/2013), Resource ID: GBIF key: <ext-link ext-link-type="uri" xlink:href="http://www.gbif.org/dataset/e2bcea8c-dfea-475e-a4ae-af282b4ea1c5">http://www.gbif.org/dataset/e2bcea8c-dfea-475e-a4ae-af282b4ea1c5</ext-link>
, Data Paper ID: doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.3897/zookeys.367.6185">10.3897/zookeys.367.6185</ext-link>
</p>
<sec sec-type="Collab"><title>Collab</title>
<p><bold>oration:</bold>
Data users might consider collaboration and/or co-authorship with the data owners.</p>
<p><bold>Storage location:</bold>
<ext-link ext-link-type="uri" xlink:href="http://ipt.pensoft.net/ipt/resource.do?r=climber">http://ipt.pensoft.net/ipt/resource.do?r=climber</ext-link>
</p>
</sec>
</sec>
</sec>
<sec><title>Data structure</title>
<sec sec-type="Dataset file"><title>Dataset file</title>
<p><bold>File name:</bold>
CLIMBER.v.1.3.csv</p>
<p><bold>Size:</bold>
398 rows, 67 columns; 183 kB.</p>
<pmc-comment>PageBreak</pmc-comment>
<p><bold>Format and storage mode:</bold>
ASCII csv, semicolon-delimited; decimal separator: ‘.’.</p>
<p><bold>Header information:</bold>
First row provides variable names.</p>
<p><bold>Alphanumeric attributes:</bold>
Mixed.</p>
<p><bold>Special characters:</bold>
Missing values are indicated by NA.</p>
</sec>
<sec sec-type="Variable definition"><title>Variable definition</title>
<p>Climatic niche characteristics are based on nine climate variables (<xref ref-type="table" rid="T3">Table 3</xref>
). All climate variables represent average values for the period of 1971–2000. Seven statistics are available for each climate variable (<xref ref-type="table" rid="T4">Table 4</xref>
).</p>
<table-wrap id="T3" orientation="portrait" position="float"><label>Table 3.</label>
<caption><p>Climatic variables used for the assessment of climatic niche characteristics of the butterflies in Europe.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Name</th>
<th rowspan="1" colspan="1">Definition</th>
<th rowspan="1" colspan="1">Unit</th>
<th rowspan="1" colspan="1">Interpretation</th>
</tr>
<tr><td rowspan="1" colspan="1">range.size</td>
<td rowspan="1" colspan="1">Distributional range size as number of occupied grids</td>
<td rowspan="1" colspan="1">Grid cells</td>
<td rowspan="1" colspan="1">Sample size</td>
</tr>
<tr><td rowspan="1" colspan="1">temp</td>
<td rowspan="1" colspan="1">Mean annual temperature</td>
<td rowspan="1" colspan="1">°C</td>
<td rowspan="1" colspan="1">Temperature (STI)</td>
</tr>
<tr><td rowspan="1" colspan="1">range.ann.temp</td>
<td rowspan="1" colspan="1">Annual range in monthly temperature (warmest month - coldest month)</td>
<td rowspan="1" colspan="1">°C</td>
<td rowspan="1" colspan="1">Continentality</td>
</tr>
<tr><td rowspan="1" colspan="1">precip</td>
<td rowspan="1" colspan="1">Annual precipitation sum</td>
<td rowspan="1" colspan="1">mm</td>
<td rowspan="1" colspan="1">Precipitation</td>
</tr>
<tr><td rowspan="1" colspan="1">range.ann.precip</td>
<td rowspan="1" colspan="1">Annual range in monthly precipitation sum (wettest month - driest month)</td>
<td rowspan="1" colspan="1">mm</td>
<td rowspan="1" colspan="1">Oceanity</td>
</tr>
<tr><td rowspan="1" colspan="1">gdd.feb</td>
<td rowspan="1" colspan="1">Accumulated growing degree days above 5°C from January to February</td>
<td rowspan="1" colspan="1">°C</td>
<td rowspan="1" colspan="1">Temperature corrected for metabolic activity preconditions</td>
</tr>
<tr><td rowspan="1" colspan="1">gdd.apr</td>
<td rowspan="1" colspan="1">Accumulated growing degree days above 5°C from January to April</td>
<td rowspan="1" colspan="1">°C</td>
<td rowspan="1" colspan="1">Temperature corrected for metabolic activity preconditions</td>
</tr>
<tr><td rowspan="1" colspan="1">gdd.june</td>
<td rowspan="1" colspan="1">Accumulated growing degree days above 5°C from January to June</td>
<td rowspan="1" colspan="1">°C</td>
<td rowspan="1" colspan="1">Temperature corrected for metabolic activity preconditions</td>
</tr>
<tr><td rowspan="1" colspan="1">gdd.aug</td>
<td rowspan="1" colspan="1">Accumulated growing degree days above 5°C from January to August</td>
<td rowspan="1" colspan="1">°C</td>
<td rowspan="1" colspan="1">Temperature corrected for metabolic activity preconditions</td>
</tr>
<tr><td rowspan="1" colspan="1">swc</td>
<td rowspan="1" colspan="1">Soil water content of the upper horizon (0.5 m)</td>
<td rowspan="1" colspan="1">No unit (0-1)</td>
<td rowspan="1" colspan="1">Water availability</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T4" orientation="portrait" position="float"><label>Table 4.</label>
<caption><p>Statistics available for each climate variable describing the niche position and breadth for the butterflies in Europe.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Name</th>
<th rowspan="1" colspan="1">Definition</th>
<th rowspan="1" colspan="1">Interpretation</th>
</tr>
<tr><td rowspan="1" colspan="1">mean</td>
<td rowspan="1" colspan="1">Mean value of climate variable across the species’ range</td>
<td rowspan="1" colspan="1">‘Optimal’ climatic conditions; niche position</td>
</tr>
<tr><td rowspan="1" colspan="1">ci.95.low</td>
<td rowspan="1" colspan="1">Lower 95% confidence interval for the mean</td>
<td rowspan="1" colspan="1">Uncertainty of the mean</td>
</tr>
<tr><td rowspan="1" colspan="1">ci.95.up</td>
<td rowspan="1" colspan="1">Upper 95% confidence interval for the mean</td>
<td rowspan="1" colspan="1">Uncertainty of the mean</td>
</tr>
<tr><td rowspan="1" colspan="1">min</td>
<td rowspan="1" colspan="1">Minimum value of the climate variable across the species range</td>
<td rowspan="1" colspan="1">Lower climatic limit</td>
</tr>
<tr><td rowspan="1" colspan="1">max</td>
<td rowspan="1" colspan="1">Maximum value of the climate variable across the species range</td>
<td rowspan="1" colspan="1">Upper climatic limit</td>
</tr>
<tr><td rowspan="1" colspan="1">sd</td>
<td rowspan="1" colspan="1">Standard deviation of the climate variable across the species range</td>
<td rowspan="1" colspan="1">Niche breadth</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>We also provide an estimation of species range size (range.size) to assess the number of grid cells used for calculating the climatic species characteristics. For a detailed description of swc see section Climate data. Annual measures are calculated over full 12 month periods, while accumulated growing degree days have been calculated for four periods from January to February, April, June and August to cover a variety of phenological aspects and life cycle stages.</p>
<p>Species range refers to the distributional range according to the 50 km × 50 km CGRS grid cells in which a species was recorded.</p>
</sec>
<sec sec-type="Data anomalies"><title>Data anomalies</title>
<p><bold>Missing values:</bold>
NA indicates that a species was only present in one grid cell and thus 95% confidence intervals and standard deviation could not be calculated.</p>
<pmc-comment>PageBreak</pmc-comment>
</sec>
</sec>
</body>
<back><ack><title>Acknowledgements</title>
<p>Most of the distributional data were compiled by Dr. Otakar Kudrna, especially within the framework of <xref rid="B33" ref-type="bibr">Kudrna (2002)</xref>
and <xref rid="B34" ref-type="bibr">Kudrna et al. (2011)</xref>
. We further acknowledge the support of European Commission Framework Programme (FP) 6 via the Integrated Project ALARM (GOCE-CT-2003-506675; <xref rid="B39" ref-type="bibr">Settele et al. 2005</xref>
) and FP 7 via the Integrated Project SCALES (grant 226 852; <xref rid="B5" ref-type="bibr">Henle et al. 2010</xref>
) and the Collaborative Project STEP (grant 244090 – STEP – CP – FP; <xref rid="B12" ref-type="bibr">Potts et al. 2011</xref>
) and the FP6 BiodivERsA Eranet project CLIMIT (funded by DLR-BMBF (Germany), NERC and DEFRA (UK), ANR (France), Formas (Sweden) and Swedish EPA (Sweden); <xref rid="B18" ref-type="bibr">Settele and Kühn 2009</xref>
; <xref rid="B20" ref-type="bibr">Thomas et al. 2009</xref>
).</p>
</ack>
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<app-group><app id="APP1"><title>Appendix</title>
<supplementary-material content-type="local-data" id="S1"><caption><p>Database of the climatic niche characteristics of the butterflies in Europe (CLIMBER). (doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.3897/zookeys.367.6185.app">10.3897/zookeys.367.6185.app</ext-link>
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<media xlink:href="ZooKeys-367-065-s001.csv" mimetype="Comma-Separated Values File" mime-subtype="csv" orientation="portrait" xlink:type="simple" id="d36e4687" position="anchor"></media>
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</record>
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