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10.3897_zookeys.100.1543.xml
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10.3897_zookeys.100.1543.xml
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<!DOCTYPE article PUBLIC "-//TaxonX//DTD Taxonomic Treatment Publishing DTD v0 20100105//EN" "tax-treatment-NS0.dtd"><article article-type="research-article" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:tp="http://www.plazi.org/taxpub">
<front>
<journal-meta>
<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="doi">10.3897/zookeys.100.1543</article-id>
<title-group>
<article-title>What do we know about winter active ground beetles (Coleoptera, Carabidae) in Central and Northern Europe?</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Jaskuła</surname>
<given-names>Radomir</given-names>
</name>
<xref ref-type="aff" rid="A1"/>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Soszyńska-Maj</surname>
<given-names>Agnieszka</given-names>
</name>
<xref ref-type="aff" rid="A1"/>
</contrib>
</contrib-group>
<aff id="A1">
<label/>Department of Invertebrate Zoology & Hydrobiology, University of Łódź, Banacha 12/16, 90-237 Łódź, Poland</aff>
<author-notes>
<fn fn-type="corresp">
<p>Corresponding authors: Radomir Jaskuła (<email xlink:type="simple">radekj@biol.uni.lodz.pl</email>), Agnieszka Soszyńska-Maj (<email xlink:type="simple">agasosz@biol.uni.lodz.pl</email>).</p>
</fn>
<fn fn-type="edited-by">
<p>Academic editor: J. Noordijk</p>
</fn>
</author-notes>
<pub-date pub-type="collection">
<year>2011</year>
</pub-date>
<pub-date pub-type="epub">
<day>20</day>
<month>5</month>
<year>2011</year>
</pub-date>
<issue>100</issue>
<fpage>517</fpage>
<lpage>532</lpage>
<history>
<date date-type="received">
<day>26</day>
<month>11</month>
<year>2009</year>
</date>
<date date-type="accepted">
<day>08</day>
<month>4</month>
<year>2010</year>
</date>
</history>
<permissions>
<copyright-statement>Radomir Jaskuła, Agnieszka Soszyńska-Maj</copyright-statement>
<license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/3.0" xlink:type="simple">
<license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License, 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>This paper summarizes the current knowledge on winter active Carabidae in Central and Northern Europe. In total 73 winter active species are listed, based on literature and own observations. Ground beetles are among the three most numerous <italic><tp:taxon-name>Coleoptera</tp:taxon-name></italic> families active during the autumn to spring period. The winter community of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> is composed both of larvae (mainly autumn breeding species) and adults, as well as of epigeic species and those inhabiting tree trunks. Supranivean fauna is characterized by lower species diversity than the subnivean fauna. The activity of ground beetles decreases in late autumn, is lowest during mid-winter and increases in early spring. <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> are noted as an important food source in the diet of insectivorous mammals. They are also predators, hunting small winter active invertebrates.</p>
</abstract>
<kwd-group>
<label>Keywords</label>
<kwd>Coleoptera</kwd>
<kwd>Carabidae</kwd>
<kwd>Central Europe</kwd>
<kwd>winter activity</kwd>
<kwd>subnivean fauna</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec sec-type="Introduction"><title>Introduction</title>
<p>During winter, invertebrates are mostly inactive in diapause as eggs, larvae or pupae, but less often as adult stages (<xref ref-type="bibr" rid="B34">Leather et al. 1993</xref>). Body fluids may freeze in low temperatures, so to avoid death, insects employ two main strategies: avoiding freezing or tolerating freezing. The adaptation to avoid freezing is the ability of supercooling by synthesizing antifreezing agents (e.g., glycerol) (<xref ref-type="bibr" rid="B43">Moore and Lee 1991</xref>). Some poikilothermic organisms may stay active in winter. These organisms use favourable atmos<!--PageBreak-->pheric condition – mild winter days with low air pressure – for migration and copulation (<xref ref-type="bibr" rid="B53">Soszyńska 2004</xref>). Their activity in low temperatures is usually related to the presence of snow and to their thermal properties. Snow cover has a high insulation capacity and low thermal conductivity due to its high air content. Low density and a greater thickness of snow (depending on geographical area) provide better insulation. The soil and litter can remain warm even if air temperature is very low (<xref ref-type="bibr" rid="B3">Aitchison 1974</xref>, <xref ref-type="bibr" rid="B11">2001</xref>).</p>
<p>Snow cover provides winter active animals with three different microhabitats. The insulating properties of snow make the space under the snow a favourable habitat for invertebrates (subnivean microenvironment). The subnivean microhabitat is relatively warm, humid, thermally stable and protects organisms from wind and lethal temperatures in contrast to the snow surface (supranivean environment), which is highly variable and completely dependent on atmospheric factors. Within the snow, the so-called intranivean habitat, temperatures are lower but organisms are still protected from the external environment (<xref ref-type="bibr" rid="B11">Aitchison 2001</xref>). Animals that are active in snow can be divided into two main groups, depending on their period of activity. The first group consists of “true winter” organisms that are active during the winter months (end of November until the beginning of April) both under and on the snow cover. The second is a nival fauna that are active on the snow cover outside the winter months. Examples of these fauna are permanent residents of high-altitude regions and glaciers. These invertebrates are adapted to permanent snow, glacier surfaces, etc. Their food source is the aeolian fauna, which consists of invertebrates passively deposited on snow fields (<xref ref-type="bibr" rid="B41">Mani 1962</xref>, <xref ref-type="bibr" rid="B11">Aitchison 2001</xref>).</p>
<p>The snow fauna is an ecological group, which consists of permanent snow active invertebrate species. The first observations regarding invertebrate activity on the snow was made in Poland in the middle of 18th century (<xref ref-type="bibr" rid="B17">Fedorowicz 1968</xref>). Since then, snow active insects have been the main subject of investigation in only a few elaborate studies. Snow activity was observed in many insect orders: Collembola, <italic><tp:taxon-name>Trichoptera</tp:taxon-name></italic>, <italic><tp:taxon-name>Plecoptera</tp:taxon-name></italic>, Blattodea, <italic><tp:taxon-name>Hemiptera</tp:taxon-name></italic>, <italic><tp:taxon-name>Mecoptera</tp:taxon-name></italic>, <italic><tp:taxon-name>Coleoptera</tp:taxon-name></italic>, <italic><tp:taxon-name>Diptera</tp:taxon-name></italic>, and <italic><tp:taxon-name>Hymenoptera</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B20">Frey 1913</xref>, <xref ref-type="bibr" rid="B59">Tahvonen 1942</xref>, <xref ref-type="bibr" rid="B58">Szulczewski 1947</xref>, <xref ref-type="bibr" rid="B63">Ulfstrand 1968</xref>, <xref ref-type="bibr" rid="B13">Brummer-Korvenkontio and Brummer-Korvenkontio 1980</xref>, <xref ref-type="bibr" rid="B35">Leinaas 1981</xref>, <xref ref-type="bibr" rid="B36">1983</xref>, <xref ref-type="bibr" rid="B23">Hågvar 1995</xref>, <xref ref-type="bibr" rid="B24">2000</xref>, <xref ref-type="bibr" rid="B11">Aitchison 2001</xref>, <xref ref-type="bibr" rid="B54">Soszyńska and Durska 2002</xref>, <xref ref-type="bibr" rid="B53">Soszyńska 2004</xref>, <xref ref-type="bibr" rid="B25">Hågvar and Greve 2003</xref>).</p>
<p>The first information about subnivean fauna appeared almost two centuries later than that of the fauna living on the snow. The subnivean microenvironment is inhabited by more numerous groups of invertebrates, such as oligochaetes, molluscs, crustaceans, arachnids and insects. Among these, insects and spiders clearly predominate, being the major representatives of the snow active fauna. The subnivean fauna was studied more often than the snow active fauna. Main studies came from Canada (<xref ref-type="bibr" rid="B4">Aitchison 1978</xref>, <xref ref-type="bibr" rid="B5">1979a</xref><xref ref-type="bibr" rid="B6"/><xref ref-type="bibr" rid="B7"/>-<xref ref-type="bibr" rid="B8">d</xref>, <xref ref-type="bibr" rid="B9">1984</xref>, <xref ref-type="bibr" rid="B11">2001</xref>), the USA (<xref ref-type="bibr" rid="B52">Schmidt and Lockwood 1992</xref>, <xref ref-type="bibr" rid="B2">Addington and Seastedt 1999</xref>), as well as from central and northern Europe (<xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B1">Ackefors 1964</xref>, <xref ref-type="bibr" rid="B45">Näsmark 1964</xref>, <xref ref-type="bibr" rid="B42">Merriam et al. 1983</xref>, <xref ref-type="bibr" rid="B29">Itämies and Lindgren 1989</xref>, <xref ref-type="bibr" rid="B40">Łęgowski and Łoziński 1995</xref>). The most common orders in terms of species diversity as well as percentage contribution to this ecological group are Collembola, <italic><tp:taxon-name>Coleoptera</tp:taxon-name></italic>, <italic><tp:taxon-name>Diptera</tp:taxon-name></italic>, but also <italic><tp:taxon-name>Hymenoptera</tp:taxon-name></italic>, and <italic><tp:taxon-name>Hemiptera</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B59">Tahvonen 1942</xref>, <xref ref-type="bibr" rid="B1">Ackefors 1964</xref>, <xref ref-type="bibr" rid="B5">Aitchison 1979a</xref><xref ref-type="bibr" rid="B6"/><xref ref-type="bibr" rid="B7"/>-<xref ref-type="bibr" rid="B8">d</xref>, <xref ref-type="bibr" rid="B9">1984</xref>, <xref ref-type="bibr" rid="B11">2001</xref>).<!--PageBreak--></p>
<p>During the last decades, global climate change has become an important scientific topic. However its influence on poikilothermic organisms has been poorly investigated. It seems that the occurrence of snow cover during the winter period plays an important role in the biology of many different invertebrate groups.</p>
<p>The aim of this paper is to summarize knowledge regarding winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> fauna from Central and Northern Europe. In the present paper we discuss only the “true winter active” ground beetles, and not members of the nival fauna occurring in high-altitude regions or glaciers.</p>
</sec>
<sec sec-type="methods"><title>Methods</title>
<p>Winter season is defined here as the period between the end of November and the beginning of April. All available literature data on winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> recorded from Central and Northern Europe were used in this study. In total, data from five countries and published in 17 papers were analyzed (see <xref ref-type="table" rid="T1">Table 1</xref>). Data of mountain <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> active on the snow and glaciers as well as species found overwintering in diapause were not included. In addition, our unpublished records of winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> from Central Poland were included. This material was collected occasionally during different field studies using pitfall traps (subnivean species) and active searching on the snow cover. The list of species analyzed in this study is given in <xref ref-type="table" rid="T1">Table 1</xref>.</p>
<p>All recorded ground beetle species were divided into three groups, according to the microenvironment in which they were noted: epigeic (subnivean), active on the snow cover (supranivean), and actively walking on tree trunks. Data on activity of both the adults and the larvae are also shown in <xref ref-type="table" rid="T1">Table 1</xref>.</p>
<p>The ecological response towards snow active ground beetle species was done according to <xref ref-type="bibr" rid="B21">Fudakowski (1959)</xref> and Pruitt (1978). These authors distinguished the following species groups according to their ecological reaction towards snow: chionobionts – stenothermic species with adaptations to survive on snow and to reproduce in winter, chionophiles – eurythermic permanent snow active group, but its members occur also in other seasons, chionoxenes – species accidentally found in winter; chionophobes – group that avoids snow.</p>
<p>For the nomenclature of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species, the Fauna Europea Web Service (2004) was followed, while the zoogeographical analysis of ground beetles was based on the study by <xref ref-type="bibr" rid="B37">Leśniak (1988)</xref>.</p>
</sec>
<sec sec-type="Results and discussion"><title>Results and discussion</title>
<sec sec-type="Winter active Carabidae – a short history of faunistic studies"><title>Winter active Carabidae – a short history of faunistic studies</title>
<p>Most studies performed on winter active ground beetles are rather recent (<xref ref-type="table" rid="T1">Table 1</xref>). The first faunistic data on winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> came from the beginning of 20th <!--PageBreak--><!--PageBreak--><!--PageBreak--><!--PageBreak-->century, when five species belonging to the genera <italic><tp:taxon-name>Leistus</tp:taxon-name></italic>, <italic><tp:taxon-name>Bradycellus</tp:taxon-name></italic>, <italic><tp:taxon-name>Dromius</tp:taxon-name></italic>, <italic><tp:taxon-name>Ocydromus</tp:taxon-name></italic> and <italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic> were noted in Finland as active on the snow surface by <xref ref-type="bibr" rid="B20">Frey (1913)</xref> and <xref ref-type="bibr" rid="B38">Levander (1913)</xref>. After more than three decades, one additional species from the genus <italic><tp:taxon-name>Agonum</tp:taxon-name></italic> was found on the snow surface by Polish entomologist <xref ref-type="bibr" rid="B58">Szulczewski (1947)</xref> in the Wielkopolski National Park (western Poland). More recently, one additional species – <italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic> – was reported by <xref ref-type="bibr" rid="B31">Jaskuła et al. (2005)</xref> from central Poland. All these papers presented only single, accidental observations. Our work summarizes up-to-date knowledge about this ecological group and gives a list of 11 species belonging to 10 genera, including first data on activity of the genera <italic><tp:taxon-name>Calathus</tp:taxon-name></italic>, <italic><tp:taxon-name>Carabus</tp:taxon-name></italic>, <italic><tp:taxon-name>Notiophilus</tp:taxon-name></italic>, and <italic><tp:taxon-name>Paradromius</tp:taxon-name></italic> from the snow surface.</p>
<p>Compared to supranivean species (which are easier to observe because of the contrast between the white colour of the snow and the dark coloured insects), the carab<!--PageBreak-->ids active under the snow surface (subnivean fauna) were discovered rather late. First data on subnivean ground beetles became available after using Barber’s traps as a collecting method, and in Central and Northern Europe were given from Germany by <xref ref-type="bibr" rid="B49">Renken (1956)</xref>. He provided information on seven species of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> from the following genera: <italic><tp:taxon-name>Anchomenus</tp:taxon-name></italic>, <italic><tp:taxon-name>Demetrias</tp:taxon-name></italic>, <italic><tp:taxon-name>Dicheirotrichus</tp:taxon-name></italic>, <italic><tp:taxon-name>Calathus</tp:taxon-name></italic>, <italic><tp:taxon-name>Nebria</tp:taxon-name></italic>, <italic><tp:taxon-name>Philorhizus</tp:taxon-name></italic> and <italic><tp:taxon-name>Trechus</tp:taxon-name></italic>. All these species were imagines. However <xref ref-type="bibr" rid="B15">Evans (1969)</xref>, using the same method of study, recorded also larvae of <italic><tp:taxon-name>Cychrus caraboides</tp:taxon-name></italic>, <italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic>, <italic><tp:taxon-name>Abax</tp:taxon-name></italic> sp., <italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic> sp., <italic><tp:taxon-name>Leistus</tp:taxon-name></italic> sp., and <italic><tp:taxon-name>Carabus</tp:taxon-name></italic> sp. as being active under the snow surface. Additional records of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> larvae were added by <xref ref-type="bibr" rid="B32">Kaczmarek (1958)</xref>, <xref ref-type="bibr" rid="B45">Näsmark (1964)</xref>, <xref ref-type="bibr" rid="B22">Greenslade (1965)</xref>, and here.</p>
<p>More adult beetles were later collected by <xref ref-type="bibr" rid="B32">Kaczmarek (1958</xref> – 6 species from 6 genera), <xref ref-type="bibr" rid="B45">(Näsmark (1964</xref> – 1 species), <xref ref-type="bibr" rid="B22">Greenslade (1965</xref> –18 species from 12 genera), <xref ref-type="bibr" rid="B44">Murdoch (1967</xref> – 17 species from 12 genera), <xref ref-type="bibr" rid="B19">Flatz and Thaler (1980</xref> – 6 species from 6 genera), <xref ref-type="bibr" rid="B12">Betz (1992</xref> – 1 species), <xref ref-type="bibr" rid="B33">Kennedy (1994</xref> – 11 species from 8 genera), <xref ref-type="bibr" rid="B61">Traugott (1998</xref> – 7 species from 5 genera), and <xref ref-type="bibr" rid="B30">Jaskuła and Grabowski (2003</xref> – 1 species). Finally, in the present paper a list of 66 <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species is given, including one genus (<italic><tp:taxon-name>Anisodactylus</tp:taxon-name></italic>) recorded for the first time as a supranivean taxon.</p>
<p>Comparing the two above-mentioned “ecological groups”, it becomes clear that in the studied area, diversity of the subnivean carabid fauna is more than five times higher than that of the supranivean species (<xref ref-type="fig" rid="F1">Fig. 1</xref>). A similar tendency was observed in Collembola, but was opposite when compared to some other insect groups like <italic><tp:taxon-name>Diptera</tp:taxon-name></italic> or <italic><tp:taxon-name>Mecoptera</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B55">Soszyńska-Maj 2005</xref>).</p>
<p>Tree trunks are the third type of microhabitat where winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> occur. The only paper on this topic known to us comes from <xref ref-type="bibr" rid="B26">Hannig et al. (2006)</xref> who noted six species in Germany: <italic><tp:taxon-name>Amara familiaris</tp:taxon-name></italic>, <italic><tp:taxon-name>Calodromius bifasciatus</tp:taxon-name></italic>, <italic><tp:taxon-name>Calodromius spilotus</tp:taxon-name></italic>, <italic><tp:taxon-name>Philorhizus melanocephalus</tp:taxon-name></italic>, <italic><tp:taxon-name>Dromius angustus</tp:taxon-name></italic>,and <italic><tp:taxon-name>Dromius quadrimaculatus</tp:taxon-name></italic>. Among them, <italic><tp:taxon-name>Dromius quadrimaculatus</tp:taxon-name></italic> predominated and the genus <italic><tp:taxon-name>Calodromius</tp:taxon-name></italic> was noted as winter active for the first time (see <xref ref-type="bibr" rid="B18">Felix and Van Wielink 2011</xref>).</p>
<table-wrap position="float" orientation="portrait" id="T1">
<label>Table 1.</label>
<caption><p>List of winter active ground beetles (<bold>A</bold> – adults, <bold>L</bold> – larvae). Roman letters indicate the month(s) of observation(s). Nomenclature after <xref ref-type="bibr" rid="B16">Fauna Europaea Web Service (2004)</xref>.</p></caption>
<table>
<tbody>
<tr>
<th rowspan="1" colspan="1"><italic>No.</italic></th>
<th rowspan="1" colspan="1"><italic>Species</italic></th>
<th rowspan="1" colspan="1"/>
<th rowspan="1" colspan="1"><italic>Snow cover</italic></th>
<th rowspan="1" colspan="1"><italic>Epigeic(subnivean)</italic></th>
<th rowspan="1" colspan="1"><italic>Treetrunks</italic></th>
<th rowspan="1" colspan="1"><italic>Source</italic></th>
</tr>
<tr>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Abax parallelepipedus</tp:taxon-name></italic> (Piller et Mitterpacher, 1783)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII, III-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Abax</tp:taxon-name></italic> sp./<italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic> sp.</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B15">Evans 1969</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Acupalpus dubius</tp:taxon-name></italic> Schilsky, 1888</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Agonum gracile</tp:taxon-name></italic> Sturm, 1824</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Agonum muelleri</tp:taxon-name></italic> (Herbst, 1784)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">I</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B58">Szulczewski 1947</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">6</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Agonum viduum</tp:taxon-name></italic> (Panzer, 1796)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">7</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Amara aulica</tp:taxon-name></italic> (Panzer, 1796)</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-I </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">8</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Amara brunnea</tp:taxon-name></italic> (Gyllenhal, 1810)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">9</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Amara communis</tp:taxon-name></italic> (Panzer, 1797)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B19">Flatz and Thaler 1980</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">10</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Amara infima</tp:taxon-name></italic> (Duftschmid, 1812)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-I (?)</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">11</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Amara familiaris</tp:taxon-name></italic> (Duftschmid, 1812)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">III-IV</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">12</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Amara lunicollis</tp:taxon-name></italic> Schiødte, 1837</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">III-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">13</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Amara</tp:taxon-name></italic> sp.</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B33">Kennedy 1994</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">14</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Anchomenus dorsalis</tp:taxon-name></italic> (Pontoppidan, 1763)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"> XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B19">Flatz and Thaler 1980</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">15</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Anisodactylus binotatus</tp:taxon-name></italic> (Fabricius, 1787)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">16</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Asaphidion flavipes</tp:taxon-name></italic> (Linné, 1761)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">17</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Asaphidion pallipes</tp:taxon-name></italic> (Schrank, 1781)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">18</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Badister sodalis</tp:taxon-name></italic> (Duftschmid, 1812)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">III-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">19</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Bradycellus caucasicus</tp:taxon-name></italic> (Chaudoir, 1846)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">I</td>
<td rowspan="1" colspan="1">XI-I</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B20">Frey 1913</xref>, <xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">20</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Bradycelus harpalinus</tp:taxon-name></italic> (Audinet-Serville, 1821)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">21</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Bradycellus verbasci</tp:taxon-name></italic> (Duftschmid, 1812)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII, II-III</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B15">Evans 1969</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">22</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Calathus erratus</tp:taxon-name></italic> (C.R. Sahlberg, 1827)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref></td>
</tr>
<tr>
<td rowspan="2" colspan="1">23</td>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Calathus fuscipes</tp:taxon-name></italic> Goeze, 1777</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B19">Flatz and Thaler 1980</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="2" colspan="1">24</td>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Calathus melanocephalus</tp:taxon-name></italic> (Linné, 1758)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XII-I </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">25</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Calathus micropterus</tp:taxon-name></italic> (Duftschmid, 1812)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">XI</td>
<td rowspan="1" colspan="1">XI-I (?)</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">26</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Calathus rotundicollis</tp:taxon-name></italic> Dejean, 1828</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">27</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Calodromius bifasciatus</tp:taxon-name></italic> (Dejean, 1825)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-III</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">28</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Calodromius spilotus</tp:taxon-name></italic> (Illiger, 1798)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-III</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">29</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Carabus convexus</tp:taxon-name></italic> Fabricius, 1775</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-III</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">30</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Carabus coriaceus</tp:taxon-name></italic> Linné, 1758</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XII-I </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">31</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Carabus hortensis</tp:taxon-name></italic> Linné, 1758</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XII </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">32</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Carabus nemoralis</tp:taxon-name></italic> O. F. Müller, 1764</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">I</td>
<td rowspan="1" colspan="1">XI-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B15">Evans 1969</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">33</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Carabus problematicus</tp:taxon-name></italic> Herbst, 1786</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B15">Evans 1969</xref>, <xref ref-type="bibr" rid="B12">Betz 1992</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">34</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Carabus</tp:taxon-name></italic> sp.</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-III </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B15">Evans 1969</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">35</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Cychrus caraboides</tp:taxon-name></italic> (Linné, 1758)</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII, III </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B15">Evans 1969</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">36</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Demetrias atricapillus</tp:taxon-name></italic> (Linné, 1758)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">37</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Dicheirotrichus cognatus</tp:taxon-name></italic> (Gyllenhal, 1827)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">38</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Dicheirotrichus placidus</tp:taxon-name></italic> (Gyllenhal, 1827)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">39</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Dromius angustus</tp:taxon-name></italic> Brullé, 1834</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">40</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Dromius quadrimaculatus</tp:taxon-name></italic> (Linné, 1758)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-III</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">41</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Dromius schneideri</tp:taxon-name></italic> Crotch, 1871</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">I</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B20">Frey 1913</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">42</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Dyschiriodes globosus</tp:taxon-name></italic> (Herbst, 1784)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B64">Weber 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">43</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Elaphrus cupreus</tp:taxon-name></italic> Duftschmid, 1812</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">III</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">44</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Epaphius secalis</tp:taxon-name></italic> (Paykull, 1790)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-I (?)</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">45</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Leistus rufomarginatus</tp:taxon-name></italic> (Duftschmid, 1812)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI, I-II</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B30">Jaskuła and Grabowski 2003</xref>, this paper</td>
</tr>
<tr>
<td rowspan="2" colspan="1">46</td>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Leistus ferrugineus</tp:taxon-name></italic> (Linné, 1758)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-I (?)</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1">I </td>
<td rowspan="1" colspan="1">XI-XII, III-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B38">Levander 1913</xref>, <xref ref-type="bibr" rid="B45">Näsmark 1964</xref>, <xref ref-type="bibr" rid="B22">Greenslade 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">47</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Leistus fulvibarbis</tp:taxon-name></italic> Dejean, 1826</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="2" colspan="1">48</td>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Leistus terminatus</tp:taxon-name></italic> (Panzer, 1793)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-II</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">49</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Leistus</tp:taxon-name></italic> sp.</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1">XII </td>
<td rowspan="1" colspan="1">II-III </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B15">Evans 1969</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">50</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Loricera pilicornis</tp:taxon-name></italic> (Fabricius, 1775)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">51</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Metallina lampros</tp:taxon-name></italic> (Herbst, 1784)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref>, this paper</td>
</tr>
<tr>
<td rowspan="2" colspan="1">52</td>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic> (Fabricius, 1792)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, <xref ref-type="bibr" rid="B15">Evans 1969</xref>, <xref ref-type="bibr" rid="B19">Flatz and Thaler 1980</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, <xref ref-type="bibr" rid="B15">Evans 1969</xref>, <xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">53</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Notiophilus biguttatus</tp:taxon-name></italic> (Fabricius, 1779)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">XII-I</td>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B15">Evans 1969</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">54</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Notiophilus rufipes</tp:taxon-name></italic> Curtis, 1829</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"> XI, I-II </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">55</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Notiophilus substriatus</tp:taxon-name></italic> C.R. Waterhouse, 1833</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"> XI, I-II </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">56</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Ocydromus tetracolus</tp:taxon-name></italic> (Say, 1823)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B20">Frey 1913</xref>, <xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, <xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">57</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Panagaeus bipustulatus</tp:taxon-name></italic> (Fabricius, 1775)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">58</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Paradromius linearis</tp:taxon-name></italic> (Olivier, 1795)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">59</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Paranchus albipes</tp:taxon-name></italic> (Fabricius, 1796)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI, II-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">60</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Philochthus aeneus</tp:taxon-name></italic> (Germar, 1824)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B33">Kennedy 1994</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">61</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Philochthus biguttatus</tp:taxon-name></italic> (Fabricius, 1779)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">62</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Philochthus guttula</tp:taxon-name></italic> (Fabricius, 1792)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI, I-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">63</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Philorhizus melanocephalus</tp:taxon-name></italic> (Dejean, 1825)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">+</td>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">64</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Phyla obtusa</tp:taxon-name></italic> (Audinet-Serville, 1821)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">65</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Poecilus versicolor</tp:taxon-name></italic> (Sturm, 1824)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI, I,II</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B19">Flatz and Thaler 1980</xref></td>
</tr>
<tr>
<td rowspan="2" colspan="1">66</td>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Pseudoofonus rufipes</tp:taxon-name></italic> (De Geer, 1774)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B64">Weber 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-III </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">67</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus diligens</tp:taxon-name></italic> (Sturm, 1824)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1">IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B20">Frey 1913</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">68</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus madidus</tp:taxon-name></italic> (Fabricius, 1775)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="2" colspan="1">69</td>
<td rowspan="2" colspan="1"><italic><tp:taxon-name>Pterostichus melanarius</tp:taxon-name></italic> (Illiger, 1798)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B19">Flatz and Thaler 1980</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-I </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B61">Traugott 1998</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">70</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus niger</tp:taxon-name></italic> (Schaller, 1783)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">71</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus nigrita</tp:taxon-name></italic> (Paykull, 1790)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1">XII</td>
<td rowspan="1" colspan="1">XI-XII, II-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">72</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus oblongopunctatus</tp:taxon-name></italic> (Fabricius, 1787)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XII, III-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="1">73</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus quadrifoveolatus</tp:taxon-name></italic> Letzner, 1852</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">74</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus strenuous</tp:taxon-name></italic> (Panzer, 1796)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, <xref ref-type="bibr" rid="B15">Evans 1969</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">75</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic> sp.</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV </td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B64">Weber 1965</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">76</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Stomis pumicatus</tp:taxon-name></italic> (Panzer, 1796)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">III</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">77</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Trechus obtusus</tp:taxon-name></italic> Erichson, 1837</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-XII, III</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B44">Murdoch 1967</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">78</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Trechus quadristriatus</tp:taxon-name></italic> (Schrank, 1781)</td>
<td rowspan="1" colspan="1">A</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B33">Kennedy 1994</xref></td>
</tr>
<tr>
<td rowspan="1" colspan="1">79</td>
<td rowspan="1" colspan="1">Larvae gen. sp.</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1">XI-IV</td>
<td rowspan="1" colspan="1"/>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref>, this paper</td>
</tr>
<tr>
<td rowspan="1" colspan="3"><italic>TOTAL</italic></td>
<td rowspan="1" colspan="1"><italic>11</italic></td>
<td rowspan="1" colspan="1"><italic>66</italic></td>
<td rowspan="1" colspan="1"><italic>6</italic></td>
<td rowspan="1" colspan="1"/>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec sec-type=""><title>Winter active carabid communities</title>
<p>The most common groups among winter active invertebrates are spiders and insects. Among hexapods, springtails (Collembola), beetles (<italic><tp:taxon-name>Coleoptera</tp:taxon-name></italic>), flies (<italic><tp:taxon-name>Diptera</tp:taxon-name></italic>) and scorpionflies (<italic><tp:taxon-name>Mecoptera</tp:taxon-name></italic>) predominate. Beetle activity under snow cover is well documented. Investigations on winter active fauna in central Poland show that the supranivean and subnivean insect winter assemblages differ in terms of percentage contribution of orders, as well as in species composition. Beetles have only a share of 13% in snow active insect communities, and 25% in material collected under the snow (<xref ref-type="bibr" rid="B55">Soszyńska-Maj 2005</xref>). Among subnivean <italic><tp:taxon-name>Coleoptera</tp:taxon-name></italic>, three families clearly predominate: <italic><tp:taxon-name>Staphylinidae</tp:taxon-name></italic>, <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> and <italic><tp:taxon-name>Cantharidae</tp:taxon-name></italic> (larvae), while carabids are only found accidentally on the snow. These three beetle groups are known as winter dominants, both in terms of species diversity and abundance (<xref ref-type="bibr" rid="B66">Wolska 1957</xref>, <xref ref-type="bibr" rid="B57">Strübing 1958</xref>, <xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B45">Näsmark 1964</xref>, <xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B6">Aitchison 1979b</xref>, <xref ref-type="bibr" rid="B9">1984</xref>, <xref ref-type="bibr" rid="B11">2001</xref>, <xref ref-type="bibr" rid="B42">Merriam et al. 1983</xref>, <xref ref-type="bibr" rid="B29">Itämies and Lindgren 1989</xref>, <xref ref-type="bibr" rid="B40">Łęgowski and Łoziński 1995</xref>, <xref ref-type="bibr" rid="B62">Traugott 2002</xref>). A total of 16 <italic><tp:taxon-name>Coleoptera</tp:taxon-name></italic> families have thus far been recorded as winter active (<xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref>, Soszyńska-Maj and Jaskuła unpublished data).</p>
<p>In general, the winter activity of <tp:taxon-name>Carabidae</tp:taxon-name> varies seasonally. Its peak – both according to the number of species and individuals – is observed in late autumn and early spring. The lowest activity is observed in mid-winter (<xref ref-type="fig" rid="F1">Fig. 1</xref>). Current analysis suggests that the diversity of ground beetles that are active under the snow cover is even several times higher than in supranivean fauna. The number of subnivean species active during the winter can be similar for months, whereas supranivean carabids occur more accidentally. As can be seen from <xref ref-type="table" rid="T1">Table 1</xref>, only a few species are regularly observed as being active during the whole winter and from many regions. For most species described as winter active only one observation of a single individual is recorded. A good example comes from a study by <xref ref-type="bibr" rid="B33">Kennedy (1994)</xref> who recorded, between 22 November and 4 April, at least 12 <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species (genus <italic><tp:taxon-name>Amara</tp:taxon-name></italic> was provided with no details about species number) in winter-wheat fields in Ireland. Among these percentages, only the proportion of <italic><tp:taxon-name>Phyla obtusa</tp:taxon-name></italic> was higher than all other recorded species – more than 70% of the caught individuals. Only three other species (<italic><tp:taxon-name>Metallina lampros</tp:taxon-name></italic>, <italic><tp:taxon-name>Philochthus aeneus</tp:taxon-name></italic> and <italic><tp:taxon-name>Trechus quadristriatus</tp:taxon-name></italic>) had a share higher than 5%. Similar results came from <xref ref-type="bibr" rid="B44">Murdoch (1967)</xref> who recorded 17 species. In this case only <italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic> and <italic><tp:taxon-name>Leistus terminatus</tp:taxon-name></italic> were caught in ‘high’ proportions: respectively 68,9% and 9,1%. <italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic> also clearly predominated among 18 ground beetle species found in winter by <xref ref-type="bibr" rid="B22">Greenslade (1965)</xref>. Dominance of only single species was noted by F<xref ref-type="bibr" rid="B19">latz and Thaler (1980</xref>; <italic><tp:taxon-name>Poecilus versicolor</tp:taxon-name></italic>) and Hanning et al. (2006; <italic><tp:taxon-name>Dromius quadrimaculatus</tp:taxon-name></italic>). All these results suggest that also among the epigeic ground beetle fauna some species are found occasionally, while at least several others can be classified as permanently winter active.</p>
<p>According to literature data, winter active carabid species are known both from forests and open habitats as well as from species living on tree trunks (<xref ref-type="table" rid="T1">Table 1</xref>). More<!--PageBreak-->over, <xref ref-type="bibr" rid="B33">Kennedy (1994)</xref> showed that at least some carabid species can be active during the winter period both during night and day. From these investigations it became clear that <italic><tp:taxon-name>Phyla obtusa</tp:taxon-name></italic> was a day active species from 2nd to 24th of January; unfortunately no data about temperature or other environmental factors were given.</p>
<p>In general, <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> can be divided into two main breeding groups: autumn breeders (eggs are laid during the last weeks of summer and first weeks of autumn) and spring breeders (eggs are laid from March to May). As a result of this division, winter and summer carabid larvae can be distinguished (<xref ref-type="bibr" rid="B39">Luff 1993</xref>). Usually winter larvae hatch from from September to November, and can be found (instars 1–3) throughout the winter and in the following spring period. Although the total number of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species that breed in the autumn period is much higher, at the moment larvae of 12 different species have been distinguished as winter active (<xref ref-type="bibr" rid="B38">Levander 1913</xref>, Näsmark 1954, <xref ref-type="bibr" rid="B49">Renken 1956</xref>, <xref ref-type="bibr" rid="B32">Kaczmarek 1958</xref>, <xref ref-type="bibr" rid="B22">Greenslade 1965</xref>, <xref ref-type="bibr" rid="B64">Weber 1965</xref>, <xref ref-type="bibr" rid="B44">Murdoch 1967</xref>, <xref ref-type="bibr" rid="B15">Evans 1969</xref>, <xref ref-type="bibr" rid="B39">Luff 1993</xref><bold>,</bold> <xref ref-type="bibr" rid="B61">Traugott 1998</xref>, this paper; <xref ref-type="table" rid="T1">Table 1</xref>). Among these, the occurrence of larvae of <italic><tp:taxon-name>Amara aulica</tp:taxon-name></italic>, <italic><tp:taxon-name>Calathus melanocephalus</tp:taxon-name></italic>, <italic><tp:taxon-name>Carabus coriaceus</tp:taxon-name></italic>, <italic><tp:taxon-name>Carabus problematicus</tp:taxon-name></italic>, <italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic>, <italic><tp:taxon-name>Pseudoophonus rufipes</tp:taxon-name></italic>, <italic><tp:taxon-name>Pterostichus melanarius</tp:taxon-name></italic>, and <italic><tp:taxon-name>Leistus</tp:taxon-name></italic> species can be explained as a result of autumn breeding (<xref ref-type="bibr" rid="B12">Betz 1992</xref>, <xref ref-type="bibr" rid="B39">Luff 1993</xref>, <xref ref-type="bibr" rid="B61">Traugott 1998</xref>). <xref ref-type="bibr" rid="B64">Weber (1965)</xref> and <xref ref-type="bibr" rid="B15">Evans (1969)</xref> did not provide any details on the identity of the <italic><tp:taxon-name>Carabus</tp:taxon-name></italic> and <italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic> larvae found during the winter period. However, these genera do have species that belong to autumnal breeders too (<xref ref-type="bibr" rid="B39">Luff 1993</xref>).</p>
<p><xref ref-type="bibr" rid="B39">Luff (1993<bold>)</bold></xref> suggested that winter larvae of ground beetles must survive not only low temperatures and food shortages, but also a long period of exposure to natural enemies, and possible flooding. He also noted that, especially at lower temperatures, some winter carabid larvae can survive without food for up to 30 days.</p>
<p>A zoogeographical analysis shows that the Central and northern European winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> most frequently belong to the Palaearctic fauna (54%). Interesting is that Euro-Siberian and Euro-Arctic species (groups that should be adapted evolutionary to low temperatures) made up only 14% of the recorded ground beetle species, while 12% of the species belong the Euro-Mediterranean fauna (<xref ref-type="fig" rid="F2">Fig. 2</xref>).</p>
<fig id="F1" position="float" orientation="portrait">
<label>Figure 1.</label>
<caption><p>Comparision of subnivean, supranivean and tree trunk fauna of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> from Central and Northern Europe during the winter season (based on different sources).</p></caption>
<graphic xlink:href="ZooKeys-100-517-g001.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</sec>
<sec sec-type="Role of carabids in the winter food chain"><title>Role of carabids in the winter food chain</title>
<p>High densities per square meter and high percentages of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> in winter active insect communities make this group an important source of food for insectivorous vertebrates, particularly shrews. Due to their very high metabolic rate these mammals must feed almost constantly to stay alive. They are active all year round, without a hibernation period in winter and their food requirement is 43% higher in winter than in summer (<xref ref-type="bibr" rid="B48">Randolph 1973</xref>). As indicated in the literature, shrews do not feed on hibernating invertebrates, but rather on winter-active species, including <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B1">Ackefors 1964</xref>, <xref ref-type="bibr" rid="B46">Pernetta 1977</xref>, <xref ref-type="bibr" rid="B9">Aitchison 1984</xref>, <xref ref-type="bibr" rid="B29">Itämies and Lindgren 1989</xref>). Ground beetles can be an attractive type of food for these mammals as they are present in relatively high densities - up to 23 individuals per square meter (<xref ref-type="bibr" rid="B33">Kennedy 1994</xref>). <xref ref-type="bibr" rid="B50">Rudge (1968)</xref> <!--PageBreak-->found that the percentage frequency of beetles in the diet of <italic><tp:taxon-name>Sorex araneus</tp:taxon-name></italic> varies from 66–72% in the autumn and spring. This increases up to 84% in the winter months. In this study only plants had a higher share during the autumn-spring period (percentage frequency 96–100%) and other small vertebrates in autumnal months (100%).</p>
<p>From the winter active Central and North European group of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic>, 79% of the species appear to be predators (<xref ref-type="table" rid="T1">Table 1</xref>). Among them there are both large zoophagous species hunting for various types of prey (e.g., <italic><tp:taxon-name>Carabus</tp:taxon-name></italic> species) and specialists collecting small but very abundant prey items, i.e., springtails and aphids (e.g., <xref ref-type="bibr" rid="B4">Aitchison 1978</xref>, <xref ref-type="bibr" rid="B8">1979</xref>, <xref ref-type="bibr" rid="B35">Leinaas 1981</xref>, <xref ref-type="bibr" rid="B36">1983</xref>, <xref ref-type="bibr" rid="B24">Hågvar 2000</xref>). Springtails are known as one of the most abundant subnivean invertebrate groups (e.g., <xref ref-type="bibr" rid="B45">Näsmark 1964</xref>, <xref ref-type="bibr" rid="B9">Aitchison 1984</xref>). They were regarded as an important food source in the winter active <italic><tp:taxon-name>Phyla obtusa</tp:taxon-name></italic> with a percentage frequency from 4 to 20% (<xref ref-type="bibr" rid="B33">Kennedy 1994</xref>). In the latter study <italic><tp:taxon-name>Phyla obtusa</tp:taxon-name></italic> was also noted as a predator of mites (8–30%) and aphids (4–33%) during January-March. Among winter active carabids, species belonging to the genera <italic><tp:taxon-name>Loricera</tp:taxon-name></italic>, <italic><tp:taxon-name>Notiophilus</tp:taxon-name></italic>, <italic><tp:taxon-name>Leistus</tp:taxon-name></italic>, and some smallspecies of <italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic> are also well known as predator of springtails. Most probably the winter activity of species belonging to <italic><tp:taxon-name>Dromius</tp:taxon-name></italic> s.l. (e.g., <xref ref-type="bibr" rid="B26">Hannig et al. 2006</xref>) group can be related to the activity of their usual type of prey, i.e., aphids. On the other hand, winter activity of omnivorous (5%) and phytophagous (16%) carabids can be explained by a relatively easy access to their food, i.e., dry or decaying wood, fungi, leaves and seeds.</p>
<p>Many <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species can change their diet according to the availability of food in the environment. Some predatory beetles (e.g., some <italic><tp:taxon-name>Carabus</tp:taxon-name></italic> species, <italic><tp:taxon-name>Pterostichus melanarius</tp:taxon-name></italic>, <italic><tp:taxon-name>Calathus fuscipes</tp:taxon-name></italic>, <italic><tp:taxon-name>Nebria brevicollis</tp:taxon-name></italic>) occasionally eat plant material. Also some typically phytophagous species (<italic><tp:taxon-name>Amara</tp:taxon-name></italic> spp., <italic><tp:taxon-name>Harpalus</tp:taxon-name></italic> spp., <italic><tp:taxon-name>Bradycellus</tp:taxon-name></italic> spp.) are able to change their diet to eggs and pupae of flies (<xref ref-type="bibr" rid="B60">Tischler 1971</xref>). When temperatures<!--PageBreak--> become too low, some species can stop feeding even if they are still active (<xref ref-type="bibr" rid="B11">Aitchison 2001</xref>). In extreme situations some beetles (including larvaal stages) can survive up to one month without food while remaining active (<xref ref-type="bibr" rid="B39">Luff 1993</xref>).</p>
<p>An important adaptation that protects winter active arthropods from freezing is non-feeding behaviour during lower temperatures (<xref ref-type="bibr" rid="B10">Aitchison 1987</xref>). The presence of food in the gut significantly increases the possibility of spontaneous freezing as ice nucleators are present in the food (<xref ref-type="bibr" rid="B51">Salt 1968</xref>). As a special adaptation to prevent freezing during eating at cold temperatures, external digestion can be seen in some <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species, including members of <italic><tp:taxon-name>Carabus</tp:taxon-name></italic>, <italic><tp:taxon-name>Cychrus</tp:taxon-name></italic>, <italic><tp:taxon-name>Pterostichus</tp:taxon-name></italic>, and ground beetle larvae (<xref ref-type="bibr" rid="B27">Hengeveld 1980a</xref>-b, <xref ref-type="bibr" rid="B14">Evans and Forsythe 1985</xref>). As was shown by <xref ref-type="bibr" rid="B10">Aitchison (1987)</xref>, who studied spiders, a group that feeds by means of external digestion, such behaviour allows the avoidance of consuming dust particles on which spontaneous ice formation can occur. One of the most common and abundant groups of winter active arthropods is Collembola, which is also a popular type of prey for some <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species. The study of <xref ref-type="bibr" rid="B4">Aitchison (1978)</xref> showed that springtails contain some cryoptotectans in their haemolymph allowing survival in cold temperatures. Feeding mechanisms observed in spiders and carabids suggest that these chemical compounds can possibly be transferred from a prey to a predator body during eating. As a result cryoprotectans of the prey may allow its predator to survive low temperatures.</p>
<fig id="F2" position="float" orientation="portrait">
<label>Figure 2.</label>
<caption><p>The relative zoogeographical structure of winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> (based on <xref ref-type="bibr" rid="B37">Leśniak 1988</xref>).</p></caption>
<graphic xlink:href="ZooKeys-100-517-g002.jpg" position="float" orientation="portrait" xlink:type="simple"/>
</fig>
</sec>
<sec sec-type="Weather conditions and winter activity of ground beetles"><title>Weather conditions and winter activity of ground beetles</title>
<p>Based on literature data we can assume that the activity of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> species decreases in late autumn. Activity will be lowest during the winter period, and increases in the early spring (e.g., <xref ref-type="bibr" rid="B15">Evans 1969</xref>, <xref ref-type="table" rid="T1">Table 1</xref>, <xref ref-type="fig" rid="F1">Fig. 1</xref>). The subnivean environment is characterized by a much higher number of carabid species compared to the supranivean one (<xref ref-type="fig" rid="F1">Fig. 1</xref>). This is observed in many other insect groups and is usually explained by the role of thermally isolated snow cover that protects the environment from wind and lethal temperatures (<xref ref-type="bibr" rid="B11">Aitchison 2001</xref>). Literature data and our own results show that <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> are active on the snow surface only from November to January, while subnivean activity occurred during the entire winter season. In general, ground beetles are only accidentally found on the snow cover and because of this, they should be classified as chionoxenes.</p>
<p>In the literature there are almost no data on the effects of weather factors on winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic>. A study by <xref ref-type="bibr" rid="B64">Weber (1965)</xref> suggests that air temperature rising from -2°C to +6°C increased activity of <italic><tp:taxon-name>Phyla obtusa</tp:taxon-name></italic> almost eight times. Similar observations were made by the same author for <italic><tp:taxon-name>Trechus quadristriatus</tp:taxon-name></italic>.</p>
<p>Interesting observations were made by Haning et al. (2006), who noted <italic><tp:taxon-name>Calodromius bifasciatus</tp:taxon-name></italic> to be active on tree trunks at -3°C and from -1 to +10°C, with males preferring lower temperatures than females (see also <xref ref-type="bibr" rid="B18">Felix and Van Wielink 2011</xref>). For supranivean active carabids, temperature data are known for only four species: <italic><tp:taxon-name>Dromius schneideri</tp:taxon-name></italic> was found at <italic>-</italic>1°C, <italic><tp:taxon-name>Pterostichus diligens</tp:taxon-name></italic> at +1°C(<xref ref-type="bibr" rid="B20">Frey 1913</xref>), <italic><tp:taxon-name>Agonum muelleri</tp:taxon-name></italic> at +2°C (<xref ref-type="bibr" rid="B58">Szulczewski 1947</xref>) and <italic><tp:taxon-name>Calathus micropterus</tp:taxon-name></italic> at -2°C (Soszyńska-Maj & Jaskuła unpublished).</p>
</sec>
</sec>
<sec sec-type="Conclusions"><title>Conclusions</title>
<p>Present knowledge on winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> from Central and Northern Europe is rather poor. Literature data are mostly from a few old papers, and usually were fragmentary. In total, 73 species have been recorded as active in winter, including 11 species belonging to 10 genera found on the snow surface, and 66 species from33 genera being subnivean. Four species were recorded for the first time as snow active and one as a subnivean carabid.</p>
<p>Ground beetles are one of the dominating <italic><tp:taxon-name>Coleoptera</tp:taxon-name></italic> groups in winter insect assemblage. The community of winter active <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> is composed of larvae and adult beetles, and consists of both epigeic species and species active on tree trunks. In general, winter active larvae are representatives of autumn breeders. A comparison of the supranivean and subnivean carabid fauna shows significant differences in species diversity. In the first group the number of species are five times lower than in the latter. It suggests that snow active species appear in supranivean microhabitats only accidentally, but they are known to be winter active in litter or soil environments. They should probably be classified as chionoxenes.</p>
<p>Winter activity of ground beetles decreases in late autumn, is lowest during mid-winter and increases in early spring. This might be correlated with weather conditions, especially air temperature. The present state of knowledge suggests that further studies are needed to confirm this hypothesis.</p>
<p>The high proportion of <italic><tp:taxon-name>Carabidae</tp:taxon-name></italic> in winter communities make this group an important food source in the diet of insectivorous mammals, especially shrews. On the other hand these carabids are predators, hunting springtails and other small winter active invertebrates.</p>
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<title>Acknowledgements</title>
<p>We would like to thank Michał Grabowski (University of Łódź) and two anonymous reviewers for their suggestions and comments.</p>
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<ref-list>
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