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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Forty years of carabid beetle research in Europe – from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation</title><author><name sortKey="Kotze, D Johan" sort="Kotze, D Johan" uniqKey="Kotze D" first="D. Johan" last="Kotze">D. Johan Kotze</name><affiliation><nlm:aff id="A1">University of Helsinki, Department of Environmental Sciences P.O. Box 65 (Biocenter 3, Viikinkaari 1), FI-00014 Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Brandmayr, Pietro" sort="Brandmayr, Pietro" uniqKey="Brandmayr P" first="Pietro" last="Brandmayr">Pietro Brandmayr</name><affiliation><nlm:aff id="A2">University of Calabria, Department of Ecology, Ponte Bucci, I-87036 Rende (CS), Italy</nlm:aff></affiliation></author><author><name sortKey="Casale, Achille" sort="Casale, Achille" uniqKey="Casale A" first="Achille" last="Casale">Achille Casale</name><affiliation><nlm:aff id="A3">Università di Sassari, Dipartimento di Zoologia e Genetica Evoluzionistica, Via Muroni 25, I-07100 Sassari, Italy</nlm:aff></affiliation></author><author><name sortKey="Dauffy Richard, Emmanuelle" sort="Dauffy Richard, Emmanuelle" uniqKey="Dauffy Richard E" first="Emmanuelle" last="Dauffy-Richard">Emmanuelle Dauffy-Richard</name><affiliation><nlm:aff id="A4">Cemagref UR EFNO, ‘Forest Ecosystems’, Domaine des Barres, F-45290 Nogent-sur-Vernisson, France</nlm:aff></affiliation></author><author><name sortKey="Dekoninck, Wouter" sort="Dekoninck, Wouter" uniqKey="Dekoninck W" first="Wouter" last="Dekoninck">Wouter Dekoninck</name><affiliation><nlm:aff id="A5">RBINS, Entomology Department, Vautierstraat 29, B-1000 Brussels, Belgium</nlm:aff></affiliation></author><author><name sortKey="Koivula, Matti J" sort="Koivula, Matti J" uniqKey="Koivula M" first="Matti J." last="Koivula">Matti J. Koivula</name><affiliation><nlm:aff id="A6">Finnish Forest Research Institute, PO Box 18, FI-01301 Vantaa, Finland</nlm:aff></affiliation></author><author><name sortKey="Lovei, Gabor L" sort="Lovei, Gabor L" uniqKey="Lovei G" first="Gábor L." last="Lövei">Gábor L. Lövei</name><affiliation><nlm:aff id="A7">Aarhus University, Faculty of Sciences & Technology, Flakkebjerg Research Centre, DK-4200 Slagelse, Denmark</nlm:aff></affiliation></author><author><name sortKey="Mossakowski, Dietrich" sort="Mossakowski, Dietrich" uniqKey="Mossakowski D" first="Dietrich" last="Mossakowski">Dietrich Mossakowski</name><affiliation><nlm:aff id="A8">Seeweg 10, D-23942 Groß Schwansee, Germany</nlm:aff></affiliation></author><author><name sortKey="Noordijk, Jinze" sort="Noordijk, Jinze" uniqKey="Noordijk J" first="Jinze" last="Noordijk">Jinze Noordijk</name><affiliation><nlm:aff id="A9">European Invertebrate Survey – Nederland, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Paarmann, Wilfried" sort="Paarmann, Wilfried" uniqKey="Paarmann W" first="Wilfried" last="Paarmann">Wilfried Paarmann</name><affiliation><nlm:aff id="A10">Meenser Weg 9, D-37124 Rosdorf/Atzenhausen, Germany</nlm:aff></affiliation></author><author><name sortKey="Pizzolotto, Roberto" sort="Pizzolotto, Roberto" uniqKey="Pizzolotto R" first="Roberto" last="Pizzolotto">Roberto Pizzolotto</name><affiliation><nlm:aff id="A2">University of Calabria, Department of Ecology, Ponte Bucci, I-87036 Rende (CS), Italy</nlm:aff></affiliation></author><author><name sortKey="Saska, Pavel" sort="Saska, Pavel" uniqKey="Saska P" first="Pavel" last="Saska">Pavel Saska</name><affiliation><nlm:aff id="A11">Czech University of Life Sciences in Prague, Department of Ecology, Kamycka 129, CZ-165 21 Prague 6 – Suchdol, Czech Republic</nlm:aff></affiliation></author><author><name sortKey="Schwerk, Axel" sort="Schwerk, Axel" uniqKey="Schwerk A" first="Axel" last="Schwerk">Axel Schwerk</name><affiliation><nlm:aff id="A12">Warsaw University of Life Sciences, Laboratory of Evaluation and Assessment of Natural Resources, Nowoursynowska street 166, PL-02-787 Warsaw, Poland</nlm:aff></affiliation></author><author><name sortKey="Serrano, Jose" sort="Serrano, Jose" uniqKey="Serrano J" first="José" last="Serrano">José Serrano</name><affiliation><nlm:aff id="A13">University of Murcia, Zoology and Physical Anthropology, Facultad Veterinaria, Campus Espinardo E-30071 Murcia, Spain</nlm:aff></affiliation></author><author><name sortKey="Szyszko, Jan" sort="Szyszko, Jan" uniqKey="Szyszko J" first="Jan" last="Szyszko">Jan Szyszko</name><affiliation><nlm:aff id="A12">Warsaw University of Life Sciences, Laboratory of Evaluation and Assessment of Natural Resources, Nowoursynowska street 166, PL-02-787 Warsaw, Poland</nlm:aff></affiliation></author><author><name sortKey="Taboada, Angela" sort="Taboada, Angela" uniqKey="Taboada A" first="Angela" last="Taboada">Angela Taboada</name><affiliation><nlm:aff id="A14">University of Leon, Department of Biodiversity & Environmental Management, Area of Ecology, Campus de Vegazana s/n, E-24071 Leon, Spain</nlm:aff></affiliation></author><author><name sortKey="Turin, Hans" sort="Turin, Hans" uniqKey="Turin H" first="Hans" last="Turin">Hans Turin</name><affiliation><nlm:aff id="A15">Loopkeverstichting (SFOC), Esdoorndreef 29, 6871 LK Renkum, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Venn, Stephen" sort="Venn, Stephen" uniqKey="Venn S" first="Stephen" last="Venn">Stephen Venn</name><affiliation><nlm:aff id="A1">University of Helsinki, Department of Environmental Sciences P.O. Box 65 (Biocenter 3, Viikinkaari 1), FI-00014 Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Vermeulen, Rikjan" sort="Vermeulen, Rikjan" uniqKey="Vermeulen R" first="Rikjan" last="Vermeulen">Rikjan Vermeulen</name><affiliation><nlm:aff id="A16">Stichting WBBS, Kanaaldijk 36, 9409 TV Loon, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Zetto, Tullia" sort="Zetto, Tullia" uniqKey="Zetto T" first="Tullia" last="Zetto">Tullia Zetto</name><affiliation><nlm:aff id="A2">University of Calabria, Department of Ecology, Ponte Bucci, I-87036 Rende (CS), Italy</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">21738408</idno><idno type="pmc">3131012</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3131012</idno><idno type="RBID">PMC:3131012</idno><idno type="doi">10.3897/zookeys.100.1523</idno><date when="2011">2011</date><idno type="wicri:Area/Pmc/Corpus">000449</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Forty years of carabid beetle research in Europe – from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation</title><author><name sortKey="Kotze, D Johan" sort="Kotze, D Johan" uniqKey="Kotze D" first="D. Johan" last="Kotze">D. Johan Kotze</name><affiliation><nlm:aff id="A1">University of Helsinki, Department of Environmental Sciences P.O. Box 65 (Biocenter 3, Viikinkaari 1), FI-00014 Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Brandmayr, Pietro" sort="Brandmayr, Pietro" uniqKey="Brandmayr P" first="Pietro" last="Brandmayr">Pietro Brandmayr</name><affiliation><nlm:aff id="A2">University of Calabria, Department of Ecology, Ponte Bucci, I-87036 Rende (CS), Italy</nlm:aff></affiliation></author><author><name sortKey="Casale, Achille" sort="Casale, Achille" uniqKey="Casale A" first="Achille" last="Casale">Achille Casale</name><affiliation><nlm:aff id="A3">Università di Sassari, Dipartimento di Zoologia e Genetica Evoluzionistica, Via Muroni 25, I-07100 Sassari, Italy</nlm:aff></affiliation></author><author><name sortKey="Dauffy Richard, Emmanuelle" sort="Dauffy Richard, Emmanuelle" uniqKey="Dauffy Richard E" first="Emmanuelle" last="Dauffy-Richard">Emmanuelle Dauffy-Richard</name><affiliation><nlm:aff id="A4">Cemagref UR EFNO, ‘Forest Ecosystems’, Domaine des Barres, F-45290 Nogent-sur-Vernisson, France</nlm:aff></affiliation></author><author><name sortKey="Dekoninck, Wouter" sort="Dekoninck, Wouter" uniqKey="Dekoninck W" first="Wouter" last="Dekoninck">Wouter Dekoninck</name><affiliation><nlm:aff id="A5">RBINS, Entomology Department, Vautierstraat 29, B-1000 Brussels, Belgium</nlm:aff></affiliation></author><author><name sortKey="Koivula, Matti J" sort="Koivula, Matti J" uniqKey="Koivula M" first="Matti J." last="Koivula">Matti J. Koivula</name><affiliation><nlm:aff id="A6">Finnish Forest Research Institute, PO Box 18, FI-01301 Vantaa, Finland</nlm:aff></affiliation></author><author><name sortKey="Lovei, Gabor L" sort="Lovei, Gabor L" uniqKey="Lovei G" first="Gábor L." last="Lövei">Gábor L. Lövei</name><affiliation><nlm:aff id="A7">Aarhus University, Faculty of Sciences & Technology, Flakkebjerg Research Centre, DK-4200 Slagelse, Denmark</nlm:aff></affiliation></author><author><name sortKey="Mossakowski, Dietrich" sort="Mossakowski, Dietrich" uniqKey="Mossakowski D" first="Dietrich" last="Mossakowski">Dietrich Mossakowski</name><affiliation><nlm:aff id="A8">Seeweg 10, D-23942 Groß Schwansee, Germany</nlm:aff></affiliation></author><author><name sortKey="Noordijk, Jinze" sort="Noordijk, Jinze" uniqKey="Noordijk J" first="Jinze" last="Noordijk">Jinze Noordijk</name><affiliation><nlm:aff id="A9">European Invertebrate Survey – Nederland, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Paarmann, Wilfried" sort="Paarmann, Wilfried" uniqKey="Paarmann W" first="Wilfried" last="Paarmann">Wilfried Paarmann</name><affiliation><nlm:aff id="A10">Meenser Weg 9, D-37124 Rosdorf/Atzenhausen, Germany</nlm:aff></affiliation></author><author><name sortKey="Pizzolotto, Roberto" sort="Pizzolotto, Roberto" uniqKey="Pizzolotto R" first="Roberto" last="Pizzolotto">Roberto Pizzolotto</name><affiliation><nlm:aff id="A2">University of Calabria, Department of Ecology, Ponte Bucci, I-87036 Rende (CS), Italy</nlm:aff></affiliation></author><author><name sortKey="Saska, Pavel" sort="Saska, Pavel" uniqKey="Saska P" first="Pavel" last="Saska">Pavel Saska</name><affiliation><nlm:aff id="A11">Czech University of Life Sciences in Prague, Department of Ecology, Kamycka 129, CZ-165 21 Prague 6 – Suchdol, Czech Republic</nlm:aff></affiliation></author><author><name sortKey="Schwerk, Axel" sort="Schwerk, Axel" uniqKey="Schwerk A" first="Axel" last="Schwerk">Axel Schwerk</name><affiliation><nlm:aff id="A12">Warsaw University of Life Sciences, Laboratory of Evaluation and Assessment of Natural Resources, Nowoursynowska street 166, PL-02-787 Warsaw, Poland</nlm:aff></affiliation></author><author><name sortKey="Serrano, Jose" sort="Serrano, Jose" uniqKey="Serrano J" first="José" last="Serrano">José Serrano</name><affiliation><nlm:aff id="A13">University of Murcia, Zoology and Physical Anthropology, Facultad Veterinaria, Campus Espinardo E-30071 Murcia, Spain</nlm:aff></affiliation></author><author><name sortKey="Szyszko, Jan" sort="Szyszko, Jan" uniqKey="Szyszko J" first="Jan" last="Szyszko">Jan Szyszko</name><affiliation><nlm:aff id="A12">Warsaw University of Life Sciences, Laboratory of Evaluation and Assessment of Natural Resources, Nowoursynowska street 166, PL-02-787 Warsaw, Poland</nlm:aff></affiliation></author><author><name sortKey="Taboada, Angela" sort="Taboada, Angela" uniqKey="Taboada A" first="Angela" last="Taboada">Angela Taboada</name><affiliation><nlm:aff id="A14">University of Leon, Department of Biodiversity & Environmental Management, Area of Ecology, Campus de Vegazana s/n, E-24071 Leon, Spain</nlm:aff></affiliation></author><author><name sortKey="Turin, Hans" sort="Turin, Hans" uniqKey="Turin H" first="Hans" last="Turin">Hans Turin</name><affiliation><nlm:aff id="A15">Loopkeverstichting (SFOC), Esdoorndreef 29, 6871 LK Renkum, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Venn, Stephen" sort="Venn, Stephen" uniqKey="Venn S" first="Stephen" last="Venn">Stephen Venn</name><affiliation><nlm:aff id="A1">University of Helsinki, Department of Environmental Sciences P.O. Box 65 (Biocenter 3, Viikinkaari 1), FI-00014 Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Vermeulen, Rikjan" sort="Vermeulen, Rikjan" uniqKey="Vermeulen R" first="Rikjan" last="Vermeulen">Rikjan Vermeulen</name><affiliation><nlm:aff id="A16">Stichting WBBS, Kanaaldijk 36, 9409 TV Loon, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Zetto, Tullia" sort="Zetto, Tullia" uniqKey="Zetto T" first="Tullia" last="Zetto">Tullia Zetto</name><affiliation><nlm:aff id="A2">University of Calabria, Department of Ecology, Ponte Bucci, I-87036 Rende (CS), Italy</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="2011">2011</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><label>Abstract</label><p>‘<italic>Carabidologists do it all</italic>’ (Niemelä 1996a) is a phrase with which most European carabidologists are familiar. Indeed, during the last half a century, professional and amateur entomologists have contributed enormously to our understanding of the basic biology of carabid beetles. The success of the field is in no small part due to regular European Carabidologists’ Meetings, which started in 1969 in Wijster, the Netherlands, with the 14th meeting again held in the Netherlands in 2009, celebrating the 40th anniversary of the first meeting and 50 years of long-term research in the Dwingelderveld. This paper offers a subjective summary of some of the major developments in carabidology since the 1960s. Taxonomy of the family <named-content content-type="taxon-name">Carabidae</named-content> is now reasonably established, and the application of modern taxonomic tools has brought up several surprises like elsewhere in the animal kingdom. Progress has been made on the ultimate and proximate factors of seasonality and timing of reproduction, which only exceptionally show non-seasonality. Triggers can be linked to evolutionary events and plausibly explained by the “taxon cycle” theory. Fairly little is still known about certain feeding preferences, including granivory and ants, as well as unique life history strategies, such as ectoparasitism and predation on higher taxa. The study of carabids has been instrumental in developing metapopulation theory (even if it was termed differently). Dispersal is one of the areas intensively studied, and results show an intricate interaction between walking and flying as the major mechanisms. The ecological study of carabids is still hampered by some unresolved questions about sampling and data evaluation. It is recognised that knowledge is uneven, especially concerning larvae and species in tropical areas. By their abundance and wide distribution, carabid beetles can be useful in population studies, bioindication, conservation biology and landscape ecology. Indeed, 40 years of carabidological research have provided so much data and insights, that among insects - and arguably most other terrestrial organisms - carabid beetles are one of the most worthwhile model groups for biological studies.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Adis, J" uniqKey="Adis J">J Adis</name></author><author><name sortKey="Paarmann, W" uniqKey="Paarmann W">W Paarmann</name></author><author><name sortKey="Erwin, Tl" uniqKey="Erwin T">TL Erwin</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Adis, J" uniqKey="Adis J">J Adis</name></author><author><name sortKey="Paarmann, W" uniqKey="Paarmann W">W Paarmann</name></author><author><name sortKey="Hofer, H" uniqKey="Hofer H">H Höfer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Alexandrovitch, Or" uniqKey="Alexandrovitch O">OR Alexandrovitch</name></author><author><name 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<front><journal-meta><journal-id journal-id-type="nlm-ta">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">21738408</article-id><article-id pub-id-type="pmc">3131012</article-id><article-id pub-id-type="doi">10.3897/zookeys.100.1523</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>Forty years of carabid beetle research in Europe – from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Kotze</surname><given-names>D. Johan</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Brandmayr</surname><given-names>Pietro</given-names></name><xref ref-type="aff" rid="A2">2</xref></contrib><contrib contrib-type="author"><name><surname>Casale</surname><given-names>Achille</given-names></name><xref ref-type="aff" rid="A3">3</xref></contrib><contrib contrib-type="author"><name><surname>Dauffy-Richard</surname><given-names>Emmanuelle</given-names></name><xref ref-type="aff" rid="A4">4</xref></contrib><contrib contrib-type="author"><name><surname>Dekoninck</surname><given-names>Wouter</given-names></name><xref ref-type="aff" rid="A5">5</xref></contrib><contrib contrib-type="author"><name><surname>Koivula</surname><given-names>Matti J.</given-names></name><xref ref-type="aff" rid="A6">6</xref></contrib><contrib contrib-type="author"><name><surname>Lövei</surname><given-names>Gábor L.</given-names></name><xref ref-type="aff" rid="A7">7</xref></contrib><contrib contrib-type="author"><name><surname>Mossakowski</surname><given-names>Dietrich</given-names></name><xref ref-type="aff" rid="A8">8</xref></contrib><contrib contrib-type="author"><name><surname>Noordijk</surname><given-names>Jinze</given-names></name><xref ref-type="aff" rid="A9">9</xref></contrib><contrib contrib-type="author"><name><surname>Paarmann</surname><given-names>Wilfried</given-names></name><xref ref-type="aff" rid="A10">10</xref></contrib><contrib contrib-type="author"><name><surname>Pizzolotto</surname><given-names>Roberto</given-names></name><xref ref-type="aff" rid="A2">2</xref></contrib><contrib contrib-type="author"><name><surname>Saska</surname><given-names>Pavel</given-names></name><xref ref-type="aff" rid="A11">11</xref></contrib><contrib contrib-type="author"><name><surname>Schwerk</surname><given-names>Axel</given-names></name><xref ref-type="aff" rid="A12">12</xref></contrib><contrib contrib-type="author"><name><surname>Serrano</surname><given-names>José</given-names></name><xref ref-type="aff" rid="A13">13</xref></contrib><contrib contrib-type="author"><name><surname>Szyszko</surname><given-names>Jan</given-names></name><xref ref-type="aff" rid="A12">12</xref></contrib><contrib contrib-type="author"><name><surname>Taboada</surname><given-names>Angela</given-names></name><xref ref-type="aff" rid="A14">14</xref></contrib><contrib contrib-type="author"><name><surname>Turin</surname><given-names>Hans</given-names></name><xref ref-type="aff" rid="A15">15</xref></contrib><contrib contrib-type="author"><name><surname>Venn</surname><given-names>Stephen</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Vermeulen</surname><given-names>Rikjan</given-names></name><xref ref-type="aff" rid="A16">16</xref></contrib><contrib contrib-type="author"><name><surname>Zetto</surname><given-names>Tullia</given-names></name><xref ref-type="aff" rid="A2">2</xref></contrib></contrib-group><aff id="A1"><label>1</label>University of Helsinki, Department of Environmental Sciences P.O. Box 65 (Biocenter 3, Viikinkaari 1), FI-00014 Helsinki, Finland</aff><aff id="A2"><label>2</label>University of Calabria, Department of Ecology, Ponte Bucci, I-87036 Rende (CS), Italy</aff><aff id="A3"><label>3</label>Università di Sassari, Dipartimento di Zoologia e Genetica Evoluzionistica, Via Muroni 25, I-07100 Sassari, Italy</aff><aff id="A4"><label>4</label>Cemagref UR EFNO, ‘Forest Ecosystems’, Domaine des Barres, F-45290 Nogent-sur-Vernisson, France</aff><aff id="A5"><label>5</label>RBINS, Entomology Department, Vautierstraat 29, B-1000 Brussels, Belgium</aff><aff id="A6"><label>6</label>Finnish Forest Research Institute, PO Box 18, FI-01301 Vantaa, Finland</aff><aff id="A7"><label>7</label>Aarhus University, Faculty of Sciences & Technology, Flakkebjerg Research Centre, DK-4200 Slagelse, Denmark</aff><aff id="A8"><label>8</label>Seeweg 10, D-23942 Groß Schwansee, Germany</aff><aff id="A9"><label>9</label>European Invertebrate Survey – Nederland, P.O. Box 9517, 2300 RA Leiden, The Netherlands</aff><aff id="A10"><label>10</label>Meenser Weg 9, D-37124 Rosdorf/Atzenhausen, Germany</aff><aff id="A11"><label>11</label>Czech University of Life Sciences in Prague, Department of Ecology, Kamycka 129, CZ-165 21 Prague 6 – Suchdol, Czech Republic</aff><aff id="A12"><label>12</label>Warsaw University of Life Sciences, Laboratory of Evaluation and Assessment of Natural Resources, Nowoursynowska street 166, PL-02-787 Warsaw, Poland</aff><aff id="A13"><label>13</label>University of Murcia, Zoology and Physical Anthropology, Facultad Veterinaria, Campus Espinardo E-30071 Murcia, Spain</aff><aff id="A14"><label>14</label>University of Leon, Department of Biodiversity & Environmental Management, Area of Ecology, Campus de Vegazana s/n, E-24071 Leon, Spain</aff><aff id="A15"><label>15</label>Loopkeverstichting (SFOC), Esdoorndreef 29, 6871 LK Renkum, The Netherlands</aff><aff id="A16"><label>16</label>Stichting WBBS, Kanaaldijk 36, 9409 TV Loon, The Netherlands</aff><author-notes><corresp>Corresponding author: D. Johan Kotze (<email>johan.kotze@helsinki.fi</email>).</corresp><fn fn-type="edited-by"><p>Academic editor: Lyubomir Penev</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>55</fpage><lpage>148</lpage><history><date date-type="received"><day>24</day><month>3</month><year>2011</year></date><date date-type="accepted"><day>24</day><month>3</month><year>2011</year></date></history><permissions><copyright-statement>D. Johan Kotze et al.</copyright-statement><license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/3.0"><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>‘<italic>Carabidologists do it all</italic>’ (Niemelä 1996a) is a phrase with which most European carabidologists are familiar. Indeed, during the last half a century, professional and amateur entomologists have contributed enormously to our understanding of the basic biology of carabid beetles. The success of the field is in no small part due to regular European Carabidologists’ Meetings, which started in 1969 in Wijster, the Netherlands, with the 14th meeting again held in the Netherlands in 2009, celebrating the 40th anniversary of the first meeting and 50 years of long-term research in the Dwingelderveld. This paper offers a subjective summary of some of the major developments in carabidology since the 1960s. Taxonomy of the family <named-content content-type="taxon-name">Carabidae</named-content> is now reasonably established, and the application of modern taxonomic tools has brought up several surprises like elsewhere in the animal kingdom. Progress has been made on the ultimate and proximate factors of seasonality and timing of reproduction, which only exceptionally show non-seasonality. Triggers can be linked to evolutionary events and plausibly explained by the “taxon cycle” theory. Fairly little is still known about certain feeding preferences, including granivory and ants, as well as unique life history strategies, such as ectoparasitism and predation on higher taxa. The study of carabids has been instrumental in developing metapopulation theory (even if it was termed differently). Dispersal is one of the areas intensively studied, and results show an intricate interaction between walking and flying as the major mechanisms. The ecological study of carabids is still hampered by some unresolved questions about sampling and data evaluation. It is recognised that knowledge is uneven, especially concerning larvae and species in tropical areas. By their abundance and wide distribution, carabid beetles can be useful in population studies, bioindication, conservation biology and landscape ecology. Indeed, 40 years of carabidological research have provided so much data and insights, that among insects - and arguably most other terrestrial organisms - carabid beetles are one of the most worthwhile model groups for biological studies.</p></abstract><kwd-group><label>Keywords</label><kwd>Carabidae</kwd><kwd>ground beetle</kwd><kwd>systematics</kwd><kwd>biology</kwd><kwd>life history</kwd><kwd>rhythms</kwd><kwd>seed feeding</kwd><kwd>ant feeding</kwd><kwd>ectoparasitism</kwd><kwd>predation on amphibians</kwd><kwd>dispersal</kwd><kwd>pitfall trapping</kwd><kwd>statistics</kwd><kwd>population dynamics</kwd><kwd>long-term research</kwd><kwd>bioindicators</kwd><kwd>conservation</kwd><kwd>habitat management</kwd><kwd>landscape ecology</kwd></kwd-group></article-meta></front><body><sec><title>1 Introduction</title><sec sec-type="1.1. General"><title>1.1. General</title><p>Carabid beetles are one of the best-known taxa in entomology. These beetles have been studied intensively by generations of coleopterists, who have clarified the taxonomy and phylogeny, geographic distribution, habitat associations and ecological requirements, life history strategies and adaptations, especially in Europe (e.g. <xref ref-type="bibr" rid="B241">Holdhaus and Lindroth 1939</xref>; <xref ref-type="bibr" rid="B450">Palmén 1944</xref>; <xref ref-type="bibr" rid="B321">Lindroth 1945a, b, 1949</xref>; <xref ref-type="bibr" rid="B551">Thiele 1977</xref>; <xref ref-type="bibr" rid="B35">Ball 1979</xref>; <xref ref-type="bibr" rid="B127">Desender 1986</xref>, <xref ref-type="bibr" rid="B136">Desender et al. 1994a</xref>; <xref ref-type="bibr" rid="B558">Turin 2000</xref>; <xref ref-type="bibr" rid="B339">Luff 2007</xref>).</p><p>This wealth of basic information has fostered a plethora of quantitative ecological studies. Indeed, the first European Carabidologists’ Meeting in Wijster, the Netherlands in 1969, touched upon one of the fascinating characteristics of carabid beetles – dispersal and dispersal power (<xref ref-type="bibr" rid="B112">Den Boer 1971</xref>). As a life history trait, dispersal has profound consequences for the dynamics and persistence of populations, the distribution and abundance of species and for community structure (<xref ref-type="bibr" rid="B148">Dieckmann et al. 1999</xref>). <pmc-comment>PageBreak</pmc-comment>Not surprisingly, a summary based on the 3rd International Carabidologists’ Meeting emphasised the role of dispersal in increasingly fragmented landscapes, and argued that much more knowledge on the effects of habitat loss and fragmentation on carabid beetle population dynamics is needed if sensible decisions are to be made regarding conservation and land-use (<xref ref-type="bibr" rid="B549">Thacker 1996</xref>).</p><p>But why study carabid beetles? The reasons are diverse: relatively stable taxonomy, high species richness, occurrence in most terrestrial environments and geographical areas, the availability of easy collection methods, known sensitivity to environmental changes, and perceived role as beneficial in agriculture (see <xref ref-type="bibr" rid="B106">Darlington 1943</xref>; <xref ref-type="bibr" rid="B334">Lövei and Sunderland 1996</xref>; <xref ref-type="bibr" rid="B473">Rainio and Niemelä 2003</xref>). Armed with such a diverse wealth of knowledge, many ecologists and taxonomists have turned to carabid beetles to test ecological research questions. In this paper we emphasise progress in some of the major fields in carabidology since the first European Carabidologists’ Meeting, 40 years ago.</p></sec><sec sec-type="1.2. Basic knowledge"><title>1.2. Basic knowledge</title><p>Modern disciplines in carabid beetle ecology, such as bioindication, conservation and habitat management, landscape ecology and urban ecology rely heavily on the work done by professional and amateur carabidologists from the more traditional fields of natural history, systematics and taxonomy. This species-rich family occurs in most terrestrial habitats and is found in the vegetation as well as high up in the trees and the canopy, not only in the tropics (<xref ref-type="bibr" rid="B16">Arndt 2005</xref>). This is probably the main reason why carabids are relatively well represented in collections around the world. In many regions, information on labels from these collections has been gathered in large databases. Combined with data from systematic sampling, such datasets enable profound faunistic work. These databases are increasingly elaborated and published as annotated checklists, red lists, catalogues and/or atlases. In combination with a clear taxonomy, mainly identification literature, these provide a sound basis for biogeographical, biological, ecological and experimental studies. <xref ref-type="table" rid="T1">Table 1</xref> shows an overview of the major publications for the European continent, which is covered well, although there is clearly need for updating in a few regions, mainly in the east (Romania, Hungary, Russia, Caucasus). In some cases, older works are mentioned in <xref ref-type="table" rid="T1">Table 1</xref>, which belong to antiquity and do not adequately cover the fauna of that region anymore (e.g. <xref ref-type="bibr" rid="B206">Ganglbauer 1892</xref>; <xref ref-type="bibr" rid="B13">Apfelbeck 1904</xref>; Porta 1923–1959). These older works are hardly in use for identification anymore. However, they still provide historical bases for modern identification works, which often have to be elaborated from numerous smaller keys or large revisions (e.g. Jeannel 1926–28; <xref ref-type="bibr" rid="B74">Breuning 1932–37</xref>), such as the keys to the <named-content content-type="taxon-name">Carabinae</named-content> (<xref ref-type="bibr" rid="B94">Casale et al. 1982</xref>) and to the supra-specific taxa of Italy (<xref ref-type="bibr" rid="B90">Casale 2005</xref>).</p><table-wrap id="T1" orientation="portrait" position="float"><label>Table 1.</label><caption><p>Overview of publications concerning the faunistics of ground beetles in Europe.</p></caption><table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1"><italic>Country</italic></th><th rowspan="1" colspan="1"><italic>Identification literature</italic></th><th rowspan="1" colspan="1"><italic>Checklist/Catalogue</italic></th><th rowspan="1" colspan="1"><italic>Atlas</italic></th></tr><tr><td rowspan="1" colspan="1">Albania</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B13">Apfelbeck 1904</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B218">Guéorguiev 2007</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Austria</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B358">Mandl 1972</xref>, <xref ref-type="bibr" rid="B359">1978</xref>; <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Baltic</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B221">Haberman 1968</xref>; <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B221">Haberman 1968</xref>; <xref ref-type="bibr" rid="B44">Barsevskis 2003</xref>; <xref ref-type="bibr" rid="B3">Alexandrovitch et al. 1996</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B221">Haberman 1968</xref></td></tr><tr><td rowspan="1" colspan="1">Belgium/Luxembourg</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B54">Boeken et al. 2002</xref>; <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref>; Muilwijk et al. (In prep.)</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B138">Desender et al. 1995</xref>; <xref ref-type="bibr" rid="B135">2008b</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B134">Desender et al. 2008a</xref></td></tr><tr><td rowspan="1" colspan="1">Bulgaria</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B13">Apfelbeck 1904</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B239">Hieke and Wrase 1988</xref>; <xref ref-type="bibr" rid="B219">Guéorguiev and Guéorguiev 1995</xref>; <xref ref-type="bibr" rid="B220">Guéorguiev et al. 1997</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Caucasus</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B256">Iablokov-Khnzorian 1976</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B302">Kryzhanovskij et al. 1995</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Czech Republic/ Slovakia</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B477">Reitter 1908</xref>; <xref ref-type="bibr" rid="B303">Kult 1947</xref>; <xref ref-type="bibr" rid="B252">Hurka 1996</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B252">Hurka 1996</xref>, <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B524">Skoupý 2004</xref></td></tr><tr><td rowspan="1" colspan="1">Denmark</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B227">Hansen 1968</xref>; <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B39">Bangsholt 1983</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B39">Bangsholt 1983</xref></td></tr><tr><td rowspan="1" colspan="1">Fennoscandia</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B325">Lindroth 1985-1986</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B321">Lindroth 1945a</xref>, <xref ref-type="bibr" rid="B323">1960</xref>, <xref ref-type="bibr" rid="B325">1985-86</xref>; <xref ref-type="bibr" rid="B528">Strand 1970</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B321">Lindroth 1945b</xref></td></tr><tr><td rowspan="1" colspan="1">France</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B262">Jeannel 1941-1942, 1949</xref>; <xref ref-type="bibr" rid="B192">Forel and Leplat 1995</xref>, <xref ref-type="bibr" rid="B194">2001</xref>, <xref ref-type="bibr" rid="B195">2003</xref>, <xref ref-type="bibr" rid="B196">2005</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B262">Jeannel 1941-1942, 1949</xref>; <xref ref-type="bibr" rid="B192">Forel and Leplat 1995</xref>, <xref ref-type="bibr" rid="B194">2001</xref>, <xref ref-type="bibr" rid="B195">2003</xref>, <xref ref-type="bibr" rid="B196">2005</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B99">Coulon et al. 2000</xref>; <xref ref-type="bibr" rid="B192">Forel and Leplat 1995</xref>, <xref ref-type="bibr" rid="B194">2001</xref>, <xref ref-type="bibr" rid="B195">2003</xref>, <xref ref-type="bibr" rid="B196">2005</xref>; <xref ref-type="bibr" rid="B87">Callot and Schott 1993</xref></td></tr><tr><td rowspan="1" colspan="1">Germany</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B477">Reitter 1908</xref>; <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref>; <xref ref-type="bibr" rid="B590">Wachmann et al. 1995</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B212">Gebert 2006</xref></td></tr><tr><td rowspan="1" colspan="1">Great Britain</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B339">Luff 2007</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B255">Hyman and Parsons 1992</xref>; <xref ref-type="bibr" rid="B339">Luff 2007</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B337">Luff 1998</xref></td></tr><tr><td rowspan="1" colspan="1">Greece</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B13">Apfelbeck 1904</xref>; <xref ref-type="bibr" rid="B19">Arndt et al. (in press)</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B19">Arndt et al. (in press)</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Hungary</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B101">Csiki 1946</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B101">Csiki 1946</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Iberia</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B193">Forel and Leplat 1998</xref>; <xref ref-type="bibr" rid="B237">Herrera and Arricibita 1990</xref>; <xref ref-type="bibr" rid="B347">Machado 1992</xref> (Canary Islands); <xref ref-type="bibr" rid="B429">Ortuño and Toribio 2005</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B237">Herrera and Arricibita 1990</xref>; <xref ref-type="bibr" rid="B601">Zaballos and Jeanne 1994</xref>; <xref ref-type="bibr" rid="B515">Serrano 2003</xref>; <xref ref-type="bibr" rid="B347">Machado 1992</xref> (Canary Islands)</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B237">Herrera and Arricibita 1990</xref>; <xref ref-type="bibr" rid="B429">Ortuño and Toribio 2005</xref></td></tr><tr><td rowspan="1" colspan="1">Iceland</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B325">Lindroth 1985, 1986</xref>; <xref ref-type="bibr" rid="B339">Luff 2007</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B320">Lindroth 1931</xref>; <xref ref-type="bibr" rid="B311">Larsson and Gigja 1959</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Ireland Italy</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B6">Anderson et al. 2000</xref><xref ref-type="bibr" rid="B466">Porta 1923-1959</xref>; <xref ref-type="bibr" rid="B94">Casale et al. 1982</xref>; <xref ref-type="bibr" rid="B90">Casale 2005</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B6">Anderson et al. 2000</xref><xref ref-type="bibr" rid="B341">Luigioni 1929</xref>; <xref ref-type="bibr" rid="B352">Magistretti 1965</xref>; <xref ref-type="bibr" rid="B582">Vigna Taglianti 1993</xref>, <xref ref-type="bibr" rid="B583">2005</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B6">Anderson et al. 2000</xref><xref ref-type="bibr" rid="B94">Casale et al. 1982</xref>, <xref ref-type="bibr" rid="B95">2007</xref>; CK Map 2006</td></tr><tr><td rowspan="1" colspan="1">Moldova/Romania</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B101">Csiki 1946</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B302">Kryzhanovskij et al. 1995</xref>; <xref ref-type="bibr" rid="B397">Neculiseanu and Matalin 2000</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">The Netherlands</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B54">Boeken et al. 2002</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B62">Brakman 1966</xref>; <xref ref-type="bibr" rid="B558">Turin 2000</xref>; <xref ref-type="bibr" rid="B389">Muilwijk and Felix 2010</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B558">Turin 2000</xref></td></tr><tr><td rowspan="1" colspan="1">Poland</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B80">Burakowski et al. 1973-1974</xref>; <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Russia/Belarus</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B301">Kryzhanovskij 1983</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B302">Kryzhanovskij et al. 1995</xref>; <xref ref-type="bibr" rid="B3">Alexandrovitch et al. 1996</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Switzerland</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B360">Marggi 1992</xref>; <xref ref-type="bibr" rid="B392">Müller-Motzfeld 2004</xref>; <xref ref-type="bibr" rid="B342">Luka et al. 2009</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B360">Marggi 1992</xref>; <xref ref-type="bibr" rid="B342">Luka et al. 2009</xref></td></tr><tr><td rowspan="1" colspan="1">Ukraine</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B301">Kryzhanovskij 1983</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B302">Kryzhanovskij et al. 1995</xref>; <xref ref-type="bibr" rid="B472">Putchkov 2011</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Former Yugoslavia</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B13">Apfelbeck 1904</xref></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B155">Drovenik 1999</xref></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">Europe, general</td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B206">Ganglbauer 1892</xref>; <xref ref-type="bibr" rid="B157">Du Chatenet 1986</xref>; <xref ref-type="bibr" rid="B556">Trautner and Geigenmüller 1987</xref>; <named-content content-type="taxon-name">Eurocarabidae</named-content>: <ext-link ext-link-type="uri" xlink:href="http://www.eurocarabidae.de">http://www.eurocarabidae.de</ext-link></td><td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B557">Turin 1981</xref>; <xref ref-type="bibr" rid="B302">Kryzhanovskij et al. 1995</xref>; <xref ref-type="bibr" rid="B327">Löbl and Smetana 2003</xref>; Fauna Europea: <ext-link ext-link-type="uri" xlink:href="http://www.faunaeur.org">http://www.faunaeur.org</ext-link></td><td rowspan="1" colspan="1">European maps: <xref ref-type="bibr" rid="B157">Du Chatenet 1986</xref> (189 European species); <xref ref-type="bibr" rid="B558">Turin 2000</xref> (380 Dutch species), <xref ref-type="bibr" rid="B561">Turin et al. 2003</xref> (<italic><named-content content-type="taxon-name">Carabus</named-content></italic>: 135 species); Fauna Europea: <ext-link ext-link-type="uri" xlink:href="http://www.faunaeur.org">http://www.faunaeur.org</ext-link></td></tr></tbody></table></table-wrap><p>A sound basic list of the <named-content content-type="taxon-name">Carabidae</named-content> of the world is the recent checklist published by <xref ref-type="bibr" rid="B330">Lorenz (2005)</xref> and a catalogue with distributional data is available for the Palaearctic<pmc-comment>PageBreak</pmc-comment> region as a whole (<xref ref-type="bibr" rid="B327">Löbl and Smetana 2003</xref>). Furthermore, many recent checklists and catalogues are available (concerning Europe, see some examples in <xref ref-type="table" rid="T1">Table 1</xref>). In particular, <xref ref-type="bibr" rid="B302">Kryzhanovskij et al. (1995)</xref> provided detailed information on the carabid fauna of Russia and adjacent countries (including central-Asiatic). In the Western Hemisphere (the Americas), detailed information is available, especially for the regions north of Mexico (<xref ref-type="bibr" rid="B324">Lindroth 1961–1969</xref>; <xref ref-type="bibr" rid="B37">Ball and Bousquet 2001</xref>; <xref ref-type="bibr" rid="B309">Larochelle and Larivière 2003</xref>; <xref ref-type="bibr" rid="B177">Erwin 2007</xref>; <xref ref-type="bibr" rid="B180">Erwin and Pearson 2008</xref>), or will soon be (Erwin in preparation), but in many tropical areas of Central and South America, many genera and species remain undescribed. Other geographical areas are less well known. Asia, as a huge continent is relatively well-known in some parts, such as Siberia, Near and Middle East and especially Japan (e.g. <xref ref-type="bibr" rid="B222">Habu 1967</xref>, 1973, 1978), whereas immense areas are a “work in progress” (China, The Himalayas and South-East Asia). Africa is well-known in some northern countries, in particular Morocco, Algeria and Tunisia, thanks to the contributions of specialists like <xref ref-type="bibr" rid="B12">Antoine (1955–1962)</xref>, <xref ref-type="bibr" rid="B47">Bedel (1899–1900)</xref> and <xref ref-type="bibr" rid="B284">Kocher (1963)</xref>. Nevertheless, in spite of the numerous papers published by Alluaud, Basilewsky, Jeannel and others, the sub-Saharan (tropical) part of the continent needs more investigation. Australia, thanks to the C.S.I.R.O. has one of the best-organised services of insect collections, and is covered by catalogues and revisions, of which we highlight the catalogue by <xref ref-type="bibr" rid="B312">Lawrence et al. (1987)</xref>. But also, recent investigations allowed the discovery of many new genera and species, including impressive, large sized <italic><named-content content-type="taxon-name">Pamborus</named-content></italic> species.</p><p>Finally, remote islands and archipelagos such as like Madagascar, Papua-New Guinea and Galápagos, for instance, have been carefully investigated by specialists like Jeannel, Darlington and Desender, respectively, but produce many new discoveries every year.</p><p>In the world catalogues of (<xref ref-type="bibr" rid="B329">Lorenz (1998</xref>, 2005) more than 35 000 ground beetle species have been listed. An estimated number of 40 000 species, which is more than 10 times the number of described mammals, has often been mentioned (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>; <xref ref-type="bibr" rid="B419">Noonan 1985</xref>). Currently, approximately 38 600 valid names occur worldwide (based on <xref ref-type="bibr" rid="B330">Lorenz 2005</xref> and an estimate of approximately 100 additional new species every year). For the Western Hemisphere only, the species count currently stands at 9 374 (Terry Erwin in litt.).</p><p>More in line with the meetings are a number of thematic treatments, but again the listed works are only examples. For a more complete and thematically arranged overview of significant work in carabidology, we refer to the excellent introduction to the proceedings of the Symposium on Phylogeny and Classification of Caraboidea by <xref ref-type="bibr" rid="B38">Ball et al. (1998)</xref>. Worth mentioning for European carabidology are the publications of the German “Gesellschaft für Angewandte Carabidologie” (GAC) with special reference to habitat studies, such as carabid beetles in river meadow habitats (GAC<xref ref-type="bibr" rid="B203"> 1999</xref>), in forests (GAC<xref ref-type="bibr" rid="B204"> 2001</xref>) and in xerothermic habitats (GAC<xref ref-type="bibr" rid="B205"> 2004</xref>). The GAC provides many carabidological papers in open access (see <ext-link ext-link-type="uri" xlink:href="http://www.laufkaefer.de/gac">http://www.laufkaefer.de/gac</ext-link>). Other published thematic studies, often including compilations of numerous papers from various authors, concern, amongst others: biotopes (<xref ref-type="bibr" rid="B238">Heydemann 1962</xref>; <xref ref-type="bibr" rid="B504">Schjøtz-Christensen 1965</xref>), larvae (<xref ref-type="bibr" rid="B68">Brandmayr and Zetto Brandmayr 1982</xref>; <xref ref-type="bibr" rid="B14">Arndt 1991</xref>; <xref ref-type="bibr" rid="B336">Luff 1993</xref>), biol<pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>ogy and periodicity (<xref ref-type="bibr" rid="B310">Larsson 1939</xref>), agroecology (Holland 2002), biogeography (<xref ref-type="bibr" rid="B36">Ball 1985</xref>; <xref ref-type="bibr" rid="B420">Noonan et al. 1992</xref>), dispersal ecology (<xref ref-type="bibr" rid="B450">Palmén 1944</xref>; <xref ref-type="bibr" rid="B113">Den Boer 1977</xref>; <xref ref-type="bibr" rid="B31">Baars 1982</xref>; <xref ref-type="bibr" rid="B129">Desender 1989b</xref>; <xref ref-type="bibr" rid="B29">Aukema 1995</xref>), morphology (<xref ref-type="bibr" rid="B519">Sharova 1981</xref>; <xref ref-type="bibr" rid="B143">Deuve 1993</xref>) and phylogeny (<xref ref-type="bibr" rid="B38">Ball et al. 1998</xref>). This listing is not exhaustive, especially in the fields of genetics and molecular biology, which are growing rapidly. We conclude with the classical works <italic>Die Fennoskandischen Carabidae</italic> (<xref ref-type="bibr" rid="B321">Lindroth 1945a, b, 1949</xref>, re-published in English as <xref ref-type="bibr" rid="B326">Lindroth 1988, 1992a, b</xref>) and <italic>Carabid beetles in their environments</italic> (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>). These inspired many carabidologists and have been, for many students, the starting point of their enthusiasm.</p></sec><sec sec-type="1.3. European Carabidologists’ Meetings (ECMs)"><title>1.3. European Carabidologists’ Meetings (ECMs)</title><p>In 1959, Piet den Boer, a zoologist at the Biological Station in Wijster, started pitfall trapping at several locations in the Dwingelderveld, a large area of heathland. His purpose was to test the model proposed by <xref ref-type="bibr" rid="B8">Andrewartha and Birch (1954)</xref>, in which animal populations could be thought of as sets of smaller local populations which periodically become extinct, their sites being subsequently reoccupied. This became known (and fashionable) under the term “metapopulation” (<xref ref-type="bibr" rid="B315">Levins 1970</xref>). By using carabid beetles as test organisms, Den Boer was able to show that in a large area many local populations or interacting groups fluctuate in numbers of individuals in space and time, developing his theory of ‘spreading of risk’ (<xref ref-type="bibr" rid="B110">Den Boer 1968</xref>). According to this theory, species occupying large areas survive more easily because the reproductive success of each separate (but interacting) group differs at different places. Dispersal between these interacting groups stabilises the number of individuals in the whole population through time. Local extinctions may occur but the chances of extinction of the entire population are minimised (<xref ref-type="bibr" rid="B111">Den Boer 1970</xref>). Den Boer eagerly wanted to discuss this topic with other carabid beetle specialists, in particular with Carl Lindroth from Sweden, who studied the significance of dispersal and Hans-Ulrich Thiele from Germany, who studied the reproduction of these animals. Consequently in 1969, a number of eminent European carabidologists were invited to Wijster. This select group of researchers focused on the topic of dispersal and the dispersal power of carabid beetles (<xref ref-type="fig" rid="F1">Fig. 1a</xref>). In 1973, Thiele invited a number of carabidologists to Rees-Grietherbush, a field station of the University of Cologne. This second ECM appeared to be an informal one and no proceedings volume was published. However, it resulted in the organisation of a now official third ECM, also at Rees-Grietherbush, by Thiele and his colleague Friedrich Weber in 1978. Most participants were German or Dutch, though Pietro Brandmayr from Italy was also present. The proceedings entitled ‘On the evolution and behaviour of carabid beetles’ was dedicated to Lindroth, who passed away in early 1979. In 1981, Weber took the initiative and organised the fourth ECM at Haus Rothenberge (Münster), on the theme ‘The synthesis of field study and laboratory experiments’. Thiele presented a lecture but his contribution fo<pmc-comment>PageBreak</pmc-comment>r the proceedings was never received. The proceedings, dedicated to Thiele, was published after his death in 1983.</p><fig id="F1" orientation="portrait" position="float"><label>Figure 1a.</label><caption><p>Participants of the first European Carabidologist Meeting in Wijster, 1969. From left to right: Vlijm, Van der Aart, Lindroth, Stein, Wijmans, Hengeveld, Palmén, Van Dijk, Richter, Venema, Mook, Thiele, Tjallingii, Den Boer, Haeck, Neumann, Meijer.</p></caption><graphic xlink:href="ZooKeys-100-055-g001"></graphic></fig><p>The first four meetings were followed by meetings organised across Europe (Table 2). As a result of political changes in Eastern Europe since the 1990s, the ECMs attained a more ‘complete’ European character. Not only did it become easier for scientists from Eastern Europe to attend these meetings, they also started to organise them. Even more noticeably during recent decades, carabidologists from beyond Europe regularly started to participate in the ECMs. Besides the official ECMs, there have been a few separate carabid beetle meetings in Europe (<xref ref-type="table" rid="T2">Table 2</xref>). Two of these (Hamburg in 1984 and Kauniainen in 1995) were not official ECM meetings, though they were mainly attended by the same carabidologists who regularly attend ECMs. The fourteen proceedings from the major ground beetle meetings that have been published before the present volume (see <xref ref-type="fig" rid="F2">Fig. 1b</xref>-<xref ref-type="fig" rid="F3">c</xref>, <xref ref-type="table" rid="T2">Table 2</xref>), comprise together more than 400 articles covering a wide range of topics. A rough classification of the articles leads to the following summary: Habitat preference, community ecology was the topic of 84 papers, Biology (development, preferences, etc.) of 55, Population biology - 46, Nature conservation - 35, Agro-ecology - 34, Dispersal ecology - 33, Evolutionary biology, phylogeny - 22, Morphology - 15, Ecology, general - 13, Genetics - 13, Biogeography - 11, Taxonomy - 11, Method-development - 10, Rest – 10, Faunistics - 9, and Palaeontology - 2. A similar series of meetings and proceedings started in America with the publication of the First International Symposium of Carabidology (<xref ref-type="bibr" rid="B178">Erwin et al. 1979</xref>). In<xref ref-type="bibr" rid="B203"> 1999</xref>, a volume consisting mainly of taxonomic papers was published, dedicated to the memory of Oleg L. Kryzhanovskij (Zamotailov and Sciaky<xref ref-type="bibr" rid="B203"> 1999</xref>).</p><fig id="F2" orientation="portrait" position="float"><label>Figure 1b.</label><caption><p>Front covers of the first European meetings, ECM 1–8 and that of Hamburg 1984 (centre cover) (see also <xref ref-type="table" rid="T2">Table 2</xref>).</p></caption><graphic xlink:href="ZooKeys-100-055-g002"></graphic></fig><fig id="F3" orientation="portrait" position="float"><label>Figure 1c.</label><caption><p>Front covers of the last five ECMs and of a few major carabidology publications (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>; <xref ref-type="bibr" rid="B38">Ball et al. 1998</xref>; <xref ref-type="bibr" rid="B178">Erwin et al. 1979</xref>; <xref ref-type="bibr" rid="B420">Noonan et al. 1992</xref>) (see also <xref ref-type="table" rid="T2">Table 2</xref>).</p></caption><graphic xlink:href="ZooKeys-100-055-g003"></graphic></fig><table-wrap id="T2" orientation="portrait" position="float"><label>Table 2.</label><caption><p>The year, location, title and editors of all the European Carabidologists’ Meetings.</p></caption><table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1"><italic>Year</italic></th><th rowspan="1" colspan="1"><italic>Location</italic></th><th rowspan="1" colspan="1"><italic>Proceedings</italic></th></tr><tr><td rowspan="1" colspan="1">1969</td><td rowspan="1" colspan="1">Wijster, The Netherlands (ECM 1)</td><td rowspan="1" colspan="1">1971. Dispersal and dispersal power of carabid beetles (Den Boer)</td></tr><tr><td rowspan="1" colspan="1">1973</td><td rowspan="1" colspan="1">Rees-Grietherbush, Germany (ECM 2)</td><td rowspan="1" colspan="1">None</td></tr><tr><td rowspan="1" colspan="1">1978</td><td rowspan="1" colspan="1">Rees-Grietherbush, Germany (ECM 3)</td><td rowspan="1" colspan="1">1979. On the evolution of behaviour in carabid beetles (Den Boer et al.)</td></tr><tr><td rowspan="1" colspan="1">1981</td><td rowspan="1" colspan="1">Münster, Germany (ECM 4)</td><td rowspan="1" colspan="1">1983. The synthesis of field study and laboratory experiments (Brandmayr et al.)</td></tr><tr><td rowspan="1" colspan="1">1982</td><td rowspan="1" colspan="1">Stara Brda Pilska, Poland(ECM 5)</td><td rowspan="1" colspan="1">1986a. Feeding behaviour and accessibility of food for carabid beetles (Den Boer et al.)</td></tr><tr><td rowspan="1" colspan="1">1984</td><td rowspan="1" colspan="1">Hamburg, Germany (17th International Entomological Congress)</td><td rowspan="1" colspan="1">1986b. Carabid beetles, their adaptations and dynamics (Den Boer et al.)</td></tr><tr><td rowspan="1" colspan="1">1986</td><td rowspan="1" colspan="1">Balatonalmadi, Hungary (ECM 6)</td><td rowspan="1" colspan="1">1987. Proceedings of the 6th ECM (Den Boer et al.)</td></tr><tr><td rowspan="1" colspan="1">1989</td><td rowspan="1" colspan="1">London, United Kingdom (ECM 7)</td><td rowspan="1" colspan="1">1990. The role of ground beetles in ecological and environmental studies (Stork)</td></tr><tr><td rowspan="1" colspan="1">1992</td><td rowspan="1" colspan="1">Louvain la Neuve, Belgium (ECM 8)</td><td rowspan="1" colspan="1">1994a. Carabid beetles, ecology and evolution (Desender et al.)</td></tr><tr><td rowspan="1" colspan="1">1995</td><td rowspan="1" colspan="1">Kauniainen, Finland (3rd International Carabidology Congress)</td><td rowspan="1" colspan="1">1996b. Population biology and conservation of carabid beetles (Niemelä)</td></tr><tr><td rowspan="1" colspan="1">1998</td><td rowspan="1" colspan="1">Camigliatello, Italy (ECM 9)</td><td rowspan="1" colspan="1">2000. Natural history and applied ecology of carabid beetles (Brandmayr et al.)</td></tr><tr><td rowspan="1" colspan="1">2001</td><td rowspan="1" colspan="1">Tuczno, Poland (ECM 10)</td><td rowspan="1" colspan="1">2002. How to protect or what we know about carabid beetles (Szyszko et al.)</td></tr><tr><td rowspan="1" colspan="1">2003</td><td rowspan="1" colspan="1">Århus, Denmark (ECM 11)</td><td rowspan="1" colspan="1">2005. European Carabidology 2003 (Lövei and Toft)</td></tr><tr><td rowspan="1" colspan="1">2005</td><td rowspan="1" colspan="1">Murcia, Spain (ECM 12)</td><td rowspan="1" colspan="1">2006. Proceedings of the XII ECM; ground beetles as a key group for biodiversity conservation studies in Europe (Serrano et al.)</td></tr><tr><td rowspan="1" colspan="1">2007</td><td rowspan="1" colspan="1">Blagoevgrad, Bulgaria (ECM 13)</td><td rowspan="1" colspan="1">2008. Back to the roots and back to the future. Towards a new synthesis between taxonomic, ecological and biogeographical approaches in carabidology (Penev et al.)</td></tr><tr><td rowspan="1" colspan="1">2009</td><td rowspan="1" colspan="1">Westerbork, Netherlands (ECM 14)</td><td rowspan="1" colspan="1">2011. Present volume (Kotze et al.)</td></tr></tbody></table></table-wrap><p><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>In 2009, the 14th ECM returned to the starting grounds in the Netherlands and was attended by five participants of the first ECM: Piet den Boer, Jaap Haeck, Rob Hengeveld, Jan Meijer and Theo van Dijk. The participants visited the permanent sampling plots in the Dwingelderveld and Mantingerveld, started 50 years earlier.</p></sec></sec><sec><title>2 Systematics, phylogeny and evolution</title><sec sec-type="2.1. Overview"><title>2.1. Overview</title><p>Regular carabidologists’ meetings have contributed significantly to our understanding of carabid phylogeny, evolution and systematics, as evidenced by the presentation of more than 60 papers on these topics. Progress has been made at different taxonom<pmc-comment>PageBreak</pmc-comment>ic ranks and in different fields of carabid systematics. At present, the integrative approach of combining morphology, molecular systematics, ethology, ecology, geographic distribution, etc., as well as the use of bioinformatics, is recognised as the best framework for solving the challenges still faced by carabidologists (<xref ref-type="bibr" rid="B24">Assmann et al. 2008</xref>), and by animal taxonomists in general.</p><p>What follows is a short overview of recent advances in carabid beetle systematics, concentrating on literature presented at ECMs and the international congresses mentioned above. As the main aim of this section is to present a general overview, only some of the main papers with a wide scope are cited.</p></sec><sec sec-type="2.2. General outline on systematics and phylogeny of the Carabidae"><title>2.2. General outline on systematics and phylogeny of the <named-content content-type="taxon-name">Carabidae</named-content></title><p><xref ref-type="bibr" rid="B35">Ball (1979)</xref> showed that the classification of <named-content content-type="taxon-name">Carabidae</named-content> is mostly based on morphological characters and that it includes both clade-based and grade-based criteria; classifications differ depending on the importance given to one or the other criterion. After this seminal revision, few advances have been made to unify the criteria to elect <named-content content-type="taxon-name">Caraboidea</named-content> (splitters) or <named-content content-type="taxon-name">Carabidae</named-content> (lumpers), and the same holds true for other high-ranked taxa. A practical synthesis of these ideas was presented by <xref ref-type="bibr" rid="B394">Nagel (1979a)</xref>, while <xref ref-type="bibr" rid="B38">Ball et al. (1998)</xref> and <xref ref-type="bibr" rid="B24">Assmann et al. (2008)</xref> revised the issue in depth. These two last-mentioned papers highlighted the need for an integrative approach to morphology, morphometrics and molecular systematics as the appropriate way of finding rapid solutions for challenging problems.</p></sec><sec sec-type="2.3. Within-species diversity"><title>2.3. Within-species diversity</title><p>An electrophoretic study on 14 Pyrenean populations of <italic><named-content content-type="taxon-name">Carabus punctatoauratus</named-content></italic> (<xref ref-type="bibr" rid="B22">Assmann 1990</xref>) revealed that the Pyrenees probably hosts an isolated relict population for this species, and that bottlenecks have affected western, central and eastern populations differentially. Subtle differences at a micro-geographic scale have also been shaped by small bottleneck phenomena in this species with low dispersal power.</p><p>Range expansion of <italic><named-content content-type="taxon-name">Carabus auronitens</named-content></italic> during the 19th century has allowed gene flow between populations in the surroundings of Münster, Germany, as evidenced by an electrophoretic study of 19 populations that showed a steep gradient of slow and fast alleles (<xref ref-type="bibr" rid="B548">Terlutter 1990</xref>). The high dispersal power of this species accounts for the observed allelic gradient (esterase-encoding gene) from source areas to recently colonised areas (<xref ref-type="bibr" rid="B405">Niehues et al. 1996</xref>). <xref ref-type="bibr" rid="B26">Assmann et al. (1994)</xref> showed that present-day populations of this species originated from three major refuges in southern France and that these putative core populations have contributed differentially to postglacial range expansion of the species.</p><p><xref ref-type="bibr" rid="B21">Ashworth (1996)</xref> showed that Quaternary climatic oscillations did not lead to enhanced rates of extinction and speciation in carabids, as inferred from 14C-dated fos<pmc-comment>PageBreak</pmc-comment>sil assemblages. The future responses of <named-content content-type="taxon-name">Carabidae</named-content> to climate change will probably be similar to that of the past, with the exception that extinction rates are expected to be higher because of human-caused habitat fragmentation.</p><p>Rasplus et al. (2000) found that populations of the threatened species <italic><named-content content-type="taxon-name">Carabus solieri</named-content></italic> consist of two distinct clusters corresponding to subspecies <italic>bonnetianus</italic> and <italic>solieri</italic>. These populations were probably isolated during the last glaciation and are worthy of protection as gene flow is restricted between these two groups. Moreover, molecular markers suggest that the subspecies <italic>curtii</italic> is a hybrid between <italic>bonnetianus</italic> and <italic>solieri</italic>.</p><p><xref ref-type="bibr" rid="B139">Desender et al. (2000)</xref> investigated the genetic diversity and wing polymorphism of the salt-marsh beetle <italic><named-content content-type="taxon-name">Pogonus chalceus</named-content></italic> in 30 populations from the Atlantic coast and nine populations from the Mediterranean Basin. These Mediterranean populations showed little differentiation associated with high dispersal power, a finding possibly related to habitat instability. A higher structuring was found in Atlantic populations, which showed varying degrees of wing polymorphism and dispersal power, possibly related to adaptation to particular conditions.</p><p><xref ref-type="bibr" rid="B271">Kamer et al. (2008)</xref> investigated variation in the 12S RNA sequence in populations at different geographic scales, namely the Baltic coast, inland populations across Central Europe, and Central plus Western Europe. Population structure varied as a result of complex factors that include past history and present dispersal power, amongst others. Cryptic taxa or a lack of molecular differences among siblings were also found, showing the usefulness of landscape genetic analyses.</p></sec><sec sec-type="2.4. Species borders and hybridisation"><title>2.4. Species borders and hybridisation</title><p><xref ref-type="bibr" rid="B283">Koch (1986)</xref> showed that <italic><named-content content-type="taxon-name">Pterostichus nigrita</named-content></italic> and its sibling <italic><named-content content-type="taxon-name">Pterostichus rhaeticus</named-content></italic> are distinct species according to habitat preferences, subtle details in male and female genitalia and karyotypic numbers. Both species are reproductively isolated, as shown by crossbreeding laboratory experiments. More recently, <xref ref-type="bibr" rid="B10">Angus et al. (2008)</xref> described a new cryptic species in the Iberian Peninsula, <italic><named-content content-type="taxon-name">Pterostichus carri</named-content></italic>, and a new subspecies of <italic><named-content content-type="taxon-name">Pterostichus nigrita</named-content></italic> from Anatolia. All taxa shared a basic 2n = 36 + X male karyotype, whereas marked variation in the number of accessory chromosomes was found within and between these taxa.</p><p><xref ref-type="bibr" rid="B588">Vogler and DeSalle (1994)</xref> analysed the relationships of 17 populations of <italic><named-content content-type="taxon-name">Cicindela dorsalis</named-content></italic> along a littoral transect from New England to Veracruz. These populations are currently ascribed to four subspecies which is difficult to ascertain. Mitochondrial DNA haplotypes showed that populations could readily be grouped into two major entities that represent well defined phylogenetic species without gene flow between them, one occupying the Atlantic coast, the other inhabiting the Gulf of Mexico. Within each of these entities, moderate diversification was found but without much geographic structure, probably because of moderate gene flow between populations.</p><p><xref ref-type="bibr" rid="B207">Galián et al. (1996)</xref> studied the karyotypes and the RFLPs resulting from digestion of total DNA with endonuclease EcoRI in four populations ascribed to <italic><named-content content-type="taxon-name">Ceroglossus chilensis</named-content></italic>. Differences between these populations in terms of chromosome number<pmc-comment>PageBreak</pmc-comment> and molecular data led to the conclusion that there are three cryptic species living in sympatry.</p><p>A clear distinction between <italic><named-content content-type="taxon-name">Abax parallelepipedus</named-content></italic> and <italic><named-content content-type="taxon-name">Abax angustatus</named-content></italic> (reported as a subspecies of the former) resulted from a morphological analysis of sympatric populations of both species, and a molecular study based on allozymes and mitochondrial DNA (<xref ref-type="bibr" rid="B160">Düring 2002</xref>). No molecular evidence of hybridisation between these two species was found.</p><p><xref ref-type="bibr" rid="B387">Mossakowski et al. (1986)</xref> carried out a field study on the frequency of hybrids between species of the subgenus <italic><named-content content-type="taxon-name">Chrysocarabus</named-content></italic>, <italic><named-content content-type="taxon-name">Chrysocarabus lineatus</named-content></italic> and <italic><named-content content-type="taxon-name">Chrysocarabus splendens</named-content></italic> in the Pyrenees. Reliable morphological characters allowed for determining the occurrence of hybrids. Both species may hybridise (up to 40% of individuals) when particular ecological conditions are met, which indicates that complete reproductive isolation has not yet been attained. However, a number of characters are fixed in each species allowing their classification as valid species. Furthermore, <xref ref-type="bibr" rid="B164">Düring et al. (2000</xref>, <xref ref-type="bibr" rid="B163">2006</xref>) studied the mitochondrial haplotype in many <italic><named-content content-type="taxon-name">Chrysocarabus splendens</named-content></italic> populations and found convincing evidence of introgressive hybridisation in <italic><named-content content-type="taxon-name">Chrysocarabus</named-content></italic> (incongruence between mitochondrial and nuclear gene trees). In contrast, nuclear ITS-2 sequences showed that populations of <italic><named-content content-type="taxon-name">Chrysocarabus splendens</named-content></italic> made up a monophyletic clade, which is sister to that made up by <italic><named-content content-type="taxon-name">Chrysocarabus lineatus</named-content></italic> and <italic><named-content content-type="taxon-name">Chrysocarabus lateralis</named-content></italic>. Shared haplotypes between <italic><named-content content-type="taxon-name">Chrysocarabus splendens</named-content></italic> and <italic><named-content content-type="taxon-name">Chrysocarabus punctatoauratus</named-content></italic> are probably the result of introgression of the latter into the former species. On the other hand, mitochondrial DNA of <italic><named-content content-type="taxon-name">Chrysocarabus rutilans</named-content></italic> was probably acquired from <italic><named-content content-type="taxon-name">Chrysocarabus splendens</named-content></italic> through introgression.</p></sec><sec sec-type="2.5. Speciation, radiation and biogeography"><title>2.5. Speciation, radiation and biogeography</title><p><xref ref-type="bibr" rid="B267">Juberthie (1979)</xref> analysed the evolutionary pathways of the genus <italic><named-content content-type="taxon-name">Aphaenops</named-content></italic> (<named-content content-type="taxon-name">Trechinae</named-content>) from putative epigean ancestors to specialised troglobionts, and noted that food must have been a major factor in promoting their morpho-functional characters. He also concluded that <italic><named-content content-type="taxon-name">Aphaenops</named-content></italic> and other hypogean <named-content content-type="taxon-name">Trechinae</named-content> are not living fossils but show highly derived characters, either regressive (loss of eyes and pigmentation) or positive (slender appendages, new chemoreceptors) with regard to ancestral epigean forms, with which they still share particular plesiomorphies.</p><p><xref ref-type="bibr" rid="B384">Mossakowski (1979)</xref> postulated that habitat preference is an evolutionary process that can be reconstructed when matching it against a phylogenetic tree of particular taxa. He tested this hypothesis by considering the subgenus <italic><named-content content-type="taxon-name">Chrysocarabus</named-content></italic> and concluded that there was an adaptive shift from Mediterranean to deciduous forests and a recent colonisation of alpine environments.</p><p><xref ref-type="bibr" rid="B317">Liebherr (1986)</xref> constructed a phylogeny of the <italic><named-content content-type="taxon-name">Agonum extensicolle</named-content></italic> group based on morphological quantitative characters and the allelic frequencies derived from the electrophoresis of soluble enzymes. The resulting tree was used to test the hypothesis of the vicariance effects of the Cochise filter/barrier separating the Sonoran and Chihuahuan deserts in SW North America. He argued that the zone between the deserts has <pmc-comment>PageBreak</pmc-comment>probably caused vicariant events between particular pairs of species and species groups, and also between subspecies of <italic><named-content content-type="taxon-name">Agonum decorum</named-content></italic> 6.5 to 2.8 million years ago. This barrier has probably led to the same phenomena in other carabid taxa.</p><p><xref ref-type="bibr" rid="B132">Desender et al. (1990)</xref> studied speciation of the genus <italic><named-content content-type="taxon-name">Pterostichus</named-content></italic> in the Galápagos using multivariate morphometric analysis and ecological data. They concluded that a combination of allopatric (stepping stone model) and parapatric events (segregation in altitude of two species inhabiting the same island) may explain radiation of the genus from ancestors related to <italic><named-content content-type="taxon-name">Pterostichus peruviana</named-content></italic>, a species presently found in South America.</p><p><xref ref-type="bibr" rid="B5">Andersen and Skorping (1990)</xref> presented a conclusive model of sympatric speciation of the genus <italic><named-content content-type="taxon-name">Bembidion</named-content></italic> (and in particular in the subgenus <italic><named-content content-type="taxon-name">Chysobracteon</named-content></italic>), in which habitat selection and the effects of parasites may give rise to disruptive selection that promotes reproductive isolation and in turn speciation. Habitat shifts in riparian carabids may have evolved in sympatry, whereas allopatry would have produced new taxa showing mere variations of the same ecological theme.</p><p><xref ref-type="bibr" rid="B33">Baehr (1994)</xref> constructed a cladistic analysis of the <named-content content-type="taxon-name">Pseudomorphinae</named-content> based on morphological characters that solved relationships of the main lineages within the subfamily. He postulated that the subfamily has an Australian-South American origin, and that it has recently spread to North America and SE Asia.</p><p><xref ref-type="bibr" rid="B71">Brandmayr and Zetto Brandmayr (1994)</xref> presented an elaborated hypothesis on the evolutionary history of the genus <italic><named-content content-type="taxon-name">Abax</named-content></italic>, based on characters of male genitalia (inflated median lobe), larval morphology, type of parental care and larval behaviour, habitat preferences and geographic distribution. Ancestors of this genus possibly inhabited lowland forests during the late Miocene, whereas most recent taxa are found in alpine grasslands and mountain forests. This suggests that there has been a major colonisation trend towards mountains during the last geological periods. A predominantly allopatric pattern was inferred for the radiation of <italic><named-content content-type="taxon-name">Abax</named-content></italic>.</p><p>The supertribe Carabitae poses major evolutionary problems because many character states are difficult to interpret due to homoplasy, and the biogeographic patterns of tribes are not congruent at first glance with relationships derived from molecular and morphological data. A synthesis of different studies (<xref ref-type="bibr" rid="B469">Prüser and Mossakowski 1998</xref>; <xref ref-type="bibr" rid="B270">Kamer et al. 2002</xref>; <xref ref-type="bibr" rid="B385">Mossakowski 2002</xref>) based on the analysis of morphological characters (adults and larvae) plus molecular data, indicates that <named-content content-type="taxon-name">Cychrini</named-content> is sister to all other tribes, and that <named-content content-type="taxon-name">Carabini</named-content> is sister to a clade made up of tribes <named-content content-type="taxon-name">Ceroglossini</named-content> and <named-content content-type="taxon-name">Pamborini</named-content>. This hypothesis also postulates a Laurasian origin of Carabitae and a single migration event across the tropics. A corollary of this hypothesis is that the <italic><named-content content-type="taxon-name">Cychrus</named-content></italic>-like mandible of <italic><named-content content-type="taxon-name">Pamborus</named-content></italic> is a homoplasy that would result from an adaptation to feed on snails (‘cychrisation<italic>’</italic>).</p><p>Of the four <italic><named-content content-type="taxon-name">Calosoma</named-content></italic> species inhabiting the Galápagos, only <italic><named-content content-type="taxon-name">Calosoma granatense</named-content></italic> is widespread among islands and altitudinal habitats. In spite of its high dispersal power and morphological stability, this species shows substantial genetic differentiation between populations on different islands and volcanoes (<xref ref-type="bibr" rid="B142">Desender and Verdyck 2000</xref>). There was probably a single colonisation event from the mainland and a stepping-stone<pmc-comment>PageBreak</pmc-comment> model of island colonisation. However, gene flow must have been enough to prevent speciation events. The other three <italic><named-content content-type="taxon-name">Calosoma</named-content></italic> species of the Galápagos are endemic to localities at high altitudes on a single island, which suggests that they have originated by convergent habitat shifts.</p><p>The phylogenetic relationship of three <italic><named-content content-type="taxon-name">Carabus</named-content></italic> species inhabiting the Tenerife and Gran Canaria (subgenus <italic><named-content content-type="taxon-name">Nesaeocarabus</named-content></italic>) was investigated by a phylogenetic analysis based of the mitochondrial <italic>nd5</italic> gene (<xref ref-type="bibr" rid="B468">Prüser et al. 2000</xref>). The hypothesis of a close relationship between <italic><named-content content-type="taxon-name">Nesaeocarabus</named-content></italic> and the subgenus <italic><named-content content-type="taxon-name">Eucarabus</named-content></italic> was rejected. Instead, Canarian taxa were closely related to the subgenus <italic><named-content content-type="taxon-name">Eurycarabus</named-content></italic> from northern Africa, southern Italy, Sardinia and Sicily. Diversification of <italic><named-content content-type="taxon-name">Nesaeocarabus</named-content></italic> in the Canaries was congruent with the geological history of the archipelago, with a diversification of ancestors beginning 14–7 million years ago.</p><p>The subgenus <italic><named-content content-type="taxon-name">Platycarabus</named-content></italic> includes five species living in the Alps and adjacent areas. <xref ref-type="bibr" rid="B93">Casale et al. (1998)</xref> tested the hypothesis of a close relationship of these species with the subgenus <italic><named-content content-type="taxon-name">Hygrocarabus</named-content></italic>, both included in the genus <italic><named-content content-type="taxon-name">Chaetocarabus</named-content></italic> sensu <xref ref-type="bibr" rid="B258">Ishikawa (1984)</xref>. Separate and combined analyses of 26 adult and larval characters, and of sequences of the <italic>nd1</italic> gene, rejected this hypothesis, as <italic><named-content content-type="taxon-name">Platycarabus</named-content></italic> is a robust monophyletic lineage distantly related to <italic><named-content content-type="taxon-name">Chaetocarabus</named-content></italic>, and is even farther from <italic><named-content content-type="taxon-name">Hygrocarabus</named-content></italic>.</p><p><xref ref-type="bibr" rid="B386">Mossakowski (2005)</xref> revised the proposal of <xref ref-type="bibr" rid="B257">Imura (2002)</xref> of grouping the genus <italic><named-content content-type="taxon-name">Carabus</named-content></italic> <italic>s. l.</italic> into 29 sections and 137 genera, based on molecular data (see also <xref ref-type="bibr" rid="B92">Casale and Mossakowski 2003</xref>). Analysis of the inflated median lobe of the male endophallus and the reassessment of DNA sets with stringent criteria of bootstrap values showed that (i) relationships of the subgenera of <italic><named-content content-type="taxon-name">Carabus</named-content></italic> were poorly solved, (ii) the results do not support the hypothesis of an explosive radiation of the ancestors of this genus, and (iii) these uncertainties do not favour the ranking of subgenera to genera proposed by <xref ref-type="bibr" rid="B257">Imura (2002)</xref>.</p></sec><sec sec-type="2.6. Phylogeny based on different types of characters"><title>2.6. Phylogeny based on different types of characters</title><p><bold>Ethology</bold></p><p><xref ref-type="bibr" rid="B67">Brandmayr and Zetto Brandmayr (1979)</xref> found that the genus <italic><named-content content-type="taxon-name">Abax</named-content></italic> shows different stages between a pure pre-social condition of merely laying eggs with a well-developed ovipositor, and the advanced construction of a chamber, laying the eggs in capsules and taking care of brood until hatching and pigmentation of the larvae. It was concluded that behavioural characters are difficult to interpret in a phylogenetic context due to convergence. However, in some instances they provide valuable clues to reconstruct the evolution of a group and give a good phylogenetic signal.</p><p><pmc-comment>PageBreak</pmc-comment>
<bold>Morphology</bold></p><p>Wing folding mechanisms have been suggested to be a character with phylogenetic value at higher taxonomic ranks (<xref ref-type="bibr" rid="B225">Hammond 1979</xref>). Differences in the structure (presence of patches of microtrichia) and mechanism (abdominal movements helping with folding) of wing folding among lineages of <named-content content-type="taxon-name">Carabidae</named-content> are not congruent with phylogenetic inferences derived from other characters. The <named-content content-type="taxon-name">Trachypachidae</named-content> is a lineage distinct from carabids, a conclusion congruent with recent molecular (<xref ref-type="bibr" rid="B349">Maddison et al. 2009</xref>) and karyotypic data (<xref ref-type="bibr" rid="B362">Martínez-Navarro et al. 2011</xref>), whereas <italic><named-content content-type="taxon-name">Gehringia</named-content></italic> was close to other carabids, as currently accepted. The basis for investigating the phylogenetic value of wing venation within <named-content content-type="taxon-name">Adephaga</named-content> and <named-content content-type="taxon-name">Carabidae</named-content> was outlined by <xref ref-type="bibr" rid="B592">Ward (1979)</xref>. This topic has received little attention, perhaps because there is a generalised model in <named-content content-type="taxon-name">Carabidae</named-content> that shows a relatively low degree of variation within particular lineages at the tribal or generic level.</p><p>Higher-ranked taxa were considered by <xref ref-type="bibr" rid="B50">Beutel (1998)</xref> when analysing the relationships of <named-content content-type="taxon-name">Trachypachidae</named-content>, based on morphological and functional characters of adults and larvae. He concluded that the family <named-content content-type="taxon-name">Gyrinidae</named-content> is sister to all other Adephagan groups. Of these clades, <named-content content-type="taxon-name">Haliplidae</named-content> was sister to the remaining families; these were in turn split into two main clades, one made up of <named-content content-type="taxon-name">Carabidae</named-content> (including <named-content content-type="taxon-name">Rhysodini</named-content> and Cicindelitae), the other made up of (<named-content content-type="taxon-name">Trachypachidae</named-content>) + (<named-content content-type="taxon-name">Noteridae</named-content>(<named-content content-type="taxon-name">Amphizoidae</named-content>+<named-content content-type="taxon-name">Dytiscidae</named-content>)). These results contradict <xref ref-type="bibr" rid="B51">Beutel and Haas (1996)</xref>, who found <named-content content-type="taxon-name">Trachypachinae</named-content> to be sister to <named-content content-type="taxon-name">Carabidae</named-content>; Beutel and Haas’ hypothesis has recently received support from molecular analyses (<xref ref-type="bibr" rid="B349">Maddison et al. 2009</xref>). Ancestors of Adephagan beetles were probably associated with riparian habitats and it has been postulated that independent colonisations of aquatic habitats gave rise to the families <named-content content-type="taxon-name">Gyrinidae</named-content>, <named-content content-type="taxon-name">Haliplidae</named-content> and <named-content content-type="taxon-name">Dytiscidae</named-content>.</p><p><xref ref-type="bibr" rid="B318">Liebherr and Will (1998)</xref> studied the phylogenetic value of characters of the female reproductive tract at an inclusive scale that covered the whole family <named-content content-type="taxon-name">Carabidae</named-content>. Surprisingly no character defined the <named-content content-type="taxon-name">Carabidae</named-content> as a monophyletic taxon; instead the Isochaeta appeared as the adelphotaxon of Anisochaeta (that included <named-content content-type="taxon-name">Gehringiini</named-content> and <named-content content-type="taxon-name">Rhysodini</named-content>). In turn, the Anisochaeta was divided into two clades separated by the evolution of a secondary spermatheca. Less inclusive clades within these two major groups of Anisochaeta showed relationships that agreed with previous hypotheses in some cases.</p><p><xref ref-type="bibr" rid="B15">Arndt (1998)</xref> analysed the phylogenetic relationships derived from larval morphology in 44 tribes of <named-content content-type="taxon-name">Carabidae</named-content>. He found support for a monophyletic <named-content content-type="taxon-name">Carabidae</named-content>+<named-content content-type="taxon-name">Tachypachidae</named-content>+<named-content content-type="taxon-name">Dytiscidae</named-content> clade. The family <named-content content-type="taxon-name">Carabidae</named-content> was also a monophyletic clade if <named-content content-type="taxon-name">Rhysodidae</named-content> were excluded. The Cicindelitae was also monophyletic and showed several autapomorphies. Metriitae and Paussitae made up a monophyletic clade. The subfamily <named-content content-type="taxon-name">Harpalinae</named-content> (“higher” carabids) appeared to be a monophyletic clade but relationships of Brachinitae were ambiguous and remain a major challenge for future studies; a close relationship with <named-content content-type="taxon-name">Harpalinae</named-content> is unlikely.<pmc-comment>PageBreak</pmc-comment></p><p>The phylogenetic relationships among basal grade <named-content content-type="taxon-name">Carabidae</named-content> was revisited by <xref ref-type="bibr" rid="B275">Kavanaugh (1998)</xref> who showed that <named-content content-type="taxon-name">Trachypachidae</named-content> is sister to all carabid taxa examined (which confirms similar conclusions reported in former works), that the supertribe Nebriitae is a grade rather than a clade (<named-content content-type="taxon-name">Nebriini</named-content> is separated from related tribes), and that cicindines are related to <named-content content-type="taxon-name">Carabini</named-content>, <named-content content-type="taxon-name">Cychrini</named-content>, <named-content content-type="taxon-name">Cicindelini</named-content> and <named-content content-type="taxon-name">Omophronini</named-content>.</p><p>Cladistic analyses based on different data sets (morphology, ethology, geographic distribution), were carried out to investigate the phylogeny of <named-content content-type="taxon-name">Paussinae</named-content> (<xref ref-type="bibr" rid="B395">Nagel 1979b</xref>), <named-content content-type="taxon-name">Ozaenini</named-content> plus <named-content content-type="taxon-name">Metriini</named-content> and <named-content content-type="taxon-name">Paussini</named-content> (<xref ref-type="bibr" rid="B584">Vigna Taglianti et al. 1998</xref>), the <italic><named-content content-type="taxon-name">Agra cayennensis</named-content></italic> group (<xref ref-type="bibr" rid="B176">Erwin 1996</xref>), the supertribe Nebriitae (<xref ref-type="bibr" rid="B274">Kavanaugh 1996</xref>), the subtribe <named-content content-type="taxon-name">Calleidina</named-content> (<named-content content-type="taxon-name">Lebiini</named-content>; <xref ref-type="bibr" rid="B89">Casale 1998</xref>), the Western Hemisphere <named-content content-type="taxon-name">Pseudomorphini</named-content> (<xref ref-type="bibr" rid="B179">Erwin and Geraci 2008</xref>), the tribe <named-content content-type="taxon-name">Rhysodini</named-content> (<xref ref-type="bibr" rid="B48">Bell 1998</xref>; which is likely a highly specialised predator of slime moulds rather than a primitive Adephagan stock), the subfamily <named-content content-type="taxon-name">Broscinae</named-content> (<xref ref-type="bibr" rid="B482">Roig-Juñent 1998</xref>), and the subfamily <named-content content-type="taxon-name">Psydrinae</named-content> (Baehr, 1998). These studies either corroborated previous ideas about relationships of taxa or shed light on new and unsuspected hypotheses about the phylogeny and classification of taxa, including the erection of new high-ranked taxa.</p><p><bold>Defence substances</bold></p><p>Characterisation of chemical compounds used for defence and the phylogenetic interest of this trait was summarised by <xref ref-type="bibr" rid="B382">Moore (1979)</xref>. The review showed that (i) compounds can be grouped into at least nine categories according to their chemical nature, (ii) there probably occurred a convergent development of the same substances in distantly related lineages, (iii) diversification of chemical types occurred within some subfamilies (e.g. <named-content content-type="taxon-name">Pterostichinae</named-content>) whereas others (<named-content content-type="taxon-name">Harpalinae</named-content>, <named-content content-type="taxon-name">Lebiinae</named-content>) are much more uniform; (iv) the phylogenetic signal of this trait is valuable at tribal level or higher ranks; some compounds seem to vary in particular lineages (Australian <named-content content-type="taxon-name">Panagaeninae</named-content>) and could be useful for assessing relationships at lower ranks; and (v) further insight into this trait would result from the study of biochemical synthetic pathways, fine structure of defensive glands and the detection of more subtle compounds.</p><p><bold>Karyotypic evolution</bold></p><p>A number of contributions have addressed the question on the ancestral karyotype of Adephaga and the <named-content content-type="taxon-name">Carabidae</named-content>, and its main patterns of evolutionary change (<xref ref-type="bibr" rid="B400">Nettmann 1986</xref>; <xref ref-type="bibr" rid="B513">Serrano 1986</xref>; <xref ref-type="bibr" rid="B516">Serrano and Galián 1998</xref>), or referred to the karyotypic evolution of particular taxa (<named-content content-type="taxon-name">Harpalini</named-content>: <xref ref-type="bibr" rid="B517">Serrano et al. 1994</xref>). The family <named-content content-type="taxon-name">Carabidae</named-content> (915 taxa analysed) is characterised by a notable variation of the diploid number (2n = 4 - 69), the occurrence of high chromosome numbers in comparison to Polyphagan beetles, and a <pmc-comment>PageBreak</pmc-comment>repeated karyotypic formula in well-studied lineages (e.g. 2n = 26 + XY in <named-content content-type="taxon-name">Carabini</named-content>; 2n = 22 + XY in <named-content content-type="taxon-name">Bembidiini</named-content>; 2n = 36 + X in <named-content content-type="taxon-name">Harpalini</named-content>).</p><p>The ancestral karyotype of <named-content content-type="taxon-name">Coleoptera</named-content>, still present in many Polyphagan lineages, 2n = 18 + Xyp, had probably undergone significant changes in the ancestors of carabids, since neither this number of autosomes nor the particular Xyp sex chromosomes are found in any carabid. The ancestral condition of a 2n = 36 + X0 male karyotype is widespread in many lineages and may be notably diversified in particular carabid lineages. The occurrence of this formula in some dytiscids and in trachypachids (<xref ref-type="bibr" rid="B362">Martínez-Navarro et al. 2011</xref>) provided further support to this hypothesis. However, it has not been found in lineages showing plesiomorphic morphological characters, which suggests that it has evolved rapidly in earlier offshoots of the <named-content content-type="taxon-name">Carabidae</named-content>.</p><p>Karyotypic data have been shown to be valuable for understanding carabid systematics though it seems that karyotypic changes are not a main driving force for speciation in carabids. This is not to deny the role of karyotypic changes in reinforcing isolation mechanisms in recently originated taxa, regardless of the occurrence of speciation processes under conditions of geographic isolation or in lowland areas (<xref ref-type="bibr" rid="B514">Serrano 1992</xref>).</p><p><xref ref-type="bibr" rid="B517">Serrano et al. (1994)</xref> summarised the karyotypic data of members of the tribe <named-content content-type="taxon-name">Harpalini</named-content>, and found that ancestors likely had a 2n = 36 + X male karyotype. Constraints to numerical variations within this tribe are similar to those found among other carabid tribes. The <named-content content-type="taxon-name">Ditomina</named-content> are peculiar because they show high chromosome numbers, which corroborates its ranking as a separate subtribe.</p><p><bold>Molecular data</bold></p><p>The number of molecular studies have increased since the 1990s, either based only on molecular data or (more recently) combined with other data sets. Inferred relationships have corroborated relationships derived from traditional taxonomy but also often contradicted these, thus emphasising the need of more holistic approaches aimed at obtaining robust and congruent phylogenies.</p><p><xref ref-type="bibr" rid="B348">Maddison et al. (1998)</xref> published the first comprehensive DNA-based phylogeny of <named-content content-type="taxon-name">Carabidae</named-content>. They studied the nuclear small subunit (18S) ribosomal DNA, sequenced in 35 carabid genera representing 26 tribes. All higher-level clades were monophyletic except for the Scrobifera (scaritines plus clivinines); the Trechitae was sister to Patrobines; <italic><named-content content-type="taxon-name">Morion</named-content></italic> and Pseudomorpha were members of <named-content content-type="taxon-name">Harpalinae</named-content>; <italic><named-content content-type="taxon-name">Psydrus</named-content></italic> and elaphrines were sisters and both were sister to trechites plus patrobines; there was a grade including scaritines immediately below <named-content content-type="taxon-name">Harpalinae</named-content>.</p><p>A combined analysis of larval morphological characters and molecular data of Cicindelitae showed a number of inferences that contradict current systematics: <named-content content-type="taxon-name">Omina</named-content> had a basal position, <named-content content-type="taxon-name">Megacephalini</named-content> was a polyphyletic taxon, and <named-content content-type="taxon-name">Cicindelinae</named-content> was not monophyletic (<xref ref-type="bibr" rid="B587">Vogler and Barraclough 1998</xref>). Use of the resulting inferences showed that there are differential diversification rates among major lineages (e.g. a<pmc-comment>PageBreak</pmc-comment> high rate of diversification was found at the base of megacephalines and collyrines, and another at the base of cicindelines).</p><p><xref ref-type="bibr" rid="B162">Düring and Brückner (2000)</xref> investigated the phylogeny and history of lineages of <named-content content-type="taxon-name">Molopina</named-content> using molecular analysis based on the sequence of two mitochondrial DNA fragments. Representatives of the genera <italic><named-content content-type="taxon-name">Percus</named-content></italic>, <italic><named-content content-type="taxon-name">Molops</named-content></italic> and <italic><named-content content-type="taxon-name">Abax</named-content></italic> were included, as well as <italic><named-content content-type="taxon-name">Pterostichus</named-content></italic> and <italic><named-content content-type="taxon-name">Carabus</named-content></italic> as outgroups. These three genera made up a monophyletic clade, and <italic><named-content content-type="taxon-name">Molops</named-content></italic> and <italic><named-content content-type="taxon-name">Abax</named-content></italic> were sister taxa. In a further step, <xref ref-type="bibr" rid="B77">Brückner and Mossakowski (2006)</xref> investigated the phylogeny of the genus <italic><named-content content-type="taxon-name">Percus</named-content></italic> by integrating previous molecular, morphological and biogeographic characters with those of nuclear 28S rRNA. This genus is likely a monophyletic taxon divided into three main clades. Relationships among the Tyrrhenian taxa remained unresolved probably as a result of recent diversification and low mutation rates of the molecular marker.</p><p>A molecular study of the tribe <named-content content-type="taxon-name">Harpalini</named-content> based on the mitochondrial <italic>cox1</italic> gene (<xref ref-type="bibr" rid="B361">Martínez-Navarro et al. 2005</xref>) showed that (i) <named-content content-type="taxon-name">Pelmatellina</named-content> should be included within <named-content content-type="taxon-name">Stenolophina</named-content>, (ii) subtribe <named-content content-type="taxon-name">Harpalina</named-content> is polyphyletic, (iii) <named-content content-type="taxon-name">Ditomina</named-content> is a valid subtribe, and (iv) Selenophori should be ranked as a valid subtribe closely related to the <named-content content-type="taxon-name">Anisodactylina</named-content>.</p><p>An analysis based on sequences of 28S and <italic>wingless</italic> genes of <italic><named-content content-type="taxon-name">Ildobates neboti</named-content></italic> (a rare hypogean species inhabiting a few caves in eastern Spain) and related taxa showed that tribes currently included in Dryptitae (<named-content content-type="taxon-name">Dryptini</named-content>, <named-content content-type="taxon-name">Galeritini</named-content> and <named-content content-type="taxon-name">Zuphiini</named-content>) made up a monophyletic clade, and that <italic><named-content content-type="taxon-name">Ildobates neboti</named-content></italic> is a member of the <named-content content-type="taxon-name">Zuphiini</named-content> (<xref ref-type="bibr" rid="B478">Ribera et al. 2006</xref>).</p><p><xref ref-type="bibr" rid="B589">Vogt et al. (2005)</xref> studied the relationships of African <italic><named-content content-type="taxon-name">Anthia</named-content></italic> and <italic><named-content content-type="taxon-name">Termophilum</named-content></italic>, and the related <italic><named-content content-type="taxon-name">Cypholoba chaudoiri</named-content></italic>, based on the sequence of the mitochondrial <italic>nd5</italic> gene. Taxa of <italic><named-content content-type="taxon-name">Anthia</named-content></italic> made up a monophyletic clade in which <italic><named-content content-type="taxon-name">Cypholoba chaudoiri</named-content></italic> was unexpectedly included. Taxa of <italic><named-content content-type="taxon-name">Termophilum</named-content></italic> made up two distinct clusters, which suggests paraphyly of this genus.</p><p>Current division of the genus <italic><named-content content-type="taxon-name">Calathus</named-content></italic> (<named-content content-type="taxon-name">Sphodrini</named-content>) was investigated on molecular grounds by sampling a <italic>cox1-cox2</italic> fragment in 44 taxa (<xref ref-type="bibr" rid="B483">Ruiz and Serrano 2006</xref>). The monophyly of the subgenus <italic><named-content content-type="taxon-name">Calathus</named-content></italic> was corroborated, as well as the distinctness of the monotypic subgenera <italic><named-content content-type="taxon-name">Bedelinus</named-content></italic> and <italic><named-content content-type="taxon-name">Iberocalathus</named-content></italic>. The subgenus <italic><named-content content-type="taxon-name">Neocalathus</named-content></italic> is polyphyletic and needs taxonomic revision and the same holds true for the Canarian <italic><named-content content-type="taxon-name">Lauricalathus</named-content></italic>. The latter subgenus should be divided into two subgenera, and one of these should include <italic><named-content content-type="taxon-name">Trichocalathus.</named-content></italic></p></sec></sec><sec><title>3 Biology</title><sec sec-type="3.1. Life history strategies and rhythms"><title>3.1. Life history strategies and rhythms</title><p>Land animals evolve strategies to optimise and synchronise their life cycle with seasonal changes of the environment. For example, reproduction usually takes place under optimal conditions, while metabolism may be reduced if conditions are suboptimal (e.g. dormancy, which in carabids has thus far only been observed for larval and adult stages).</p><p><pmc-comment>PageBreak</pmc-comment>
<bold>Ultimate (limiting) factors regulating ground beetle life histories</bold></p><p>Ultimate factors determining beetle life cycles include variation in temperature and rainfall. Optimal development of the immature stages requires an estimated temperature range of 4–35°C. Rainfall, in combination with temperature, affects soil humidity, which is critical because eggs absorb water from their surroundings to complete embryonic development (<xref ref-type="bibr" rid="B441">Paarmann 1986</xref>) and larvae are sensitive to desiccation (<xref ref-type="bibr" rid="B432">Paarmann 1973</xref>).</p><p>Food can also be critical. Reproduction of, for example, seed-feeding carabid species may be governed by ripe seeds that usually appear at the end of the wet or warm season. Only very few habitats offer suitable conditions for polyvoltine development throughout the year, for example, lake shores, swamps and some lowland rainforests with very short dry spells.</p><p>The only ultimate factor determining carabid beetle life cycles in the Arctic, Subarctic and Antarctic, as well as in montane habitats of the temperate zone is temperature (e.g. <xref ref-type="bibr" rid="B551">Thiele 1977</xref>). In the summer, only a short time window exists for reproduction and development. All species in these habitats are summer developers. Species with rapid larval development, such as <italic><named-content content-type="taxon-name">Pterostichus adstrictus</named-content></italic> (<xref ref-type="bibr" rid="B443">Paarmann 1994</xref>), are true summer breeders with adult hibernation only. Species with slow larval development hibernate as larvae as well as adults and require more than one season to complete their life cycle (<xref ref-type="bibr" rid="B273">Kaufmann 1971</xref>; <xref ref-type="bibr" rid="B108">Davies 1972</xref>; <xref ref-type="bibr" rid="B371">Matalin 2008</xref>). In the mountains of temperate Europe (altitude of 2200–2600 m) the favourable season is reduced to 3–4 months. In forest Pterostichines, especially in the large genus <italic><named-content content-type="taxon-name">Pterostichus</named-content></italic> and in the Molopines <italic><named-content content-type="taxon-name">Abax</named-content></italic> and <italic><named-content content-type="taxon-name">Percus</named-content></italic>, cycles are often biennial (<xref ref-type="bibr" rid="B63">Brandmayr 1977</xref>). In the genus <italic><named-content content-type="taxon-name">Molops</named-content></italic>, where embryonic development can last for more than one month and the eggs are guarded in a subterranean hole, the females disappear from the soil surface during summer, and reappear in the autumn. The subterranean larvae are active during winter, and the new generation requires a further year to reach maturity (<xref ref-type="bibr" rid="B70">Brandmayr and Zetto Brandmayr 1991</xref>).</p><p><xref ref-type="bibr" rid="B310">Larsson (1939)</xref> recognised different reproductive strategies in carabid beetles of the temperate zone by studying museum material. He divided them into Frühlingstiere (spring breeders) and Herbsttiere (autumn breeders). Spring breeders reproduce during the spring and hibernate as adults only. Autumn breeders reproduce during the autumn and hibernate mainly as larvae. In a number of species, adults may hibernate after reproduction to enter a second reproductive period (<xref ref-type="bibr" rid="B213">Gilbert 1956</xref>; <xref ref-type="bibr" rid="B586">Vlijm et al. 1968</xref>; <xref ref-type="bibr" rid="B506">Schjøtz-Christensen 1968</xref>; <xref ref-type="bibr" rid="B299">Krehan 1970</xref>). (<xref ref-type="bibr" rid="B504">Schjøtz-Christensen (1965</xref>, 1966) showed that in some <italic><named-content content-type="taxon-name">Harpalus</named-content></italic> species spring and autumn breeding populations co-occure in the same habitat. Other examples include <italic><named-content content-type="taxon-name">Abax parallelepipedus</named-content></italic> (<xref ref-type="bibr" rid="B331">Löser 1970</xref>), <italic><named-content content-type="taxon-name">Poecilus lepidus</named-content></italic> (<xref ref-type="bibr" rid="B442">Paarmann 1990</xref>), <italic><named-content content-type="taxon-name">Pseudophonus rufipes</named-content></italic> (<xref ref-type="bibr" rid="B366">Matalin 1997a</xref>) and <italic><named-content content-type="taxon-name">Harpalus affinis</named-content></italic> (<xref ref-type="bibr" rid="B369">Matalin 1998</xref>). A third breeding category – spring-autumn breeder (<xref ref-type="bibr" rid="B367">Matalin 1997b</xref>) – is found in the genus <italic><named-content content-type="taxon-name">Stenolophus</named-content></italic>. In 1990 Den Boer and Den Boer-Daanje, summarising the life history strategies of 68 common carabid beetles in Drenthe (the Netherlands), found a continuum of reproduction from early spring to late autumn,<pmc-comment>PageBreak</pmc-comment> and seven of them reproduced during winter. Den Boer and Den Boer-Daanje distinguished species with summer larvae (summer developers, 40 species) and species with winter larvae (winter developers, 28 species). Drenthe is located in an area with Atlantic climate: warm winters and wet summers, thus offering a broad reproductive window. In areas with a continental climate, however, this window is much narrower.</p><p>Cave environments are buffered against climatic variation and can have (i) a constant temperature throughout the year, or (ii) distinct seasonality. Trechines living in caves are mostly autumnal reproducers with winter larvae. The rhythms of <italic><named-content content-type="taxon-name">Aphaenops</named-content></italic> and related genera may show distinct seasonality at least in the activity of adults, influenced by the cave’s air humidity (<xref ref-type="bibr" rid="B266">Juberthie 1969</xref>), and sometimes with two distinct annual peaks (<xref ref-type="bibr" rid="B85">Cabidoche 1963</xref>, 1966). Reproduction may coincide with a peak in food, as found between <italic><named-content content-type="taxon-name">Neaphenops tellkampfi</named-content></italic> and the eggs of the orthopteran <italic><named-content content-type="taxon-name">Hadenoecus subterraneus</named-content></italic> (<xref ref-type="bibr" rid="B272">Kane et al. 1975</xref>).</p><p>The seven winter breeding species found in the Netherlands (see above) connect the carabid fauna of the temperate zone with the life history strategy typical for the subtropics with winter rain. In Palestine, <xref ref-type="bibr" rid="B53">Bodenheimer (1934)</xref> only caught beetles from October to June. Winter breeding (rainy season breeding) is a typical reproductive strategy in habitats that are dry in the summer, such as North Africa (<xref ref-type="bibr" rid="B431">Paarmann 1970</xref>, 1975). In specific habitats with moist soil during the dry summer period, propagation and reproduction occur throughout the year (<xref ref-type="bibr" rid="B434">Paarmann 1975</xref>, 1976d). <italic><named-content content-type="taxon-name">Thermophilum sexmaculatum</named-content></italic> and <italic><named-content content-type="taxon-name">Graphipterus serrator</named-content></italic>, with specialised larvae that feed on ants and their brood, reproduce in the summer (<xref ref-type="bibr" rid="B440">Paarmann 1985</xref>; <xref ref-type="bibr" rid="B441">Paarmann et al. 1986</xref>; <xref ref-type="bibr" rid="B152">Dinter et al. 2002</xref>), but only in sandy soil that acts as a moisture trap.</p><p>In Mediterranean Europe, which is also dry in the summer, some seed-feeding carabids - the ditomines <italic><named-content content-type="taxon-name">Carterus calydonius</named-content></italic>, <italic><named-content content-type="taxon-name">Ditomus clypeatus</named-content></italic>, and harpaline carabid beetles that provide <italic><named-content content-type="taxon-name">Daucus</named-content></italic> or <italic><named-content content-type="taxon-name">Plantago</named-content></italic> seeds to their larvae (<xref ref-type="bibr" rid="B66">Brandmayr and Zetto Brandmayr 1974</xref>; <xref ref-type="bibr" rid="B507">Schremmer 1960</xref>) - show summer reproduction. Other seed-feeders (<italic><named-content content-type="taxon-name">Ophonus</named-content></italic>, <italic><named-content content-type="taxon-name">Pseudoophonus</named-content></italic>) are adapted to more humid soils and normally reproduce in the autumn (winter larvae; <xref ref-type="bibr" rid="B605">Zetto Brandmayr 1983a</xref>, b).</p><p>No information is available on the reproductive strategies of <named-content content-type="taxon-name">Carabinae</named-content> from the subtropics with summer rain. However, it seems reasonable to suggest that they show rainy season propagation (summer breeding) in habitats which are dry during winter. A number of studies on carabid beetle life histories are available from the tropics. In Central Africa (Kivu district), which is characterised by low variation in median air temperature (0.9 °C) and low rainfall from June-August (<xref ref-type="bibr" rid="B591">Walter and Lieth 1960</xref>), the majority of species avoid reproduction during and around the dry season (<xref ref-type="bibr" rid="B436">Paarmann 1976b</xref>). Dry season propagation was only found in two species, one living in a swamp and one in a cultivated area. North Sulawesi (Indonesia) is without a dry period, yet the appearance of gonad dormancies was widespread among 155 carabid beetle species: 65% had at least one dormant female (<xref ref-type="bibr" rid="B449">Paarmann and Stork 1987</xref>; <xref ref-type="bibr" rid="B527">Stork and Paarmann 1992</xref>). Females of the canopy dweller <italic><named-content content-type="taxon-name">Colpodes buchanani</named-content></italic> also synchronise reproduction with annual temperature changes typical of the subtropical climate (<xref ref-type="bibr" rid="B448">Paarmann and Paarmann 1997</xref>).</p><p><pmc-comment>PageBreak</pmc-comment>Along the Amazon River in Brazil, forests are often inundated for up to seven months of the year. This flooding is independent of the rainy season in central Amazonia. During flooding, carabid beetles occur on tree trunks or in the canopy in the inundated site, reproducing when the water level is low (<xref ref-type="bibr" rid="B1">Adis et al. 1986</xref>; <xref ref-type="bibr" rid="B2">Adis et al. 1990</xref>). In lowland rainforests, carabids aggregate in areas with an accumulated amount of organic matter, such as fruit falls (<xref ref-type="bibr" rid="B175">Erwin 1979b</xref>). These fruit falls are unpredictable in space and time, lasting only for a few weeks. Fig fruit falls play an important role in these rainforests, as they occur virtually throughout the year. Distinct carabid assemblages have been found at fig fruit falls in lowland rainforests of the Amazon basin (<xref ref-type="bibr" rid="B445">Paarmann et al. 2001</xref>), Brunei (<xref ref-type="bibr" rid="B57">Borcherding et al. 2000</xref>), Australia and Africa (<xref ref-type="bibr" rid="B447">Paarmann et al. 2006</xref>). Female gonad maturation starts immediately after locating a fruit fall, with some females carrying ripe eggs combined with the undeveloped ovaries. These ‘transport eggs’ can be deposited directly after arrival at the fruit fall, providing larvae more time for development. While moving between patches of fruit fall, females experience short gonad dormancy induced by food shortages (<xref ref-type="bibr" rid="B447">Paarmann et al. 2001</xref>; <xref ref-type="bibr" rid="B17">Arndt and Kirmse 2002</xref>).</p><p><bold>Proximate factors and endogenous rhythms</bold></p><p>During unstable temperatures, soil humidity and resources, proximate factors and endogenous rhythms play a major role in controlling carabid beetle life cycles. At temperate latitudes, many species, especially species with summer larvae, use photoperiodic changes to synchronise gonad maturation (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>). Autumn breeding species display thermic parapause (<xref ref-type="bibr" rid="B391">Müller 1970</xref>): an obligatory dormancy at a genetically fixed developmental stage, where the phase of induction cannot be recognised. Larval development can only be completed after passing a certain period of time at low temperatures. Larvae of other species with winter larvae, such as <italic><named-content content-type="taxon-name">Abax ovalis</named-content></italic> and <italic><named-content content-type="taxon-name">Abax parallelepipedus</named-content></italic>, only pass a thermic quiescence (<xref ref-type="bibr" rid="B391">Müller 1970</xref>): a facultative delay or suspension of development. This may also be the case for species with winter larvae at higher latitudes (and montane regions): Subarctic populations of <italic><named-content content-type="taxon-name">Pterostichus nigrita</named-content></italic> were still under photoperiodic control in terms of gonad maturation, yet displayed a shift of the response curve to longer day lengths (<xref ref-type="bibr" rid="B188">Ferenz 1975</xref>).</p><p>Annual day length amplitudes decrease from higher latitudes to the equator, as does the importance of photoperiodic changes as a proximate factor. However, day length changes of 1 h can control imaginal diapause (<xref ref-type="bibr" rid="B424">Norris 1959</xref>, 1965). Two carabid species from North Africa synchronise their life cycle with annual rainfall, triggered by a decrease in temperature and a decline in the photoperiod (<xref ref-type="bibr" rid="B433">Paarmann 1974</xref>, 1976c). This control mechanism in a rainy season breeder (or winter breeder) of the subtropics with winter rain shows marked similarities with temperate autumn breeders and aestivation (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>).</p><p>In the Kivu region, Central Africa (see <xref ref-type="bibr" rid="B436">Paarmann 1976b</xref>), the maximum change in daylight is 16 min only, and the maximum annual temperature change is 0.9°C.<pmc-comment>PageBreak</pmc-comment> Under such climatic conditions, temperature plays a role as a proximate factor. The temperature of the upper soil layers and the soil surface is influenced by the water content of the soil. With water loss in the upper soil layers, daily temperature fluctuations increase. Some hours of higher temperatures per day induce gonad dormancy. With the onset of rainfall, temperature fluctuations decline and dormancy is terminated. Synchronised maturation is stimulated by the increase in average temperatures (<xref ref-type="bibr" rid="B441">Paarmann 1986</xref>).</p><p><bold>Endogenous control of gonad dormancies</bold></p><p>The synchronisation of gonad maturation with seasonal change in ultimate factors is possible only if proximate factors influence the endocrine system controlling this maturation. <xref ref-type="bibr" rid="B170">Emmerich and Thiele (1969)</xref> and <xref ref-type="bibr" rid="B240">Hoffmann (1969)</xref> were the first to study the hormonal control of gonad maturation in spring breeders. They found a connection between proximate factors, neurosecretions and the activity of the <italic>corpora allata</italic>, which produces juvenile hormones (JH). JHs are necessary to complete gonad maturation in males (<xref ref-type="bibr" rid="B190">Ferenz and Hölters 1975</xref>). In females, only previtellogenesis is controlled by JHs. To complete ovarian maturation, the production of a second hormone is postulated. Applications of JHs to dormant beetles of the winter breeder <italic><named-content content-type="taxon-name">Orthomus barbarus</named-content></italic> have confirmed a similar control mechanism for this breeding type (<xref ref-type="bibr" rid="B435">Paarmann 1976a</xref>). The same application to dormant beetles of the summer breeder <italic><named-content content-type="taxon-name">Pogonus chalceus</named-content></italic> resulted in complete maturation of both sexes, even complete gonad maturation in females, meaning that either complete maturation is controlled by JHs only, or high temperatures suppress only the production of JHs but not of vitellogenic hormones.</p><p>Endogenous rhythms are involved in gonad maturation. Under constant environmental conditions gonad maturation is controlled by an endogenous rhythm, synchronised by an external cue such as soil temperature (<xref ref-type="bibr" rid="B441">Paarmann 1986</xref>). In the desert-dwelling carabid beetle <italic><named-content content-type="taxon-name">Thermophilum sexmaculatum</named-content></italic> thermoregulational behaviour is controlled by a circannual rhythm, resulting in lower body temperatures at the end of the optimal reproductive period, which causes an inactive stage of the gonads (<xref ref-type="bibr" rid="B171">Erbeling and Paarmann 1986</xref>).</p><p>As part of the taxon pulse theory (<xref ref-type="bibr" rid="B175">Erwin 1979b</xref>), ground beetles from tropical areas undergo latitudinal and altitudinal expansion, leading to climatic specialisation, including the development of dormancy to survive unfavourable climatic conditions. If all carabid beetle dormancies are based on a uniform hormonal system, manifold convergent evolution is possible. The use of gonad dormancies to synchronise life cycles with changing environmental conditions is widespread among tropical carabid beetles. Only one <italic><named-content content-type="taxon-name">Abacetus</named-content></italic> species, living under stable humidity and temperature conditions (the shore of Lake Kivu, Central Africa), seems to develop without dormancy. With the exception of short gonad dormancies, triggered by food shortages in the seed-feeding guild, all studied gonad dormancies are under the control of temperature as a proximate factor.</p><p>Specialisation along riparian habitats (pathway i) leads to a synchronisation of the life cycle with seasons with stable moisture conditions, especially along riverbanks. <pmc-comment>PageBreak</pmc-comment>Specialisation in seasonally dry habitats (pathway ii) leads to a synchronisation of the life cycle with the period of optimal soil humidity, e.g. rainy season propagation (<xref ref-type="bibr" rid="B439">Paarmann 1979</xref>). While larvae of winter breeders in the subtropics with winter rainfall are adapted to comparable temperatures, a small group requires high temperatures for successful development. These specialists, whose larvae feed on ants and ant brood, have evolved along pathway (ii) in the subtropics with summer rainfall and spending the winter in gonad dormancy. Such species have yet to be reported in the temperate zone.</p><p><xref ref-type="bibr" rid="B310">Larsson (1939)</xref> found no autumn breeders among 21 studied species of the old genus <italic><named-content content-type="taxon-name">Agonum</named-content></italic>. These species are possibly all descendants of one common ancestor that reached the temperate zone along pathway (i) after which some descendant species adapted to non-riparian habitats. One member of this group, namely <italic><named-content content-type="taxon-name">Platynus</named-content></italic> (<italic><named-content content-type="taxon-name">Agonum</named-content></italic>, <italic><named-content content-type="taxon-name">Limodromus</named-content></italic>) <italic>assimilis</italic> is a spring breeder, but its gonad dormancy is controlled in a fundamentally different way than in other spring breeders, by a photoperiodic quiescence (<xref ref-type="bibr" rid="B401">Neudecker and Thiele 1974</xref>).</p><p>Gaps in our current understanding of carabid beetle life history strategies include (i) a lack of knowledge on life history strategies in the subtropics with summer rainfall, in the tropics with long dry seasons and in areas with unpredictable rainfall, (ii) whether canopy dwelling carabid beetles in tropical rainforests display seasonal patterns, and (iii) a detailed study on the hormonal control of dormancies in carabid beetles, as no such studies have been performed since <xref ref-type="bibr" rid="B189">Ferenz (1977)</xref>.</p></sec><sec sec-type="3.2. Carabid beetle food"><title>3.2. Carabid beetle food</title><p>Carabid beetles are generally considered polyphagous predators. However, in line with their enormous species richness and diversity in body shapes and biotopes they inhabit, a whole range of trophic specialisations occurs in the <named-content content-type="taxon-name">Carabidae</named-content> (<xref ref-type="bibr" rid="B234">Hengeveld 1980a</xref>; <xref ref-type="bibr" rid="B611">Zetto Brandmayr et al. 1998b</xref>). Although carabid feeding ecology and biology has been studied frequently (also during ECM meetings), it is surprising how many basic questions on carabid food remain unanswered. Except for <xref ref-type="bibr" rid="B308">Larochelle (1990)</xref>, who mentioned food preferences of 1054, mainly North-American, European and Japanese species, basic information on food preferences or requirements is often lacking, even for many common species. This chapter does not attempt to review all trophic specialisations of <named-content content-type="taxon-name">Carabidae</named-content>; it has been done before (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>; <xref ref-type="bibr" rid="B234">Hengeveld 1980a</xref>; <xref ref-type="bibr" rid="B555">Toft and Bilde 2002</xref>). Instead, it focuses on recent advances in the domains of seed and ant feeding, as well as unique life history strategies, such as ectoparasitism and the predation of amphibians.</p><p><bold>Seed feeding</bold></p><p>Carabid beetles accept a variety of plant foods such as leaves, fruits, pollen, seeds and fungi (<xref ref-type="bibr" rid="B555">Toft and Bilde 2002</xref> and references therein). Seed feeding, or granivory, occurs in many species including polyphagous ones that prefer animal prey (<xref ref-type="bibr" rid="B343">Lund and Turpin 1977</xref>;<pmc-comment>PageBreak</pmc-comment> <xref ref-type="bibr" rid="B235">Hengeveld 1980b</xref>; <xref ref-type="bibr" rid="B555">Toft and Bilde 2002</xref>). True granivory, i.e. where seeds are central to the species’ food budget, has evolved in two tribes of <named-content content-type="taxon-name">Carabidae</named-content>, <named-content content-type="taxon-name">Zabrini</named-content> and <named-content content-type="taxon-name">Harpalini</named-content>. The ecology of granivorous carabids is of great interest since granivory required the evolution of morphological, physiological and behavioural adaptations associated with crushing, digesting and foraging for seeds. To crush hard seeds, adults and larvae of granivorous species have evolved broad mandibles with massive adductors (<xref ref-type="bibr" rid="B611">Zetto Brandmayr et al. 1998b</xref>; <xref ref-type="bibr" rid="B447">Paarmann et al. 2006</xref>). Sclerotised structures in the adult proventriculus are then used for fine grinding of the ingested seed fragments (<xref ref-type="bibr" rid="B181">Evans and Forsythe 1985</xref>). Behavioural adaptations have involved, for example, climbing plants and storing seeds in burrows (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>). Physiological adaptations to seed feeding are understudied but recent evidence shows that digestion of seeds is facilitated by endosymbionts (<xref ref-type="bibr" rid="B344">Lundgren and Lehman 2010</xref>).</p><p>The amount of seeds eaten by carabids in the field may be substantial. Based on seed losses of artificially exposed seeds, <xref ref-type="bibr" rid="B245">Honek et al. (2003)</xref> estimated that up to 4000 seeds m-2 d-1 may be removed by carabid beetles in arable fields in the Czech Republic. <xref ref-type="bibr" rid="B246">Honek et al. (2005)</xref> reported that carabids, mainly <italic><named-content content-type="taxon-name">Amara montivaga</named-content></italic>, destroyed about 83–88% of the annual seed production of <italic><named-content content-type="taxon-name">Taraxacum officinale</named-content></italic> spp. agg., and <xref ref-type="bibr" rid="B280">Kjellsson (1985)</xref> showed that approximately 65% of the annual seed production of <italic><named-content content-type="taxon-name">Carex pilulifera</named-content></italic> L. was consumed by a single species, <italic><named-content content-type="taxon-name">Harpalus solitaris</named-content></italic>. However, individual capacity for eating seeds varies with season (<xref ref-type="bibr" rid="B249">Honek et al. 2006</xref>) as a result of natural phenological changes (transition from dormancy to reproduction, dispersal, breeding and searching for overwintering sites). Consumption is also affected by temperature (<xref ref-type="bibr" rid="B502">Saska et al. 2010</xref>). Clearly, carabid beetles may have an important impact on the reproductive success and dispersal of plant species, but more research is needed on how these affect the population dynamics of plants in the longer term. Larvae should also be considered in these studies, as their consumption of seeds can be comparable to that of adults (<xref ref-type="bibr" rid="B282">Klimeš and Saska 2010</xref>).</p><p>The consumption of particular seed species is ultimately determined by the preferences of the carabids in question. During the last 30 years, a number of authors have investigated carabid preferences for seeds in the laboratory using choice (cafeteria) experiments (<xref ref-type="bibr" rid="B343">Lund and Turpin 1977</xref>; <xref ref-type="bibr" rid="B79">Brust and House 1988</xref>; <xref ref-type="bibr" rid="B264">Jørgensen and Toft 1997a</xref>). Most studies, however, have established preferences based on a limited number of seed species (usually 2–5). Only <xref ref-type="bibr" rid="B245">Honek et al. (2003</xref>, <xref ref-type="bibr" rid="B249">2006</xref>, 2007) tested seed preferences in carabids using 64 or 28 species of herbaceous seed. <xref ref-type="bibr" rid="B245">Honek et al. (2003</xref>, 2007, 2011) demonstrated that the preference for seeds correlates with carabid body size: on average, smaller species prefer smaller seeds, and <italic>vice versa</italic>. Larger carabids also consume a greater variety of seed species and <named-content content-type="taxon-name">Harpalini</named-content> are less specialised than <named-content content-type="taxon-name">Zabrini</named-content> (<xref ref-type="bibr" rid="B248">Honek et al. 2007</xref>). However, there are other characters such as seed shape, thickness of the testa (<xref ref-type="bibr" rid="B345">Lundgren and Rosentrater 2007</xref>) and nutrient content of the seed that affect preference. Similarly to other seed-cracking organisms (e.g. <xref ref-type="bibr" rid="B147">Diaz 1994</xref>), mandible size and shape determine the seed preferences of <italic><named-content content-type="taxon-name">Notiobia</named-content></italic> species occupying fruit fall sites in tropical forests (<xref ref-type="bibr" rid="B17">Arndt and Kirmse 2002</xref>; <xref ref-type="bibr" rid="B447">Paarmann et al. 2006</xref>), and these preferences are consistent throughout the season (<xref ref-type="bibr" rid="B249">Honek et al. 2006</xref>).</p><p><pmc-comment>PageBreak</pmc-comment>Taxonomic affiliation constrains the preferences for food in many insect groups. Earlier research as well as direct field observations have indicated that species of certain genera had specific affinities with respect to their seed preferences. For example, <xref ref-type="bibr" rid="B69">Brandmayr and Zetto Brandmayr (1987)</xref> and <xref ref-type="bibr" rid="B607">Zetto Brandmayr (1990)</xref> suggested that most <italic><named-content content-type="taxon-name">Ditomina</named-content></italic> and <italic><named-content content-type="taxon-name">Ophonus</named-content></italic> (both <named-content content-type="taxon-name">Harpalini</named-content>) are associated with <named-content content-type="taxon-name">Apiaceae</named-content>, while <italic><named-content content-type="taxon-name">Harpalus</named-content></italic> (<named-content content-type="taxon-name">Harpalini</named-content>) is unspecialised in this sense (<xref ref-type="bibr" rid="B607">Zetto Brandmayr 1990</xref>). <xref ref-type="bibr" rid="B252">Hurka (1996)</xref> reported that species of the subgenus <italic><named-content content-type="taxon-name">Zezea</named-content></italic> (<named-content content-type="taxon-name">Zabrini</named-content>: <italic><named-content content-type="taxon-name">Amara</named-content></italic>) may be associated with <named-content content-type="taxon-name">Poaceae</named-content>. The existence of a taxonomic constraint has been experimentally confirmed by <xref ref-type="bibr" rid="B248">Honek et al. (2007)</xref>, who carried out a cafeteria experiment that included 28 seed species and 30 carabid species. They demonstrated that species of <named-content content-type="taxon-name">Zabrini</named-content> mostly prefer seeds of <italic><named-content content-type="taxon-name">Taraxacum</named-content></italic>, while species of <named-content content-type="taxon-name">Harpalini</named-content> prefer seeds of <italic><named-content content-type="taxon-name">Cirsium</named-content></italic> and Viola. Carabids not only distinguished seeds from different families, but they were also able to discriminate between seeds at a finer taxonomic scale, i.e. seeds of different sections of the <italic><named-content content-type="taxon-name">Taraxacum officinale</named-content></italic> species complex (<xref ref-type="bibr" rid="B247">Honek et al. 2011</xref>). The origin of seeds plays a role in some carabid species. For example, <xref ref-type="bibr" rid="B247">Honek et al. (2011)</xref> fed Czech carabids with Italian and Czech seeds of the same plant species and found that the beetles preferred the latter. It is likely that the existence of specialisation on particular seeds reduces the competition for food and allows the coexistence of species in the same habitat.</p><p>Seeds are nutritious, but their value as food for carabids has not been appropriately recognised until recently. The value of food is best defined by its contribution to the fitness of the consumer (<xref ref-type="bibr" rid="B555">Toft and Bilde 2002</xref>). Fitness parameters that are commonly used as criteria for the evaluation of food quality are female fecundity, survival and duration of larval development, and the attainable body size. <xref ref-type="bibr" rid="B604">Zetto Brandmayr (1976)</xref> showed better survival in larvae of several species of the genus <italic><named-content content-type="taxon-name">Ophonus</named-content></italic> when provided with seeds of <named-content content-type="taxon-name">Apiaceae</named-content> compared to other seeds or insects. Although Jørgensen and Toft (1997a, b) stimulated further research on this topic (mainly in Europe and Japan), information on how seed diet affects fitness is only available for a small number of species. Adaptations to granivory have evolved to varying degrees in different taxa, and even closely related species may show different strategies (for <italic><named-content content-type="taxon-name">Amara</named-content></italic>, subgenus <italic><named-content content-type="taxon-name">Amara</named-content></italic>, compare e.g. <xref ref-type="bibr" rid="B265">Jørgensen and Toft 1997b</xref>; Saska and Jarošík<xref ref-type="bibr" rid="B204"> 2001</xref>; <xref ref-type="bibr" rid="B253">Hurka and Jarošík 2003</xref>; <xref ref-type="bibr" rid="B186">Fawki and Toft 2005</xref>; <xref ref-type="bibr" rid="B497">Saska 2008</xref>; for <italic><named-content content-type="taxon-name">Amara</named-content></italic>, subgenus <italic><named-content content-type="taxon-name">Curtonotus</named-content></italic>, compare e.g. <xref ref-type="bibr" rid="B496">Saska 2005</xref>; <xref ref-type="bibr" rid="B494">Sasakawa 2007</xref>; 2009); for <italic><named-content content-type="taxon-name">Notiobia</named-content></italic>, see <xref ref-type="bibr" rid="B18">Arndt et al. 1996</xref>; <xref ref-type="bibr" rid="B447">Paarmann et al. 2001</xref>; <xref ref-type="bibr" rid="B17">Arndt and Kirmse 2002</xref>). More interestingly, particular seed diets may have contrasting effects on different fitness traits (<xref ref-type="bibr" rid="B186">Fawki and Toft 2005</xref>). The effects of maternal diet (Saskawa 2009) or diet of the previous generations (<xref ref-type="bibr" rid="B253">Hurka and Jarošík 2003</xref>) on larval performance are poorly studied. Also, worthy of mention here is the scoring system of <xref ref-type="bibr" rid="B444">Paarmann (2002)</xref> used to evaluate larval performance under different dietary regimes. In general, larvae are more specific in their food preferences than adults (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>) because of increased selection pressures on larvae (<xref ref-type="bibr" rid="B494">Sasakawa 2007</xref>) and due to morphological constraints on the suitability of the available food during the early stages of development (<xref ref-type="bibr" rid="B447">Paarmann et al. 2006</xref>). <xref ref-type="bibr" rid="B282">Klimeš and Saska (2010)</xref> argued that this selection pressure is highest in the first instar larva and decreases in older instars, with increasing the head width/seed size ratio in larvae and widening the range of edible food items.<pmc-comment>PageBreak</pmc-comment></p><p><bold>Ant feeding</bold></p><p>Ants are the most abundant group of organisms on Earth in terms of biomass (<xref ref-type="bibr" rid="B243">Hölldobler and Wilson 1990</xref>). Not surprisingly they represent an important food source for many other taxa, including carabid beetles. Polyphagous carabid species frequently prey on ants (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>; <xref ref-type="bibr" rid="B235">Hengeveld 1980b</xref>), and several clades have adapted to ant feeding with some having evolved the highest degree of specialisation, i.e. myrmecophily. In general, biological information is very limited and needs systematic study.</p><p>Species that have adapted to feeding on ants have evolved interesting behavioural and morphological adaptations, including chemical mimicry that reduces the risk of being attacked by their hosts (<xref ref-type="bibr" rid="B608">Zetto Brandmayr et al. 2000a</xref>; <xref ref-type="bibr" rid="B152">Dinter et al. 2002</xref>). Larvae of <italic><named-content content-type="taxon-name">Sphallomorpha</named-content></italic> (<named-content content-type="taxon-name">Pseudomorphini</named-content>) form burrows close to ant nests and attack ants that pass by (<xref ref-type="bibr" rid="B381">Moore 1974</xref>). Associations with ants and termites seem to be a joint character for the entire tribe of <named-content content-type="taxon-name">Pseudomorphini</named-content>, though evidence is limited (<xref ref-type="bibr" rid="B33">Baehr 1994</xref>). Species of the <named-content content-type="taxon-name">Siagonini</named-content> also prey on ants ( 1998a, <xref ref-type="bibr" rid="B609"></xref>). Species of the genus <italic><named-content content-type="taxon-name">Siagona</named-content></italic> inhabit crevices in the soil near ant nests and attack ants both as adults and larvae, but do not seem to enter ant nests frequently (<xref ref-type="bibr" rid="B46">Bauer et al. 2005</xref>). The larvae of some <named-content content-type="taxon-name">Ozaeini</named-content> use so-called terminal disks (modified last abdominal segments) for attracting and capturing ants (<xref ref-type="bibr" rid="B150">Di Giulio and Vigna Taglianti 2001</xref>; <xref ref-type="bibr" rid="B383">Moore and Di Giulio 2006</xref>).</p><p>Adults of the North African <named-content content-type="taxon-name">Anthiini</named-content> and <named-content content-type="taxon-name">Graphipterini</named-content> are free-living but larvae enter ant nests where they prey upon ants to complete their development (<xref ref-type="bibr" rid="B440">Paarmann 1985</xref>; <xref ref-type="bibr" rid="B441">Paarmann et al. 1986</xref>). The larva of <italic><named-content content-type="taxon-name">Thermophilum</named-content></italic> (<named-content content-type="taxon-name">Anthiini</named-content>) moves freely in the nest after it gains chemical mimicry from ants it has previously attacked (<xref ref-type="bibr" rid="B152">Dinter et al. 2002</xref>), and consumes both ants and ant brood (Paarmann and Erbeling 1986). In contrast, the larva of <italic><named-content content-type="taxon-name">Graphipterus serrator</named-content></italic> forms a chamber inside the ant nest where it stores ant brood before consumption, and hides against ant attacks (<xref ref-type="bibr" rid="B152">Dinter et al. 2002</xref>). Species of <italic><named-content content-type="taxon-name">Thermophilum</named-content></italic>, as well as <italic><named-content content-type="taxon-name">Graphipterus serrator</named-content></italic>, show preferences for particular ant species, <italic><named-content content-type="taxon-name">Graphipterus</named-content></italic> being the least selective (<xref ref-type="bibr" rid="B152">Dinter et al. 2002</xref>).</p><p>True myrmecophily (and perhaps termitophily) evolved in the tribe <named-content content-type="taxon-name">Paussini</named-content>, in which morphological and behavioural adaptations are prominent in both adults and larvae (<xref ref-type="bibr" rid="B395">Nagel 1979b</xref>; <xref ref-type="bibr" rid="B149">Di Giulio and Moore 2004</xref>; <xref ref-type="bibr" rid="B383">Moore and Di Giulio 2006</xref>). Although this association is well known, data on food requirements or trophic associations are known for a limited number of taxa only, and this requires further investigation.</p><p><bold>Unique life history strategies – ectoparasitism and the predation of amphibians</bold></p><p>The variety of life history strategies in carabid beetles includes ectoparasitoidism, a strategy otherwise rare in beetles. Parasitoids are insects whose larvae develop at the expense of a single prey individual (a host), which ultimately dies as a result of parasitoid feeding (<xref ref-type="bibr" rid="B585">Vinson 1976</xref>). Ectoparasitoid larvae attach to the host body and feed externally on it, while their adults are free-living (<xref ref-type="bibr" rid="B585">Vinson 1976</xref>).</p><p>Ectoparasitoidism has been described from four carabid genera: <italic><named-content content-type="taxon-name">Brachinus</named-content></italic> (<named-content content-type="taxon-name">Brachinini</named-content>), <italic><named-content content-type="taxon-name">Pelecium</named-content></italic> (<named-content content-type="taxon-name">Peleciini</named-content>), <italic><named-content content-type="taxon-name">Lebia</named-content></italic> and <italic><named-content content-type="taxon-name">Lebistina</named-content></italic> (both <named-content content-type="taxon-name">Lebiini</named-content>) (<xref ref-type="bibr" rid="B593">Weber et al. 2008</xref>), but several related genera show tendencies towards parasitoidism (<xref ref-type="bibr" rid="B174">Erwin 1979a</xref>; <xref ref-type="bibr" rid="B199">Frank et al. 2009</xref>). The life cycle of a typical carabid ectoparasitoid includes (i) a female depositing eggs in the host habitat when hosts are present; (ii) mobile early instar larva searching for and attaching to a suitable host; (iii) after attachment, a short physogastric feeding phase, typically with rapid ingestion; and (iv) a distinct pre-pupal “resting” phase during which the host is consumed.</p><p>Despite the early discovery of ectoparasitoidism in <named-content content-type="taxon-name">Carabidae</named-content> (e.g. <xref ref-type="bibr" rid="B597">Wickham 1893</xref>; <xref ref-type="bibr" rid="B520">Silvestri 1904</xref>), known host associations are few. With one known exception, beetle pupae are the hosts. Larvae of <italic><named-content content-type="taxon-name">Lebia</named-content></italic> (five species known to be parasitoids) and <italic><named-content content-type="taxon-name">Lebistina</named-content></italic> (one species) parasitise leaf beetle (<named-content content-type="taxon-name">Chrysomelidae</named-content>) pupae (<xref ref-type="bibr" rid="B593">Weber et al. 2008</xref>). Larvae of a single undetermined species of <italic><named-content content-type="taxon-name">Pelecium</named-content></italic> have been observed developing on chrysomelid pupae and millipedes (<xref ref-type="bibr" rid="B489">Salt 1928</xref>). Nearctic wetland species of <italic><named-content content-type="taxon-name">Brachinus</named-content></italic> (seven species) parasitise the pupae of water beetles (<named-content content-type="taxon-name">Dytiscidae</named-content>, <named-content content-type="taxon-name">Gyrinidae</named-content>, <named-content content-type="taxon-name">Hydrophilidae</named-content>) (Saska and Honek<xref ref-type="bibr" rid="B205"> 2004</xref>). Despite suggestions proposed by Jeannel (1942), the discovery of the hosts for dryland species of <italic><named-content content-type="taxon-name">Brachinus</named-content></italic> from Europe was only made 60 years later. Saska and Honek(<xref ref-type="bibr" rid="B205"> (2004</xref>, 2005) successfully reared two species (<italic><named-content content-type="taxon-name">Brachinus explodens</named-content></italic> and <italic><named-content content-type="taxon-name">Brachinus crepitans</named-content></italic>) on the pupae of another carabid genus, <italic><named-content content-type="taxon-name">Amara</named-content></italic>, a finding that has recently been confirmed for <italic><named-content content-type="taxon-name">Brachinus elegans</named-content></italic> by <xref ref-type="bibr" rid="B357">Makarov and Bokhovko (2005)</xref>.</p><p>Besides direct observations, host-parasite associations have frequently been suggested simply on the basis of co-occurrence of the carabid parasitoid and potential host species. In some cases, however, these observations have led to erroneous predictions (Jeannel 1942; <xref ref-type="bibr" rid="B457">Perez-Zaballos 1985</xref>), subsequently refuted because the life cycles of the two suggested partners are not synchronous. Such synchrony has so far been demonstrated only for <italic><named-content content-type="taxon-name">Brachinus explodens</named-content></italic> and <italic><named-content content-type="taxon-name">Brachinus crepitans</named-content></italic> (<xref ref-type="bibr" rid="B500">Saska and Honek 2008</xref>). Thus, when looking for hosts of <italic><named-content content-type="taxon-name">Mastax</named-content></italic> or <italic><named-content content-type="taxon-name">Aptinus</named-content></italic> (both <named-content content-type="taxon-name">Brachinini</named-content>) or wetland Palaearctic <italic><named-content content-type="taxon-name">Brachinus</named-content></italic> species, both co-occurrence and synchrony should be taken into account. More discoveries are probably to be made in the tropics, as that climatic zone contains a vast diversity of lebiine carabids (<xref ref-type="bibr" rid="B426">Ober and Maddison 2008</xref>). Research is also needed on the ecology of ectoparasitic carabids to determine the adaptive significance of life history traits of this peculiar strategy. In most cases, available information relates to a brief description of development; only a few species have been studied in detail (<xref ref-type="bibr" rid="B173">Erwin 1967</xref>; <xref ref-type="bibr" rid="B268">Juliano 1985</xref>; Saska and Honek<xref ref-type="bibr" rid="B205"> 2004</xref>, 2005, 2008; <xref ref-type="bibr" rid="B594">Weber et al. 2006</xref>). Host selection, food utilisation or the adaptive significance of variation in the number of instars (2–5 instead of the typical 3) could produce interesting results. Mimetic complexes have been described between adults of <italic><named-content content-type="taxon-name">Lebia</named-content></italic> and chrysomelids, including species for which parasitoidism is unknown (<xref ref-type="bibr" rid="B231">Hemenway and Whitcomb 1967</xref>), suggesting further trophic associations between the two groups. Focusing on taxa representing transitional evolutionary steps to parasitoidism (<xref ref-type="bibr" rid="B174">Erwin 1979a</xref>; <xref ref-type="bibr" rid="B199">Frank et al. 2009</xref>) may shed light on the evolution of parasitoidism in <named-content content-type="taxon-name">Carabidae</named-content> and in <named-content content-type="taxon-name">Coleoptera</named-content> in general.</p><p>Carabid beetle larval and adult predation on amphibians has recently been described in Israel. <xref ref-type="bibr" rid="B169">Elron et al. (2007)</xref> have shown that larvae of the carabid <italic><named-content content-type="taxon-name">Epomis dejeani</named-content><pmc-comment>PageBreak</pmc-comment></italic> preyed upon two amphibian species (<italic><named-content content-type="taxon-name">Bufo viridis</named-content></italic> and <italic><named-content content-type="taxon-name">Hyla savignyi</named-content></italic>), confirming an earlier brief note by <xref ref-type="bibr" rid="B380">Moore (1971)</xref> from Australia. Subsequently, <xref ref-type="bibr" rid="B598">Wizen and Gasith (2011)</xref> performed laboratory experiments, and showed that adults of the two sympatric <italic><named-content content-type="taxon-name">Epomis</named-content></italic> species in Israel, <italic><named-content content-type="taxon-name">Epomis dejeani</named-content></italic> and <italic><named-content content-type="taxon-name">Epomis circumscriptus</named-content></italic>, prey upon five and four amphibian species, respectively. <xref ref-type="bibr" rid="B598">Wizen and Gasith (2011)</xref> argue that little is known about the feeding habits of sympatric congeneric insects, and that the partial food overlap of these <italic><named-content content-type="taxon-name">Epomis</named-content></italic> species warrants further investigation.</p></sec><sec sec-type="3.3. Dispersal"><title>3.3. Dispersal</title><p>Carabid beetles found a rich niche in ecological research through the peculiarities of their dispersal power. <xref ref-type="bibr" rid="B488">Sahlberg (1868)</xref> recognised that carabid species exhibit a variety of wing attributes, including wing dimorphism, and that this has implications for their powers of dispersal. Darwin was probably the first to consider the evolutionary and ecological implications of wing polymorphism in <named-content content-type="taxon-name">Coleoptera</named-content> after recording high proportions of flightless beetles on the island of Madeira. He hypothesized that flight ability might be evolutionarily disadvantageous for species from insular populations, as they would be more likely to get carried away from the island (<xref ref-type="bibr" rid="B109">Darwin 1859</xref>). A few decades later, Darlington turned his attention to the low proportions of macropterous carabids in isolated locations such as islands and mountain tops, and concluded that wing reduction must confer enhanced viability (<xref ref-type="bibr" rid="B105">Darlington 1936</xref>, 1943). <xref ref-type="bibr" rid="B326">Lindroth (1988, 1992a, b)</xref> studied the wing morphology of carabid assemblages from islands in the Baltic Sea in comparison to control assemblages from nearby mainland sites. He found that the proportions of brachypterous and macropterous species were both lower in insular than in mainland assemblages, even whilst macropterous species were predominant in all of the studied assemblages (see also <xref ref-type="bibr" rid="B20">Ås 1984</xref>, <xref ref-type="bibr" rid="B294">Kotze et al. 2000</xref>). Dimorphic species, on the other hand, were more numerous in insular than in mainland faunas (<xref ref-type="bibr" rid="B326">Lindroth 1988, 1992a, b</xref>). These observations were of fundamental importance to Lindroth’s epic zoogeographical studies, published posthumously in 1992 (<xref ref-type="bibr" rid="B326">Lindroth 1988, 1992a, b</xref>). After determining the frequencies of the different wing morphologies in populations of wing-dimorphic carabid species across the Fennoscandian region, Lindroth was able to estimate the relative ages of these populations. On that basis, he theorised about the routes of post-glacial colonisation of Fennoscandia by different species. He was subsequently able to divide the fauna into three elements: Wűrm hibernators, immigrants from a southern route to the west of the Baltic Sea and immigrants from the east. Both <xref ref-type="bibr" rid="B326">Lindroth (1988, 1992a, b)</xref> and <xref ref-type="bibr" rid="B111">Den Boer (1970)</xref> came to the conclusion that macropterous specimens dominate in recently established populations of dimorphic species, which gradually shift to an increasing proportion of brachypterous individuals as these populations grow older. Observations of pioneering populations of the invasive species <italic><named-content content-type="taxon-name">Pterostichus melanarius</named-content></italic> in Canada support this model (<xref ref-type="bibr" rid="B418">Niemelä and Spence 1991</xref>).</p><p>Largely thanks to the work of Piet den Boer and colleagues, subsequent to the Dutch land reclamation projects of the late 1950s, research interest in the dispersal of carabid beetles flourished, and this provided the theme for the first meeting of European carabidologists in Wijster in 1969, which Piet den Boer hosted (see above). Dispersal power was also the theme of the subsequently published proceedings volume, edited by Den Boer (1971, see also <xref ref-type="table" rid="T2">Table 2</xref>).</p><p>Lindroth was keen to determine the genetic mechanism behind wing dimorphism and conducted breeding experiments with the wing-dimorphic species <italic><named-content content-type="taxon-name">Pterostichus anthracinus</named-content></italic> (<xref ref-type="bibr" rid="B326">Lindroth 1988, 1992a, b</xref>). The results he obtained, supported by similar results from studies of other coleopteran taxa, led him to conclude that wing dimorphism is inherited in a simple Mendelian pattern, in which brachyptery is dominant. The late Konjev Desender, in whose honour the 14th ECM was held, performed similar breeding experiments using the wing polymorphic species <italic><named-content content-type="taxon-name">Pogonus chalceus</named-content></italic>. In this species, crosses between macropterous and brachypterous adults produced offspring with intermediate wing length, suggesting that the genetic control of wing length in this species is polygenic (<xref ref-type="bibr" rid="B128">Desender 1989a</xref>). Desender also conducted an exhaustive biometric study of wing development in 300 carabid species indigenous to Belgium and demonstrated that, in addition to brachypterous individuals, also a large proportion of macropterous individuals do not possess functioning flight muscles and are therefore incapable of flight. In the wing-polymorphic <italic><named-content content-type="taxon-name">Pterostichus vernalis</named-content></italic>, for instance, some populations are entirely macropterous, with functional flight (but see below) muscles even in relatively short-winged individuals, whereas in some other populations even macropterous individuals lack functional flight muscles (<xref ref-type="bibr" rid="B129">Desender 1989b</xref>, see also <xref ref-type="bibr" rid="B399">Nelemans 1987</xref>). Desender also studied wing morphology in the genus <italic><named-content content-type="taxon-name">Calosoma</named-content></italic> after research trips to Easter Island and the Galapagos archipelago. Three endemic species appeared to be brachypterous, whereas the supposedly introduced species, <italic><named-content content-type="taxon-name">Calosoma granatense,</named-content></italic> appeared to be wing polymorphic (<xref ref-type="bibr" rid="B139">Desender et al. 2000</xref>).</p><p>Berend Aukema conducted breeding experiments with the <italic><named-content content-type="taxon-name">Calathus melanocephalus</named-content></italic> group to shed further light on the inheritance of dispersal characteristics. <xref ref-type="bibr" rid="B27">Aukema (1990)</xref> demonstrated that these species show a simple Mendelian pattern of inheritance of wing morphology, as described by Lindroth, i.e. simple inheritance with brachyptery dominant over macroptery for the two wing dimorphic species <italic><named-content content-type="taxon-name">Calathus cinctus</named-content></italic> and <italic><named-content content-type="taxon-name">Calathus melanocephalus</named-content></italic>. However, he also demonstrated that certain environmental factors, such as temperature and food supply, influence expression, with higher temperatures and better food availability resulting in both greater proportion of macropterous individuals (<xref ref-type="bibr" rid="B27">Aukema 1990</xref>), and the development of flight muscles (<xref ref-type="bibr" rid="B399">Nelemans 1987</xref>). Moreover, long-winged females of these two species had greater fecundity than short-winged females, both in terms of quantity of egg production and duration of egg production (<xref ref-type="bibr" rid="B28">Aukema 1991</xref>). This result was somewhat counterintuitive, as a number of other studies of wing dimorphic insects, e.g. <xref ref-type="bibr" rid="B481">Roff (1986)</xref> found that brachypterous females are generally more fecund, suggesting that the advantage conferred by brachyptery is enhanced fecundity for females. Furthermore, macropterous females of <italic><named-content content-type="taxon-name">Pogonus chalceus</named-content></italic> have greater fecundity, suggesting<pmc-comment>PageBreak</pmc-comment> that long wings and functional flight muscles are associated with large body size (<xref ref-type="bibr" rid="B129">Desender 1989b</xref>; <xref ref-type="bibr" rid="B28">Aukema 1991</xref>).</p><p>Work from other invertebrate taxa has suggested that there is a cost in terms of reproductive capacity for flight, with some macropterous females lysing their flight muscles and shedding their wings prior to reproduction, resulting in enhanced reproductive capacity. Among carabids, <italic><named-content content-type="taxon-name">Amara plebeja</named-content></italic> autolyses its wings and can subsequently regenerate them to facilitate migration between breeding and over-wintering habitats (<xref ref-type="bibr" rid="B567">van Huizen 1977</xref>, 1979). This is supported by Matalin’s (1994) observation that reproductive females from window traps invariably have fewer ova than those from pitfall traps. <xref ref-type="bibr" rid="B365">Matalin (1994)</xref> also concluded that the choice between flying and walking varies considerably between species and with different stages in the life cycle, with flight activity being favoured by dispersive young adults, shortly after emergence and, in <italic><named-content content-type="taxon-name">Harpalus rufipes</named-content></italic> and <italic><named-content content-type="taxon-name">Harpalus calceatus</named-content></italic>, by mature males. Mature adults exhibit the highest walking activity during the breeding season, apparently being the favoured form of locomotion when seeking a mate (<xref ref-type="bibr" rid="B365">Matalin 1994</xref>).</p><p>Wing morphology alone is not sufficient to describe dispersal ability in carabids. <xref ref-type="bibr" rid="B130">Desender (2000)</xref> and <xref ref-type="bibr" rid="B370">Matalin (2003)</xref> studied the phenology of carabids in relation to flight muscle development. <xref ref-type="bibr" rid="B130">Desender (2000)</xref> investigated the trade-off between dispersal and reproduction in female carabids from the Belgian fauna, and most of the species he studied supported the oogenesis-flight syndrome, i.e. females with ripe ovaries tend not to possess functional flight musculature. This phenomenon was most pronounced for species that reproduce in late summer or autumn and emerge in late spring (<xref ref-type="bibr" rid="B130">Desender 2000</xref>). <xref ref-type="bibr" rid="B370">Matalin (2003)</xref> concluded that in females of large species, wing muscles decline during a period of increasing body mass, after development of the gonads.</p><p>In addition to the wealth of material on dispersal by flight, carabidologists have also investigated running activity, demonstrating that larger <italic><named-content content-type="taxon-name">Carabus</named-content></italic> species run slower than smaller carabids, though in <named-content content-type="taxon-name">Pterostichinae</named-content> and <named-content content-type="taxon-name">Harpalinae</named-content>, larger species are faster (<xref ref-type="bibr" rid="B388">Mossakowski and Stier 1983</xref>). Temperature has a significant effect on running activity in <italic><named-content content-type="taxon-name">Carabus auronitens</named-content></italic> (<xref ref-type="bibr" rid="B4">Althoff et al. 1994</xref>). Clearly the expression of dispersal ability in carabid beetles is highly complex, being governed by environmental and life cycle factors, in addition to genetic control. It is equally clear that there are still many unresolved issues regarding the dispersal of carabids and we are likely to see studies on this topic at future ECMs. In particular, ongoing land-use change and habitat fragmentation, exacerbated by the influence of climate change, mean stronger selective advantages for species with better powers of dispersal. A major challenge for the scientific community will be to discern evolutionary changes in response to this selective pressure. In conservation, the main challenge will be to develop strategies for the conservation of species with poor powers of dispersal.</p></sec></sec><sec><title>4 Methods</title><sec sec-type="4.1. Methodological approaches"><title>4.1. Methodological approaches</title><p>Methods influence the way we approach, perceive, and understand the world. All methods have strengths and weaknesses, which make certain things to be easily noticed while others remain hidden or un-emphasised – and such effects of the methods on knowledge often go unnoticed or are unappreciated by researchers. Carabid research has long been dominated by observation and description, but there still remains much to be observed and described about carabids. However, the prevalence of certain methods in carabid research (e.g. pitfall trapping as a collection method, see below) has put a strong stamp on the amount and structure of our knowledge about carabids. Some of the resulting biases are mentioned below; this list is illustrative, not exhaustive.</p><p><italic>Prevalence of knowledge about adults:</italic> Due to the epigaeic activity of the adults, and the fact that they are more easily collected, manipulated and kept in the laboratory, there is an overwhelming disparity about our knowledge on the ecology of the different life stages of carabid beetles. Our knowledge on carabids was (<xref ref-type="bibr" rid="B334">Lövei and Sunderland 1996</xref>) and remains primarily determined by knowledge about adults. A search on Web of Science with the term “carabid* OR ground beetl*” between 2000–2009 yielded 3186 papers, only 460 remaining when this was combined with the term “larv*” (search made by G Lövei, on 4 February 2011).</p><p><italic>Geographical unevenness in the origin of our knowledge:</italic> This is a general phenomenon: we know that the tropics is more species rich, in general, than the temperate region (already mentioned by <xref ref-type="bibr" rid="B109">Darwin 1859</xref>), yet most of our research effort is still directed towards temperate ground beetles. Of the above computer search on ground beetles, only 80 of the original 3186 papers remained when the additional term “tropic*” was introduced. We can safely predict important new understanding emerging from more detailed studies performed in more southerly regions; many of the techniques formerly restricted to developed countries can now be usefully employed in more tropical areas.</p><p><italic>Biased perception of carabids as predators:</italic> Predators and predation keep us fascinated, possibly because early humans have been both hunters and hunted. However, this colours our perception of the world (see <italic>Carabid beetle food</italic> above). In the case of carabids, the fact that many species will attack prey offered to them, especially in the laboratory, and that many beetles are indeed fast-moving predators, has led to a widely-held belief that carabids are predators. Carabidologists (mostly) know better, but we have been a bit lax to actively dispel this notion among ecologists, natural historians, and the general public. In relatively recent literature, one still comes across this perception (<xref ref-type="bibr" rid="B72">Braun et al. 2004</xref>), and in some cases, elaborate theories are built on such shaky grounds (<xref ref-type="bibr" rid="B332">Lövei and Magura 2006</xref>).</p><p><italic>The rarity of testable hypotheses</italic>: Due to a history of descriptive studies, there seems to be a general rarity of precisely formulated, testable hypotheses. Many studies have<pmc-comment>PageBreak</pmc-comment> the only justification that “we do not yet know, so let’s find out”. With increasingly fierce competition for funding and publication, such arguments do not carry much weight. An additional advantage of formulating hypotheses is that it forces us to think ahead: what is to be expected? Why? However, hypotheses should be well formulated (see <xref ref-type="bibr" rid="B191">Ford 2009</xref>; <xref ref-type="bibr" rid="B562">Underwood 2009</xref>). In the literature (not only in carabidology) one often encounters the “null hypothesis” formulated as “we expect no differences will be found”. Do researchers really expect that “nothing will happen”? If so, why is the experiment worth performing? Indeed, in the real world the null hypothesis is rarely if ever true as there will always be differences between effects. What is of importance is the magnitude, i.e. effect size, and precision, i.e. confidence interval of the effect (<xref ref-type="bibr" rid="B396">Nakagawa and Cuthill 2007</xref>; <xref ref-type="bibr" rid="B304">Läärä 2009</xref>). The careful separation of hypothesis formulation vs. the Popperian way of arriving at scientific evidence should not be confused – but often is.</p><p>The overall task is unchanged: to understand what made carabids such an evolutionarily successful group. In order to answer this question, one has to quantitatively continue to document the patterns of occurrence of members of this group – this is a logistical, not a methodological challenge. Among the promising “methodological approaches”, modern population genetical toolkits are well used, with several interesting results – it would be good to take these and use them in extra-European habitats as well. Gene expression study methods have recently developed and simplified considerably (<xref ref-type="bibr" rid="B430">Ouborg and Vriezen 2007</xref>), and facilitate the study of some interesting ecological questions, such as reaction to such factors as stress and food selection. Modern methods, such as those of ecological immunity, also allow a more refined characterisation of ground beetle reactions to habitat quality.</p></sec><sec sec-type="4.2. Analysing pitfall-trapped carabid data"><title>4.2. Analysing pitfall-trapped carabid data</title><p>Pitfall trapping is the best-known collection method used by carabidologists, especially in ecological studies (<xref ref-type="bibr" rid="B334">Lövei and Sunderland 1996</xref>). The method, originally described nearly 80 years ago (<xref ref-type="bibr" rid="B43">Barber 1931</xref>) and later often referred to as Barber traps (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>), is cheap, easy to use and once set up, operates by itself. It allows for adequate replication in field-based studies, and collects large samples (see <xref ref-type="fig" rid="F4">Fig. 2</xref> for examples of a few commonly used pitfall traps).</p><fig id="F4" orientation="portrait" position="float"><label>Figure 2.</label><caption><p>Different pitfall types. <bold>A</bold> = Jar or yoghurt can. <bold>B</bold> and <bold>C</bold> = traps with an outer can to make collecting of the sample easier. <bold>B</bold> = funnel trap with small jar. <bold>C</bold> = trap for moist biotopes (the outer can contains gravel or stones to prevent the can from being pushed up by groundwater). <bold>V</bold> = preservative (usually formaldehyde 3–4% or propylene glycol), <bold>S</bold> = stones or gravel.</p></caption><graphic xlink:href="ZooKeys-100-055-g004"></graphic></fig><p>One of the most convenient features of pitfall trapping is also its main disadvantage, because the resulting catch, although beguilingly countable, is not a measure of density, but of activity density. Carabidologists have recognised this and other drawbacks of pitfall trapping, which have often been discussed in the literature from <xref ref-type="bibr" rid="B215">Greenslade (1964)</xref>, <xref ref-type="bibr" rid="B551">Thiele (1977)</xref> and <xref ref-type="bibr" rid="B334">Lövei and Sunderland (1996)</xref> to Holland (2002) and regularly at ECMs. However, the method has not been subject to rigorous, thorough testing, nor to a systematic review, and consequently, most carabidologists tip their hat at the problem, then proceed to ignore it, and often use sophisticated evaluation methods to answer important research questions. Need<pmc-comment>PageBreak</pmc-comment>less to say that if these drawbacks in pitfall-trapped samples remain unresolved, this brings into question any analysis using assemblage data, such as ordination techniques, diversity indices, the determination of dominance structure and any ecological analysis or testing of theory.</p><p>While the sharpening of research questions before starting trapping is a salutary piece of advice, which will also influence the type and arrangement of traps, some problems associated with pitfall traps for general carabid beetle studies have reached a general consensus. Several of the aspects below are, however, still ignored but could be easily fixed. These include that (i) an odourless preservative is preferred, because formalin, for example, seems to attract some species and repel others (<xref ref-type="bibr" rid="B551">Thiele 1977</xref>); (ii) the traps should have a cover to prevent flooding, desiccation, scavenging and bycatch – a funnel to prevent escape and reduce bycatch also helps (<xref ref-type="bibr" rid="B306">Lange et al. 2011</xref>); (iii) traps should preferably not be used solitarily, but placed in series of at least three to five traps at distances of less than 10 m apart in order to optimise the catch and to overcome occasional trap losses; (iv) distances between sampling plots (single traps or trap groups) should be large enough to allow for sample independence (this distance will, of course, depend on the dispersal power of the focal species, see e.g. <xref ref-type="bibr" rid="B151">Digweed et al. 1995</xref>); and (v) the question of missing samples that inevitably occur when large numbers of traps are used over long time periods (see below). Important challenges that await study and resolution are: (i) that trap numbers and length of the trapping period do not contribute equally to the catch (<xref ref-type="bibr" rid="B333">Lövei and Magura 2011</xref>); (ii) how to reliably minimise the impact of trapping on assemblages and protected species (the methods of partial seasonal samples and pulsating samples, for example, have been suggested: Sapia et al. 2005);<pmc-comment>PageBreak</pmc-comment> and (iii) the challenge of non-destructive carabid sampling (<xref ref-type="bibr" rid="B61">Bowie and Frampton 2004</xref>), such as radiotelemetry (see <xref ref-type="bibr" rid="B398">Negro et al. 2008</xref>).</p><p>The arrangement of pitfall traps in the field depends on the research question asked. The most popular research questions include: (i) <italic>Faunistic investigations</italic> intended to obtain an accurate species list of a given area. Here many pitfall traps should be used, also along gradients and at biotope edges; it seems that the spatial aspect is more important than the temporal one, i.e. it is better to have many traps for shorter periods of time than fewer traps for longer time periods (<xref ref-type="bibr" rid="B333">Lövei and Magura 2011</xref>). (ii) <italic>Community or gradient studies</italic> intended to investigate the (typical) fauna of different biotopes or at different positions along a gradient. In this case series of traps per biotope or gradient position can be used (<xref ref-type="fig" rid="F5">Fig. 3a</xref>) with independent replicates (with sufficient distances between the series, see above). An example of this is the Globenet project (<xref ref-type="bibr" rid="B416">Niemelä et al. 2002</xref>). In some cases a row design with repeats will generate more precise information (<xref ref-type="fig" rid="F5">Fig. 3b</xref>), especially when short-term movements of species along gradients are expected. The same holds for different treatments in an experimental design, such as (iii) <italic>Biological studies</italic> investigating e.g. the periodicity of one or more species within a year, to be eventually compared with different biotopes or years in phenological and/or climate studies (e.g. do species reproduce earlier or later during warmer periods or in different biotopes?). In the case of (iv) <italic>Biological studies</italic> investigating diurnal rhythms or movements of adults and larvae, a grid or matrix design (<xref ref-type="fig" rid="F5">Fig. 3c</xref>) is recommended; and (v) <italic>Population studies</italic> intended to investigate the response of populations to biotic and abiotic environmental factors. Here, estimates of population densities are required and, as such, pitfall-trapped data need to be interpreted with caution.</p><fig id="F5" orientation="portrait" position="float"><label>Figure 3.</label><caption><p>Examples of pitfall trap placements across a forest edge.</p></caption><graphic xlink:href="ZooKeys-100-055-g005"></graphic></fig><p>The fact that pitfall catches are a function of the species’ true population size and its activity (activity-density: <xref ref-type="bibr" rid="B215">Greenslade 1964</xref>; <xref ref-type="bibr" rid="B553">Thomas et al. 1998</xref>), creates specific statistical problems. Continuous sampling over the whole activity period can cause a potential<pmc-comment>PageBreak</pmc-comment>ly serious problem when the catch is analysed. Trap losses can occur at any time during the activity period and have traditionally been dealt with by standardising the catch to 100 trapping days without taking into account variability in activity across the season (<xref ref-type="bibr" rid="B296">Kotze and Niemelä 2002</xref>; <xref ref-type="bibr" rid="B416">Niemelä et al. 2002</xref>). For example, some species are more active in the spring or autumn (see <italic>Life history strategies and rhythms</italic> above), while others are active throughout the summer months. As such, a trap lost at the beginning of the continuous sampling period will have a different effect on the estimated activity-density of a spring-active species, for example, than if the trap is lost at a later stage when activity is low. When the research question involves study of the response of separate species to an environmental gradient, statistical models in which seasonality (or visit) is added as a free factor and sampling effort (number of trapping days per visit) as an offset term, and in which the response variable is specified as following a negative binomial distribution, seem to correct for seasonality and trap losses appropriately.</p><p>The reason for specifying activity-density (or abundance) data as following a negative binomial distribution (and not a Gaussian distribution, as is often done) is that ecological field data (here counts of individuals or species) seldom follow the assumptions of classical parametric statistics (<xref ref-type="bibr" rid="B104">Dalthorp 2004</xref>). Carabid beetles (both in terms of abundance and species) are often aggregated in space (<xref ref-type="bibr" rid="B412">Niemelä et al. 1986</xref>, <xref ref-type="bibr" rid="B410">1992</xref>; <xref ref-type="bibr" rid="B553">Thomas et al. 1998</xref>) and sampling them is likely to produce an expected variance that is greater than the expected mean. Such ‘clumped’ counts data appear to be most appropriately analysed by models that incorporate extra variation, such as the negative binomial distribution (see <xref ref-type="bibr" rid="B595">White and Bennetts 1996</xref>; <xref ref-type="bibr" rid="B104">Dalthorp 2004</xref>), or quasi-Poisson methods (<xref ref-type="bibr" rid="B571">Ver Hoef and Boveng 2007</xref>; e.g. <xref ref-type="bibr" rid="B167">Elek et al. 2010</xref>). Another important advantage of using methods designed for dealing with count data (negative binomial, Poisson) is that the response variable (number of individuals or species) does not need to be transformed to comply with the assumptions of parametric test statistics, such as analysis of variance, t-test or linear regression. Surprisingly, abundance and species richness data are often log-transformed for subsequent use in parametric test procedures, even though textbooks on statistical methods in ecology (<xref ref-type="bibr" rid="B525">Sokal and Rohlf 1995</xref>; <xref ref-type="bibr" rid="B100">Crawley 2003</xref>) recommend the use of the square-root transformation to normalise count data. Nevertheless, neither square root nor log-transformed count data (for use in parametric tests) performed as well as <italic>non</italic>-transformed data (for use in a negative binomial model) (<xref ref-type="bibr" rid="B428">O’Hara and Kotze 2010</xref>). A possible reason for this is that count data often contain many zero values, which have to be fudged (when a log transformation is performed) by adding 0.1 or 1 to every observation – which may have unforeseen effects on estimates.</p><p>Another problem occurs when the activity density results for different species are compared. Since each species reacts differently to pitfall traps, their “catchability” will also differ, subsequently with more or less incomparable results between species. A possible solution, suggested by Den Boer, is to standardise the catches per species over the sampling sites (<xref ref-type="bibr" rid="B559">Turin et al. 1991</xref>). After standardisation, with the obtained “relative abundances”, multivariate methods (calculating (dis)similarities, clustering and ordination) can be used to analyse the data. Similar classifications have been carried out for Britain (<xref ref-type="bibr" rid="B340">Luff et al. 1989</xref>; <xref ref-type="bibr" rid="B183">Eyre and Luff, 1990</xref>; <xref ref-type="bibr" rid="B373">Mccracken 1994</xref>; <xref ref-type="bibr" rid="B6">Anderson et al. 2000</xref>). Although the approach of correcting and standardising the data was quite different from the Dutch method, the results for classification of the carabid habitats in the Netherlands and Britain were very similar. A study of the carabid fauna of Trento, Italy (<xref ref-type="bibr" rid="B56">Bonavita and Chemini 1996</xref>) in a deviating trans-alpine fauna, revealed highly corresponding results for the classification of the 48 (out of 57) species common to Italy and northern Europe. A relatively simple and flexible method developed by <xref ref-type="bibr" rid="B159">Dufrêne and Legendre (1997)</xref> to classify a Belgian dataset (the IndVal procedure), has the advantage that it is insensitive to the relative abundances of species. We contend that the problems associated with the comparison of assemblages sampled by pitfall trapping are still not fully resolved, but the above confirm that this method has merit in many types of investigations.</p></sec></sec><sec><title>5 Population dynamics and long-term research</title><p>Since the 1960s the population dynamics of carabid beetles has been subject to the study of population persistence. During this time, discussion has revolved around how the size of populations and their fluctuations have been established, resulting in two popular theories. The first theory postulates that population sizes are balanced within narrow limits by density dependent processes, a feedback mechanism in which predators, parasites, competitors for food and other biotic aspects of the environment are involved, resulting in the regulation of population size (see <xref ref-type="bibr" rid="B404">Nicholson 1958</xref>). The second theory argues that the founding and re-founding of local populations take place, driven by dispersal, small population size and extinction, heterogeneity of the environment, the distinction between local (sub) and natural (entire) populations, and the genetic plasticity of species in relation to different components of the environment and to fluctuations of population size (<xref ref-type="bibr" rid="B8">Andrewartha and Birch 1954</xref>). Den Boer tested the latter theory by using carabid beetles as a model group. In 1959 he started pitfall trapping at several locations in the Dwingelderveld, a large area of heathland in the Netherlands, which he regarded as home to large natural populations of several carabid species. This founding/re-founding theory, the concept of metapopulation, states that natural populations consist of many local populations or colonies. Indeed, Den Boer was able to show that in a large area many local populations or interacting groups of carabids fluctuated in numbers of individuals in space and time. From these results the “spreading of risk” theory was derived (<xref ref-type="bibr" rid="B110">Den Boer 1968</xref>, see <italic>European Carabidologists’ Meetings (ECMs)</italic> above).</p><p>The significance of dispersal in founding, re-founding and establishment of populations was confirmed during the first ECM (see Introduction). However, the role of density dependent processes was not resolved. In 1970 in Oosterbeek, the Netherlands, an entire symposium on the <italic>Dynamics of Populations</italic> (<xref ref-type="bibr" rid="B118">Den Boer and Gradwell 1971</xref>) was devoted to whether or not populations were regulated. Some contributors showed examples in which density-dependent processes seemed to govern the abundance of a species, whereas others showed the opposite, so the discus<pmc-comment>PageBreak</pmc-comment>sion continued. Later on, again using carabid beetles, several studies were conducted to test the density dependence hypothesis. For instance <xref ref-type="bibr" rid="B32">Baars and Van Dijk (1984)</xref> were able to show that the number of eggs in the ovaries of females was negatively correlated with the mean density around pitfall traps. However, later on <xref ref-type="bibr" rid="B565">Van Dijk and Den Boer (1992)</xref> demonstrated that egg and larval mortality were too high to compensate for egg production. It was concluded that the density dependent relationship could hardly play an important role in the dynamics of the populations of <italic><named-content content-type="taxon-name">Calathus melanocephalus</named-content></italic>, as shown by <xref ref-type="bibr" rid="B32">Baars and Van Dijk (1984)</xref>. In <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic> the amount of food available affects the number of eggs laid. <xref ref-type="bibr" rid="B230">Heessen (1981)</xref> suggested that this would regulate population dynamics of this species. However, <xref ref-type="bibr" rid="B532">Szyszko (1981)</xref> and <xref ref-type="bibr" rid="B116">Den Boer (1986)</xref> observed that population explosions of certain prey species lead to a strong decline in some carabids. <xref ref-type="bibr" rid="B580">Vermeulen and Szyszko (1992)</xref> were able to show that in order to maintain a high level of egg production, <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic> has to switch prey. Presumably the right mixture of amino acids and the quality of nitrogen (<xref ref-type="bibr" rid="B596">White 1993</xref>) are essential for a high level of egg production. Another study on regulation in carabid populations was carried out by <xref ref-type="bibr" rid="B78">Brunsting et al. (1986)</xref>. They showed that cannibalism occurs between larvae of <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic> and suggested that this phenomenon would regulate population size. However, <xref ref-type="bibr" rid="B572">Vermeulen (1986)</xref> could not find differences in raising <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic> under circumstances in which cannibalism was included and excluded, suggesting that larvae may not be actively searching for other larvae of the same species to feed upon. Cannibalism might take place only under extremely high, unnatural densities. Also, the role of competition in the population dynamics of carabid beetles has not been convincingly demonstrated so far. For example, <xref ref-type="bibr" rid="B328">Loreau (1990)</xref> found only weak evidence for competitive regulation in <italic><named-content content-type="taxon-name">Abax ater</named-content></italic> populations. He suggested that competition might only be significant in dominant species. On the other hand, (<xref ref-type="bibr" rid="B114">Den Boer (1980</xref>, <xref ref-type="bibr" rid="B115">1985</xref>) and <xref ref-type="bibr" rid="B406">Niemelä (1993)</xref> showed that competition hardly plays any role in determining population size. The discussion on whether or not regulation plays an important role in population dynamics led to a second symposium on population dynamics, this time held in Poland in 1992 (<xref ref-type="bibr" rid="B122">Den Boer et al. 1993</xref>). However, again only a discussion for and against regulation resulted. After this meeting the subject quickly went out of fashion and was not discussed in this way again. In 1996, Den Boer and Reddingius wrote a book in which they reviewed all the population dynamic theories so far.</p><p>At present, it is generally accepted that the persistence of carabid populations depends on the availability of sufficient suitable habitat over long periods of time, as well as on habitat quality. The latter was nicely illustrated in the Dwingelderveld, the Netherlands. These heathlands have nitrified from the 1970s onwards, associated with an almost complete disappearance of <italic><named-content content-type="taxon-name">Carabus nitens</named-content></italic> there. A few years after the removal of the nutrient-rich topsoil layer by sod cutting, however, this species was again recorded in high numbers (<xref ref-type="bibr" rid="B566">Van Essen 1993</xref>). A similar recovery is now seen in the Mantingerveld, the Netherlands, for the same species since 2007 (Rikjan Vermeulen, pers. obs.). Because of the turnover in local populations, dispersal is necessary for a given<pmc-comment>PageBreak</pmc-comment> species to (re)colonise areas where habitat patches are small. The classical technique for investigating these processes has been mark and recapture. Using modern techniques, e.g. simulation programmes (<xref ref-type="bibr" rid="B458">Persigehl et al. 2004</xref>) and genetic techniques (e.g. <xref ref-type="bibr" rid="B153">Drees et al. 2011</xref>), the relationships between populations can be demonstrated more easily.</p><p>Permanently set-up pitfall traps give an impression of the activity of different species during different seasons and between years, and produce relative estimates of population fluctuations for a particular species. In 1959, several series of permanent pitfall traps were initiated in the Dwingelderveld and later, in 1963, in the Mantingerveld (<xref ref-type="bibr" rid="B125">Den Boer and Van Dijk 1994</xref>). Carabid beetles from these series were collected on a weekly basis. Results from the first 6–7 years showed considerable fluctuations in the total number of individuals of a particular species collected per series between successive years. This fluctuating pattern was also different between each separate catching series within an area in the same year. These observations of asynchronous fluctuations in catches of a particular species were instrumental in the development of the “spreading of risk” theory by Piet den Boer (see above). Environmental conditions since the establishment of these series also changed. At the end of the 1960s the ground-water table gradually receded and during the 1970s the effects of air pollution became apparent: increasing acidification and eutrophication of the upper soil layers and the subsequent replacement of both <italic><named-content content-type="taxon-name">Calluna</named-content></italic> and <italic><named-content content-type="taxon-name">Erica</named-content></italic> by grasses. At the end of the 1980s, the local nature management authority started to artificially raise the water table, which subsequently reached its pre-1960s level during 2010–2011. At the same time the grassy vegetation, together with the polluted top soil layer, was removed by sod cutting, and grazing by cows and sheep has subsequently been introduced. Moreover, the average temperature of the area had increased by 1 oC in the last few decades. Both the increase in temperature and the hours of sunshine appear to be significant from 1988 onwards (<xref ref-type="bibr" rid="B467">Prins et al. 2007</xref>).</p><p>Since the establishment of these series of pitfall traps, the composition of the carabid beetle fauna has changed continuously. In the beginning of the 1970s species such as <italic><named-content content-type="taxon-name">Agonum krynickii</named-content></italic>, <italic><named-content content-type="taxon-name">Carabus cancellatus</named-content></italic>, <italic><named-content content-type="taxon-name">Cicindela sylvatica</named-content></italic> and <italic><named-content content-type="taxon-name">Cicindela germanica</named-content></italic> disappeared completely from the catches, followed by <italic><named-content content-type="taxon-name">Amara quenseli</named-content></italic> and <italic><named-content content-type="taxon-name">Amara praetermissa</named-content></italic>. During the same period, species such as <italic><named-content content-type="taxon-name">Carabus nitens</named-content></italic>, <italic><named-content content-type="taxon-name">Harpalus solitaris</named-content></italic> and <italic><named-content content-type="taxon-name">Amara infima</named-content></italic> decreased significantly in numbers. The climate did not change significantly during this period, and it can be speculated that changes in the environment, as mentioned above, and habitat fragmentation (in the case of Hullenzand, Mantingerveld) may be responsible for these local extinctions and changes in population numbers. From the end of the 1990s, species such as <italic><named-content content-type="taxon-name">Agonum ericeti</named-content></italic>, <italic><named-content content-type="taxon-name">Cymindis vaporariorum</named-content></italic> and <italic><named-content content-type="taxon-name">Cymindis macularis</named-content></italic> disappeared from the catches. This may be a consequence of climate change, since during this period environmental conditions in the heathlands improved. This is well illustrated for <italic><named-content content-type="taxon-name">Carabus nitens</named-content></italic>, which became rather abundant during this period, as well as for <italic><named-content content-type="taxon-name">Carabus arvensis</named-content></italic>, <italic><named-content content-type="taxon-name">Nebria salina</named-content></italic> and <italic><named-content content-type="taxon-name">Harpalus solitaris</named-content></italic>. From 1990 to<xref ref-type="bibr" rid="B205"> 2004</xref>, ten species not previously recorded from these areas have been collected (<xref ref-type="bibr" rid="B579">Vermeulen et al. 2004</xref>). Recently, two records of <italic><named-content content-type="taxon-name">Agonum viridicupreum</named-content></italic> can be added to this list (Rikjan Vermeulen pers. obs.). Apart from one, all of these newly recorded species have their <pmc-comment>PageBreak</pmc-comment>center of distribution south of the Netherlands, suggesting that their appearance is related to climate warming. Similarly, the virtual disappearance of the northern species, <italic><named-content content-type="taxon-name">Agonum ericeti</named-content></italic>, may be related to this phenomenon. Adequate management may, to a limited extent, compensate for the effects of climate change. The northerly distributed <italic><named-content content-type="taxon-name">Carabus nitens</named-content></italic> that almost disappeared from both the Mantinger- and Dwingelderveld, made a rapid comeback after top-soil removal and sod-cutting.</p><p>However, the dramatic decline and extinction of the highly hygrophylic <italic><named-content content-type="taxon-name">Carabus clatratus</named-content></italic> in Italy may not be entirely related to climate change. <italic><named-content content-type="taxon-name">Carabus clatratus</named-content></italic> is one of the most localised and endangered carabid species in Europe, and its disappearance from Italy, and possibly also France, is possibly a consequence of the colonisation of its wet biotopes by the alien red swamp crayfish, <italic><named-content content-type="taxon-name">Procambarus clarkii</named-content></italic>, which preys on adults of <italic><named-content content-type="taxon-name">Carabus clatratus</named-content></italic> (<xref ref-type="bibr" rid="B91">Casale and Busato 2008</xref>).</p><p>Long-term data on weekly catches can also be used to monitor phenological changes in species. For example, compared to the period prior to 1988, the activity of <italic><named-content content-type="taxon-name">Amara equestris</named-content></italic>, <italic><named-content content-type="taxon-name">Carabus arvensis</named-content></italic>, <italic><named-content content-type="taxon-name">Poecilus lepidus</named-content></italic> and <italic><named-content content-type="taxon-name">Poecilus versicolor</named-content></italic> started earlier in the season.</p><p>As far as the consequences of climate change, management and other environmental changes are concerned, it is of great importance to continue long-term observational studies of carabid beetles, such as that in Drenthe, so that future changes can be monitored and possibly explained. Such long-term sampling programmes for carabid beetles are also known from Poland, Germany and Italy.</p></sec><sec><title>6 Bioindicators</title><p>Carabids are excellent model organisms for research on ecological and conservation theory. These beetles readily respond to abiotic and biotic variation, and to disturbances and management (e.g. <xref ref-type="bibr" rid="B334">Lövei and Sunderland 1996</xref>; <xref ref-type="bibr" rid="B473">Rainio and Niemelä 2003</xref>). This evidence has led many to suggest carabids to function as ‘indicators’. An indicator is a taxon or a structure “<italic>whose characteristics</italic> (...) <italic>are used as an index of attributes too difficult, inconvenient, or expensive to measure for other species or environmental conditions of interest</italic>” (<xref ref-type="bibr" rid="B305">Landres et al. 1988</xref>). However, using this definition many, if not most, of carabid ‘indicator’ studies appear to only demonstrate individualistic responses to environmental variation. But instead of investing resources for finding new indicator taxa, environmental managers should test and select taxa that are already well known and easily sampled, and that cover multiple dimensions of biodiversity (Taylor and Doran<xref ref-type="bibr" rid="B204"> 2001</xref>), and critically evaluate their indicator functioning (<xref ref-type="bibr" rid="B307">Langor and Spence 2006</xref>). Carabids fulfil the former but the latter aspect requires further attention.</p><p>European carabids have certain qualities that make them good candidates for indicators. They are taxonomically well known, with relatively stable systematics, and their ecology has been widely studied (<xref ref-type="bibr" rid="B334">Lövei and Sunderland 1996</xref>). Variation in carabid morphology, life history strategies and small-scale abiotic and biotic requirements are extensively documented (e.g. <xref ref-type="bibr" rid="B324">Lindroth 1961–1969</xref>, 1985, 1986). Carabids also respond predictably to not only small-scale but also to landscape- and even continent-level phenomena (e.g.<pmc-comment>PageBreak</pmc-comment> <xref ref-type="bibr" rid="B236">Hengeveld 1987</xref>; <xref ref-type="bibr" rid="B298">Kotze and O’Hara 2003</xref>; <xref ref-type="bibr" rid="B292">Koivula and Spence 2006</xref>). Moreover, they are relatively easy to collect in high numbers using standard methods. But can carabids reflect environmental variation in ways useful for conservation assessment purposes? Knowledge of carabid indicator functioning, using the categories listed in <xref ref-type="bibr" rid="B319">Lindenmayer et al. (2000)</xref>, is briefly summarised below (see <xref ref-type="bibr" rid="B288">Koivula 2011</xref> for a complete evaluation).</p><p><bold>i. Taxon indicators.</bold> The presence of a taxon indicator reflects the presence of a set of other species, and its absence indicates the lack of the entire set of species. Perfect multi-taxon richness overlaps may be rare (e.g. Jonsson and Jonsell<xref ref-type="bibr" rid="B203"> 1999</xref>; <xref ref-type="bibr" rid="B487">Sætersdal et al. 2005</xref>; <xref ref-type="bibr" rid="B521">Similä et al. 2006</xref>), which highlights the importance of using multiple taxa in environmental assessments (Taylor and Doran<xref ref-type="bibr" rid="B204"> 2001</xref>; <xref ref-type="bibr" rid="B158">Duelli and Obrist 2003</xref>). Carabid functioning as taxon indicators mostly relies on weak correlations among taxa.</p><p><bold>ii. Keystone indicators.</bold> These species affect their environment disproportionately strongly relative to their abundance. In field and laboratory conditions, carabids forage on slugs and pest insects (e.g. Kromp<xref ref-type="bibr" rid="B203"> 1999</xref>). <xref ref-type="bibr" rid="B226">Hance (1987)</xref> showed that, using enclosures with different carabid densities, carabids have the potential to significantly prey on pest insects foraging on crop plants with economic benefits.</p><p><bold>iii. Pollution indicators.</bold> These taxa reflect human-altered abiotic conditions. Heavy metals in the soil negatively affect carabids (e.g. <xref ref-type="bibr" rid="B364">Maryański et al. 2002</xref>; <xref ref-type="bibr" rid="B172">Ermakov 2004</xref>), and in agro-ecosystems, pesticides and fertilizers affect carabids, at least in the short term (e.g. <xref ref-type="bibr" rid="B251">Huusela-Veistola 1996</xref>; Kromp<xref ref-type="bibr" rid="B203"> 1999</xref>).</p><p><bold>iv. Dominant indicators.</bold> These taxa make up much of the total biomass or the number of individuals in an area of interest and predict particular ecosystems or assemblages. Many common carabid species are succession and habitat-type generalists (Lindroth 1985, 1986; <xref ref-type="bibr" rid="B413">Niemelä et al. 2007</xref>), so their numbers may not indicate aspects useful for conservation or management. Mean Individual Biomass (MIB), on the other hand, links carabid biomass to succession without considering species entities (<xref ref-type="bibr" rid="B541">Szyszko et al. 2000</xref>). However, the ‘behaviour’ of MIB along succession should be examined in detail before applying it in conservation and management.</p><p><bold>v. Environmental indicators.</bold> These should reliably reflect particular environmental conditions. Although carabids have the potential to reflect soils, wetness and habitat-type variation (e.g. <xref ref-type="bibr" rid="B551">Thiele 1977</xref>; Lindroth 1985, 1986), they cannot currently compete with plants as indicators of these factors.</p><p><bold>vi. Early-warning signallers (true bio-indicators).</bold> These taxa are extremely sensitive to changing environmental conditions. Carabid evidence is scarce, but some carabids have apparently undergone shifts of tens of metres in altitude over 10–20 years (<xref ref-type="bibr" rid="B23">Assmann 2009</xref>; <xref ref-type="bibr" rid="B463">Pizzolotto 2009</xref>, David Kavanaugh, pers. comm.), coinciding with climate warming (<xref ref-type="bibr" rid="B452">Parry et al. 2007</xref>, see <italic>Population dynamics and long-term research</italic> <pmc-comment>PageBreak</pmc-comment>above). These observations suggest good potential in, for example, climate-change and urban-spread research.</p><p><bold>vii. Disturbance indicators.</bold> These taxa reflect natural and human-caused disturbances. Carabids readily respond to agriculture and forestry (for reviews, see <xref ref-type="bibr" rid="B334">Lövei and Sunderland 1996</xref>; Kromp<xref ref-type="bibr" rid="B203"> 1999</xref>; <xref ref-type="bibr" rid="B413">Niemelä et al. 2007</xref>). Their indicator functioning may hold at a general level: they respond similarly to environmental change as many other taxa do (e.g. <xref ref-type="bibr" rid="B40">Barbaro et al. 2005</xref>). But indicators should not be used for self-evident patterns: the ecological impact of clear-cutting, for example, does not require an indicator.</p><p>Clearly, carabids have good potential for becoming useful indicators for conservationists and environmental managers. Certain obstacles still need to be overcome. First, the functioning and accuracy of carabids to predict habitats or species requiring conservation action should be critically evaluated. According to the indicator definition of <xref ref-type="bibr" rid="B305">Landres et al. (1988)</xref>, none of the above examples indicate that carabids function as particularly useful indicators. Thus, for a conservationist, carabid responses should be considered as individualistic as long as there is no evidence for their responses to reliably predict responses of threatened taxa or particular, difficult-to-observe conditions. This is important because there is very little room for error if threatened species or habitats are at stake. Strict tests must thus be applied to evaluate indicator functioning (<xref ref-type="bibr" rid="B307">Langor and Spence 2006</xref>). Second, the relationship between carabid responses and other taxa should be considerably clarified (<xref ref-type="bibr" rid="B473">Rainio and Niemelä 2003</xref>) before using these beetles in environmental assessments. Third, it is unclear whether carabids reflect aspects not attainable using other indicators (apart from their individualistic response) and whether conditions exist under which carabids really are the most cost-efficient indicator taxon. Currently widely used, easy-to-use, relatively cheap and economic tools for assessing the state of the environment include vegetation, habitat structural elements, satellite and aerial photos, as well as weather and land-use inventory data.</p><p>The focus of carabidologists should perhaps be changed from total species richness to the indicator potential of single species, groups of specialists or functional groups. We lack an explicitly defined ‘niche’ of these beetles in environmental assessment protocols. Cases for carabids fulfilling the conservationists’ definition for a useful indicator (<xref ref-type="bibr" rid="B305">Landres et al. 1988</xref>) will possibly be documented in the near future, but their indicator functioning may always remain context specific.</p></sec><sec><title>7 Carabid conservation, protection and habitat management</title><p>Conservation may mean protecting particular species or patches of habitat against alteration, generally human-caused, but the term may also include operations characterised by an active human role (e.g. <xref ref-type="bibr" rid="B200">Freitag and Kavanaugh 1993</xref>; <xref ref-type="bibr" rid="B125">Den Boer and Van Dijk 1994</xref>; Sutherland 1998; Gaston and Spicer<xref ref-type="bibr" rid="B205"> 2004</xref>). Examples include the<pmc-comment>PageBreak</pmc-comment> maintenance of areas of high natural value, the restoration of patches to a state they are presumed to once have represented (often referred to as ‘natural’ state), and the artificial conversion of one habitat type to another. The latter may be required in landscapes where habitat for a threatened species has become rare (see <xref ref-type="bibr" rid="B398">Negro et al. 2008</xref>) and new habitat patches are unlikely to appear through natural processes. Such cases might be found, for example, within urban areas. These active operations of patch maintenance, restoration and creation are collectively called ‘conservation management’.</p><p>Insect conservation management is a relatively new research discipline, both generally and in the context of carabid beetles (e.g. <xref ref-type="bibr" rid="B316">Lewis et al. 2007</xref>; <xref ref-type="bibr" rid="B313">Leather et al. 2008</xref>; <xref ref-type="bibr" rid="B403">New 2010</xref>). The restoration and artificial creation of habitats − two elements of conservation management − have been important components of carabid conservation since the 1980s (e.g. <xref ref-type="bibr" rid="B554">Thomas 1990</xref>; <xref ref-type="bibr" rid="B198">Främbs 1990</xref>; <xref ref-type="bibr" rid="B52">Blake et al. 1996</xref>). Conservation became an important topic for the ECMs since the Hungarian meeting in 1986. Before that meeting, conservation issues were only occasionally discussed, but from then on, both conservation in general (e.g. identifying diversity hotspots and gathering data on endemic and rare species) and practical conservation management in particular have been among key topics and have altogether consistently made up over 20% of papers in the proceedings. Generally, almost any piece of knowledge on carabid ecology can be applied in conservation-management policy and action to support these beetles and associated epigaeic fauna. In Europe and North America, information necessary for efficient conservation − on carabid ecology and threats − is readily available (<xref ref-type="bibr" rid="B351">Maelfait et al. 1994</xref>; <xref ref-type="bibr" rid="B334">Lövei and Sunderland 1996</xref>; see also national lists of threatened species). However, the functioning of active management for the benefit of threatened carabid species urgently demands critical evaluation and detailed information. For instance, according to <xref ref-type="bibr" rid="B133">Desender et al. (2010)</xref>, the decline of carabid beetles in Belgium between the period <1950 and 1950–1985, had halted for a considerable number of species. During the period 1986–2008, however, 60% of these species still had not reached the same distribution area as in the first half of the 20th century, notwithstanding many initiatives and large scale active management. Most of these species now only occur in large and high-quality nature reserves with the last remnants of semi-natural biotopes and have, at present, little or no possibilities to further increase their distribution range.</p><p>Here the advances in conservation management, mostly as derived from the proceedings of the previous ECMs are discussed under four topics: (i) Which species characteristics are particularly associated with threatened species? (ii) In which habitat types can conservation of carabids best be realised? (iii) What do we know about habitat connectivity as a way to conserve carabids? (iv) How does conservation management of habitats affect carabids?</p><p><bold>i. Ecological and habitat characteristics of threatened species.</bold> To study which ecological and habitat characteristics of carabids are associated with species being threatened, national species lists and their IUCN categories for five countries are used as examples: Belgium, Sweden, Denmark, Norway and Finland (respectively <xref ref-type="bibr" rid="B134">Desender et al. 2008a</xref>, <xref ref-type="bibr" rid="B135">b</xref>; <xref ref-type="bibr" rid="B208">Gärdenfors 2005</xref>; <xref ref-type="bibr" rid="B455">Pedersen and Wind 2009</xref>; <xref ref-type="bibr" rid="B269">Kålås et al. 2006</xref>; <xref ref-type="bibr" rid="B476">Rassi et al. 2001</xref>). This dataset is complemented with four regional lists of threatened species from Niedersachsen and Bremen, Germany; Nordrhein-Westfalen, Germany; Wadden Sea area; and a preliminary red list for Drenthe, the Netherlands (respectively <xref ref-type="bibr" rid="B25">Assmann et al. 2002</xref>; <xref ref-type="bibr" rid="B508">Schüle and Terlutter 1998</xref>; <xref ref-type="bibr" rid="B356">Mahler et al. 1996</xref>; <xref ref-type="bibr" rid="B423">Noordijk and Vermeulen 2009</xref>). For analytical purposes, species characteristics were collected from Lindroth (1985, 1986), <xref ref-type="bibr" rid="B127">Desender (1986)</xref>, <xref ref-type="bibr" rid="B560">Turin and Den Boer (1988)</xref>, <xref ref-type="bibr" rid="B141">Desender and Turin (1989)</xref>, <xref ref-type="bibr" rid="B558">Turin (2000)</xref>, <xref ref-type="bibr" rid="B11">Anonymous (2006)</xref> and <xref ref-type="bibr" rid="B134">Desender et al. (2008a)</xref>. Several characteristics were evaluated, such as the roles of body size, wing morphology, and associations with shadiness and moisture. This evaluation was done by calculating percentages per size, wing morphology, shadiness and moisture classes for all species (for the five countries), for species classified as threatened by IUCN categories NT (Near Threatened), VU (VUlnerable), EN (ENdangered), CR (CRitically endangered) and EW (Extinct in the Wild; also RE, i.e. Regionally Extinct, in some national lists), and also the proportion of threatened species over all species within a given class. Occasionally, certain information for some species was lacking and these were (partly) removed from the analysis. For example, if for a certain species information was unavailable on wing morphology, it was omitted from the wing morphology analysis but retained in other analyses.</p><p>Carabids mostly fell into mid-size classes (43–46% of all species were 4.1–8.0 mm and 28–31% were 8.1–16.0 mm), were macropterous (64–71%) and were associated with open areas (63–64%), but were quite evenly distributed among moisture-association classes (see columns “All” in <xref ref-type="table" rid="T3">Table 3</xref>). Carabids classified as being threatened roughly complied with these figures (columns “IUCN” in <xref ref-type="table" rid="T3">Table 3</xref>): also these species were mostly mid-sized (26–50% were 4.1–8.0 mm and 24–39% were 8.1–16.0 mm), macropterous (64–81%) and open-area associated (63–79%; very shady habitats had only 2–12%). However, threatened species were more often associated with either very wet (34–53%) or very dry habitats (32–47%) than with “average” or moist/dryish conditions (12–30%). This dichotomous association with both very dry and very wet habitats was much more pronounced in the four Nordic countries and in the two areas in Germany than in Belgium or in Drenthe (<xref ref-type="table" rid="T3">Table 3</xref>).</p><table-wrap id="T3" orientation="portrait" position="float"><label>Table 3.</label><caption><p>Morphological and habitat-association characteristics of all carabid species found in a given area (“All” columns; % of species), of species classified as threatened according to the IUCN (“IUCN”; categories NT, EN, VU, CR and EW pooled; % of species), and proportion of threatened species of all species within a given category (“% IUCN”). For example, the value “50” for BEL % IUCN >16 mm indicates that in Belgium, of all species with body size >16 mm, 50% are considered threatened. Values for “All“ and “IUCN“ columns make up 100% for each area/country. BEL = Belgium; SWE = Sweden; DEN = Denmark; NOR = Norway; FIN = Finland; Niede = Niedersachsen and Bremen, Germany; Nordr = Nordrhein-Westfalen, Germany; Wadde = Wadden Sea area; and Drent = Drenthe, the Netherlands (proposed Red Data list). For the last four areas, only species classified as threatened according to the IUCN are shown. For species and their characteristics data, see text.</p></caption><table frame="hsides" rules="groups"><tbody><tr><th rowspan="2" colspan="1"><italic>Classes</italic></th><th rowspan="1" colspan="3"><italic>BEL</italic></th><th rowspan="1" colspan="3"><italic>SWE</italic></th><th rowspan="1" colspan="3"><italic>DEN</italic></th><th rowspan="3" colspan="1"></th></tr><tr><th rowspan="1" colspan="1"><italic>All</italic></th><th rowspan="1" colspan="1"><italic>IUCN</italic></th><th rowspan="1" colspan="1"><italic>% IUCN</italic></th><th rowspan="1" colspan="1"><italic>All</italic></th><th rowspan="1" colspan="1"><italic>IUCN</italic></th><th rowspan="1" colspan="1"><italic>% IUCN</italic></th><th rowspan="1" colspan="1"><italic>All</italic></th><th rowspan="1" colspan="1"><italic>IUCN</italic></th><th rowspan="1" colspan="1"><italic>% IUCN</italic></th></tr><tr><th rowspan="1" colspan="10"><italic>Body size</italic></th></tr><tr><td rowspan="1" colspan="1">0.1-4.0 mm</td><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1">13</td><td rowspan="16" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">4.1-8.0 mm</td><td rowspan="1" colspan="1">44</td><td rowspan="1" colspan="1">40</td><td rowspan="1" colspan="1">25</td><td rowspan="1" colspan="1">44</td><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">43</td><td rowspan="1" colspan="1">38</td><td rowspan="1" colspan="1">25</td></tr><tr><td rowspan="1" colspan="1">8.1-16.0 mm</td><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1">35</td><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1">29</td><td rowspan="1" colspan="1">40</td><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">41</td><td rowspan="1" colspan="1">39</td></tr><tr><td rowspan="1" colspan="1">>16 mm</td><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">28</td><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">46</td></tr><tr><td rowspan="1" colspan="10"><italic>Wing morphology</italic></td></tr><tr><td rowspan="1" colspan="1">Macropt</td><td rowspan="1" colspan="1">71</td><td rowspan="1" colspan="1">69</td><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1">68</td><td rowspan="1" colspan="1">69</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">66</td><td rowspan="1" colspan="1">68</td><td rowspan="1" colspan="1">29</td></tr><tr><td rowspan="1" colspan="1">Poly/dimo</td><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">33</td></tr><tr><td rowspan="1" colspan="1">Brachypt</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">20</td><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">23</td></tr><tr><td rowspan="1" colspan="10"><italic>Shadiness</italic></td></tr><tr><td rowspan="1" colspan="1">Shady (forest)</td><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1">24</td><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">25</td></tr><tr><td rowspan="1" colspan="1">Generalist</td><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1">29</td><td rowspan="1" colspan="1">25</td><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">25</td><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">29</td></tr><tr><td rowspan="1" colspan="1">Open</td><td rowspan="1" colspan="1">63</td><td rowspan="1" colspan="1">69</td><td rowspan="1" colspan="1">36</td><td rowspan="1" colspan="1">64</td><td rowspan="1" colspan="1">78</td><td rowspan="1" colspan="1">20</td><td rowspan="1" colspan="1">64</td><td rowspan="1" colspan="1">64</td><td rowspan="1" colspan="1">28</td></tr><tr><td rowspan="1" colspan="10"><italic>Moisture</italic></td></tr><tr><td rowspan="1" colspan="1">Water/wet</td><td rowspan="1" colspan="1">39</td><td rowspan="1" colspan="1">35</td><td rowspan="1" colspan="1">29</td><td rowspan="1" colspan="1">38</td><td rowspan="1" colspan="1">35</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">37</td><td rowspan="1" colspan="1">42</td><td rowspan="1" colspan="1">32</td></tr><tr><td rowspan="1" colspan="1">Moist-dryish</td><td rowspan="1" colspan="1">29</td><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1">22</td><td rowspan="1" colspan="1">20</td></tr><tr><td rowspan="1" colspan="1">Dry</td><td rowspan="1" colspan="1">32</td><td rowspan="1" colspan="1">42</td><td rowspan="1" colspan="1">43</td><td rowspan="1" colspan="1">32</td><td rowspan="1" colspan="1">47</td><td rowspan="1" colspan="1">24</td><td rowspan="1" colspan="1">32</td><td rowspan="1" colspan="1">36</td><td rowspan="1" colspan="1">30</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="3"><italic>NOR</italic></td><td rowspan="1" colspan="3"><italic>FIN</italic></td><td rowspan="1" colspan="1"><italic>Niede</italic></td><td rowspan="1" colspan="1"><italic>Nordr</italic></td><td rowspan="1" colspan="1"><italic>Wadde</italic></td><td rowspan="1" colspan="1"><italic>Drent</italic></td></tr><tr><td rowspan="1" colspan="1"><italic>Classes</italic></td><td rowspan="1" colspan="1"><italic>All</italic></td><td rowspan="1" colspan="1"><italic>IUCN</italic></td><td rowspan="1" colspan="1"><italic>% IUCN</italic></td><td rowspan="1" colspan="1"><italic>All</italic></td><td rowspan="1" colspan="1"><italic>IUCN</italic></td><td rowspan="1" colspan="1"><italic>% IUCN</italic></td><td rowspan="1" colspan="1"><italic>IUCN</italic></td><td rowspan="1" colspan="1"><italic>IUCN</italic></td><td rowspan="1" colspan="1"><italic>IUCN</italic></td><td rowspan="1" colspan="1"><italic>IUCN</italic></td></tr><tr><td rowspan="1" colspan="11"><italic>Body size</italic></td></tr><tr><td rowspan="1" colspan="1">0.1-4.0 mm</td><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">16</td></tr><tr><td rowspan="1" colspan="1">4.1-8.0 mm</td><td rowspan="1" colspan="1">46</td><td rowspan="1" colspan="1">38</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">45</td><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">44</td><td rowspan="1" colspan="1">40</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">33</td></tr><tr><td rowspan="1" colspan="1">8.1-16.0 mm</td><td rowspan="1" colspan="1">28</td><td rowspan="1" colspan="1">36</td><td rowspan="1" colspan="1">24</td><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1">53</td><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">34</td><td rowspan="1" colspan="1">24</td><td rowspan="1" colspan="1">39</td></tr><tr><td rowspan="1" colspan="1">>16 mm</td><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">6</td><td rowspan="1" colspan="1">6</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">12</td></tr><tr><td rowspan="1" colspan="11"><italic>Wing morphology</italic></td></tr><tr><td rowspan="1" colspan="1">Macropt</td><td rowspan="1" colspan="1">64</td><td rowspan="1" colspan="1">63</td><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">70</td><td rowspan="1" colspan="1">81</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">71</td><td rowspan="1" colspan="1">74</td><td rowspan="1" colspan="1">74</td><td rowspan="1" colspan="1">64</td></tr><tr><td rowspan="1" colspan="1">Poly/dimo</td><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">21</td></tr><tr><td rowspan="1" colspan="1">Brachypt</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">15</td></tr><tr><td rowspan="1" colspan="11"><italic>Shadiness</italic></td></tr><tr><td rowspan="1" colspan="1">Shady (forest)</td><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">6</td><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">12</td></tr><tr><td rowspan="1" colspan="1">Generalist</td><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">27</td><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1">28</td><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">21</td></tr><tr><td rowspan="1" colspan="1">Open</td><td rowspan="1" colspan="1">64</td><td rowspan="1" colspan="1">70</td><td rowspan="1" colspan="1">20</td><td rowspan="1" colspan="1">63</td><td rowspan="1" colspan="1">76</td><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">66</td><td rowspan="1" colspan="1">63</td><td rowspan="1" colspan="1">79</td><td rowspan="1" colspan="1">67</td></tr><tr><td rowspan="1" colspan="11"><italic>Moisture</italic></td></tr><tr><td rowspan="1" colspan="1">Water/wet</td><td rowspan="1" colspan="1">37</td><td rowspan="1" colspan="1">47</td><td rowspan="1" colspan="1">22</td><td rowspan="1" colspan="1">40</td><td rowspan="1" colspan="1">47</td><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">46</td><td rowspan="1" colspan="1">51</td><td rowspan="1" colspan="1">53</td><td rowspan="1" colspan="1">31</td></tr><tr><td rowspan="1" colspan="1">Moist-dryish</td><td rowspan="1" colspan="1">32</td><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">30</td></tr><tr><td rowspan="1" colspan="1">Dry</td><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1">36</td><td rowspan="1" colspan="1">20</td><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">41</td><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">38</td><td rowspan="1" colspan="1">32</td><td rowspan="1" colspan="1">33</td><td rowspan="1" colspan="1">39</td></tr></tbody></table></table-wrap><p>The proportion of threatened species over all species in the five countries revealed some important issues (columns “% IUCN” in <xref ref-type="table" rid="T3">Table 3</xref>). First of all, relative to the total number of species per category, larger species tended to be more often threatened than smaller species (see <xref ref-type="bibr" rid="B298">Kotze and O’Hara 2003</xref>). For size classes 8.1–16.0 mm and >16 mm, the proportions of threatened species were 21–39% and 12–50%, respectively, whereas for the size classes 0.1–4.0 mm and 4.1–8.0 mm, they were 10–18% and 7–25%, respectively. After pooling species into larger (>8.1 mm) and smaller (0.1–8.0 mm) size classes, proportions of these were between 19–40% (mean 28%) and 8–23% (mean 16%), respectively. Regarding wing morphology, the proportions of threatened species were rather even among the categories. In this respect, wing morphology was not clearly related to species being threatened, except for the slight tendency of wing-polymorphic species being proportionally more frequently threatened in Denmark and brachypterous species in Norway. Regarding shadiness associations, open-area species<pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment> included proportionally slightly more threatened species than did species of very shady habitats or shadiness generalists. Regarding moisture associations, species associated with very dry habitats included proportionally more threatened species (17–43%) than wet-habitat species (14–29%) or moist/dryish-habitat species (5–26%).</p><p>To the extent one can generalise from these figures, in northern and western Europe (see also <xref ref-type="bibr" rid="B91">Casale and Busato 2008</xref> for southern Europe) particular attention should be paid to large carabids, species associated with very dry, open habitats (e.g. sand dunes, heathlands and calcareous meadows; see national Red Lists) and water-associated species (e.g. freshwater stream specialists and salt-marsh species; see national Red Lists). However, as is evident from the variation in percentages presented in <xref ref-type="table" rid="T3">Table 3</xref>, particular targets of conservation and management (habitat types and species) should vary from one area to another. Below, research-based evidence on how to protect these carabids by the application of conservation management is reviewed.</p><p><bold>ii. Habitat selection for conservation efforts.</bold> Undisturbed mature ecosystems, particularly nature reserves, are vital for the conservation of many carabid species (e.g. <xref ref-type="bibr" rid="B131">Desender 2005</xref>; <xref ref-type="bibr" rid="B522">Skłodowski 2006</xref>). Also edge habitats and habitat mosaics may be important for carabid conservation (e.g. <xref ref-type="bibr" rid="B294">Kotze 2000</xref>; <xref ref-type="bibr" rid="B185">Falke et al. 2000</xref>; <xref ref-type="bibr" rid="B229">Hatteland et al. 2005</xref>; <xref ref-type="bibr" rid="B7">Andorkó and Kádár 2006</xref>). The scarcity of certain habitat types has increased the need for active maintenance of remaining patches. For example, <xref ref-type="bibr" rid="B49">Bérces et al. (2008)</xref> used field and museum data, original research and communication among entomologists, and showed that <italic><named-content content-type="taxon-name">Carabus hungaricus</named-content></italic> can best be protected by active management of open meadows. Due to the on-going loss of natural and semi-natural areas and the intensification of agricultural practices in many countries, also particular anthropogenic habitats have become important for carabid conservation. Examples include roadside verges, former agricultural fields, urban waste-grounds, and sand and gravel pits (<xref ref-type="bibr" rid="B464">Plachter 1986</xref>; <xref ref-type="bibr" rid="B182">Eversham et al. 1996</xref>; <xref ref-type="bibr" rid="B547">Telfer and Eversham 1996</xref>; <xref ref-type="bibr" rid="B510">Schwerk 2000</xref>; <xref ref-type="bibr" rid="B581">Versteirt et al. 2002</xref>; <xref ref-type="bibr" rid="B289">Koivula and Kotze 2005</xref>).</p><p><bold>iii. Habitat connectivity.</bold> Habitat-patch isolation and fragmentation may be of major concern for carabids (<xref ref-type="bibr" rid="B145">De Vries 1994</xref>; <xref ref-type="bibr" rid="B277">Kinnunen et al. 1996</xref>; <xref ref-type="bibr" rid="B422">Noordijk et al. 2006</xref>; <xref ref-type="bibr" rid="B232">Hendrickx et al. 2009</xref>). A number of means of reducing the impact of fragmentation have been suggested (<xref ref-type="bibr" rid="B577">Vermeulen et al. 2002</xref>). For forested environments, <xref ref-type="bibr" rid="B548">Terlutter (1990)</xref> discovered for <italic><named-content content-type="taxon-name">Carabus auronitens</named-content></italic> two different gene flows from two old forest remnants into a recent, regenerated forest-field mosaic. Later on, <xref ref-type="bibr" rid="B459">Petit (1994)</xref> underlined the importance of hedgerow networks for forest carabid assemblages. However, more recent hedges may be sub-optimal for this purpose (<xref ref-type="bibr" rid="B550">Thiele 1971</xref>; <xref ref-type="bibr" rid="B217">Gruttke 1994</xref>). For open areas, on the other hand, <xref ref-type="bibr" rid="B575">Vermeulen and Opsteeg (1994)</xref> showed that roadside verges might be used either as habitat or as movement corridors connecting heathland patches. Both purposes may be served by roadsides, as shown by (<xref ref-type="bibr" rid="B285">Koivula (2002a</xref>, 2005) for Finnish forest roads and <xref ref-type="bibr" rid="B421">Noordijk (2009)</xref> for highway verges in the Netherlands. Moreover, <xref ref-type="bibr" rid="B578">Vermeulen and Spee (2005)</xref> stressed the importance of source habitats for nature restoration sites. Hence, the remaining patches of natural <pmc-comment>PageBreak</pmc-comment>habitat, corridors of similar, often man-made environments, and artificially created, larger patches may together form an efficient patch network for carabid conservation.</p><p><bold>iv. Habitat management.</bold> Because many important natural processes (wildfire, flooding, wind, grazing and insect outbreaks) are effectively prevented in many areas, particularly in urban environments, active maintenance is considered necessary to preserve certain vegetation types. Carabids respond varyingly to these efforts (<xref ref-type="bibr" rid="B581">Versteirt et al. 2002</xref>; <xref ref-type="bibr" rid="B102">Cuesta et al. 2006</xref>; <xref ref-type="bibr" rid="B542">Taboada et al. 2006a</xref>). The effects of grassland management were discussed in depth by <xref ref-type="bibr" rid="B484">Rushton et al. (1990)</xref> who found that some species avoid intensively managed sites, some are favoured by these, while others showed intermediate or no detectable responses. Similarly, <xref ref-type="bibr" rid="B52">Blake et al. (1996)</xref> showed that vegetation management in wildflower meadows resulted in a decrease in large species and an increase in xerophilous species, while species characteristic of areas with ‘natural’ conditions were absent. Like mowing, grazing also profoundly affects carabids: its intensity determines assemblage composition (<xref ref-type="bibr" rid="B374">McFerran et al. 1994</xref>). <xref ref-type="bibr" rid="B98">Cole et al. (2006)</xref> showed that intensive grazing decreases the abundance of large <italic><named-content content-type="taxon-name">Carabus</named-content></italic> species more than less intensive grazing. These studies indicate that variation in management leads to variation in carabid beetle assemblages. In riparian environments, <xref ref-type="bibr" rid="B201">Fuellhaas (2000)</xref> showed that raising the water-table level increases the number of hygrophilic species. <xref ref-type="bibr" rid="B198">Främbs (1990)</xref> studied regenerating peat bogs and found that although carabid diversity increased, the peat-bog specialist <italic><named-content content-type="taxon-name">Agonum ericeti</named-content></italic> remained absent. <xref ref-type="bibr" rid="B154">Drees et al. (2007)</xref> argued that this might be related to habitat quality, in this case the lack of peat-producing vegetation.</p><p>To summarise, (i) Carabid conservation should give special attention to very large species, and species associated with both very wet and very dry, exposed conditions, (ii) Old and undisturbed natural areas are important for many specialists, but conservation of pioneer or open-habitat species can be realised in many anthropogenic areas as well (<xref ref-type="fig" rid="F6">Fig. 4</xref>), (iii) Fragmentation potentially isolates local populations, but its effects can be decreased by maintaining large, inter-connected areas, corridor networks, and designing restoration areas near potential source areas, and (iv) Guidelines for active management of carabid habitats are difficult to draft, as some species respond negatively to any disturbance, including conservation management. However, many species urgently need small-scale management that keeps habitats constantly at some preferred successional phase; most of these species are subject to severe stress in modern, fragmented landscapes.</p><fig id="F6" orientation="portrait" position="float"><label>Figure 4.</label><caption><p><italic><named-content content-type="taxon-name">Cylindera germanica</named-content></italic> (Photo by Jinze Noordijk)</p></caption><graphic xlink:href="ZooKeys-100-055-g006"></graphic></fig><p>The conservation of carabids and their habitats is far from perfect. This issue is complicated by the fact that, due to these beetles’ mobility, occupation of varyingly sized habitat patches, varying degrees of specialisation, and development through numerous developmental phases, their ecological requirements vary in time and place. As habitat patches of carabid assemblages usually include several vegetation types and/or physical structures, a conservation approach targeted for maintaining only particular vegetation types or high plant diversity may not always be appropriate for the conservation of arthropod assemblages (<xref ref-type="bibr" rid="B451">Panzer and Schwartz 1998</xref>; <xref ref-type="bibr" rid="B126">Dennis et al. 2007</xref>). Moreover, the<pmc-comment>PageBreak</pmc-comment> high number of carabid species, each with specific demands, makes it difficult to define a single conservation strategy. Protection of whole landscapes with mosaics of distinct habitat types may prove efficient for carabid conservation. Simultaneously, the natural variety of successional stages should be conserved, as particular stages can be crucial for certain species (cf. <xref ref-type="bibr" rid="B413">Niemelä et al. 2007</xref>). Moreover, some species – such as <italic><named-content content-type="taxon-name">Amara plebeja</named-content></italic> (<xref ref-type="bibr" rid="B567">van Huizen 1977</xref>) – possibly require more than just one habitat type and/or successional stage to persist in a landscape (‘landscape species’; Szyszko<xref ref-type="bibr" rid="B205"> 2004</xref>; <xref ref-type="bibr" rid="B540">Szyszko et al. 2011</xref>; Axel Schwerk, pers. comm.). Habitat patches within these mosaics should include particular structures, such as micro-relief, patches of bare sand, stony patches, small water bodies, heaps of decaying plant material, and dead wood, features that are often of no special importance for plants and vertebrates and therefore often ignored if conservation management is based on vegetation data alone. Thus, a broad landscape approach, supplemented by these small-scale structures, may produce good results for the conservation of carabid beetles (<xref ref-type="bibr" rid="B279">Kirby 1992</xref>; <xref ref-type="bibr" rid="B402">New 1995</xref>, 2010; <xref ref-type="bibr" rid="B490">Samways 2005</xref>, 2007; <xref ref-type="bibr" rid="B228">Haslett 2007</xref>).</p></sec><sec><title>8 Landscape ecology</title><p>How carabid beetles perceive space may influence habitat selection, home ranges, the dispersal of individuals and the dynamics and distributions of populations. Furthermore, the amount, extent and spatial arrangement of suitable habitats within a landscape (i.e. landscape composition and configuration) may affect long-term population <pmc-comment>PageBreak</pmc-comment>persistence. Thus, although the spatial distribution of carabid beetles may be primarily determined by microhabitat conditions and biotic interactions at the local scale, identifying general patterns of carabid responses to landscape features may help us to understand how species, functional groups and assemblages effectively distribute, and to predict how they will cope with current and future land-use and climatic changes. As such, the spatial context related to a species’ distribution patterns is an essential component when studying how global changes affect carabid species conservation.</p><p>Over the last 40 years, investigations of ground beetle landscape ecology have demonstrated that landscape features influence not only the spatial distribution of these beetles, but also their population dynamics (<xref ref-type="bibr" rid="B368">Matalin 1997c</xref>; <xref ref-type="bibr" rid="B55">Bommarco 1998</xref>) and genetic structure (<xref ref-type="bibr" rid="B76">Brouat et al. 2003</xref>; <xref ref-type="bibr" rid="B276">Keller et al. 2004</xref>; <xref ref-type="bibr" rid="B140">Desender et al. 2005</xref>; <xref ref-type="bibr" rid="B492">Sander et al. 2006</xref>). From the late 1950s until the mid 1970s, contributions to carabid beetle ecology aimed at characterising the structure and composition of communities occurring in specific types of landscapes, which were, at that stage, considered as homogeneous entities (e.g. forested vs. open landscapes; see <xref ref-type="bibr" rid="B551">Thiele 1977</xref>). Research developed in the 1980s and 1990s confirmed the significance of heterogeneity within landscapes and thus addressed the role of singular landscape elements or habitat types for the carabid fauna in a variety of either natural or highly-modified and simplified landscapes. In tests of the application of the theory of island biogeography (<xref ref-type="bibr" rid="B346">MacArthur and Wilson 1967</xref>) to carabid communities, it has been shown that local communities are not simply a passive random sample of the regional species pool, but that species are filtered according to the association of their life history traits to habitat quality, configuration and biotic interactions (e.g. <xref ref-type="bibr" rid="B474">Ranta and Ås 1982</xref>; <xref ref-type="bibr" rid="B417">Niemelä et al. 1985</xref>; <xref ref-type="bibr" rid="B146">De Vries et al. 1996</xref>). Studies that have looked for an island effect in carabid assemblages of patches of terrestrial habitats have generally concluded that such patches are not sufficiently isolated to represent islands, due to the strong dispersal capacity of many carabid species (<xref ref-type="bibr" rid="B107">Davies and Margules 1998</xref>; <xref ref-type="bibr" rid="B353">Magura et al. 2001</xref>; <xref ref-type="bibr" rid="B75">Brose 2003</xref>). Studies have also been conducted on carabid assemblages of real islands, and these too have concluded that a simple species-area relationship explains the differences in carabid species richness between islands of different size better than distance from mainland populations (<xref ref-type="bibr" rid="B296">Kotze and Niemelä 2002</xref>; Zalewski<xref ref-type="bibr" rid="B205"> 2004</xref>).</p><p>In recent literature, studies on the importance of the landscape context in determining the occurrence of carabid species based on different aspects of landscape composition, configuration, connectivity, history, land-use type and intensity have proliferated (e.g. <xref ref-type="bibr" rid="B470">Purtauf et al. 2004</xref>; <xref ref-type="bibr" rid="B73">Bräuniger et al. 2010</xref>; <xref ref-type="bibr" rid="B209">Gardiner et al. 2010</xref>; <xref ref-type="bibr" rid="B393">Nabe-Nielsen et al. 2010</xref>; <xref ref-type="bibr" rid="B599">Woodcock et al. 2010</xref>). Many studies analysed the influence of the landscape context on overall carabid beetle activity density and species richness, often finding no statistically significant effect. Mostly, changes in landscape features have been related to shifts in carabid species composition, and variations in the activity density of individual species and ecologically meaningful groups (e.g. Niemelä<xref ref-type="bibr" rid="B204"> 2001</xref>; <xref ref-type="bibr" rid="B298">Kotze and O’Hara 2003</xref>; <xref ref-type="bibr" rid="B413">Niemelä et al. 2007</xref>; <xref ref-type="bibr" rid="B414">Niemelä and Kotze 2009</xref>).</p><p>Agricultural landscapes in particular, driven by daily, seasonal and annual fluctuations, soon became the subject of many carabid beetle surveys, followed by an extensive<pmc-comment>PageBreak</pmc-comment> number of publications to date (e.g. <xref ref-type="bibr" rid="B278">Kinnunen et al. 2001</xref>; <xref ref-type="bibr" rid="B242">Holland 2002</xref>). In general, the basic composition of the carabid fauna of agricultural mosaic landscapes appears to be surprisingly similar across countries (<xref ref-type="bibr" rid="B338">Luff 2002</xref>), dominated by eurytopic species, which are highly tolerant to disturbance. However, the size, amount, isolation and spatial arrangement of agricultural patches, the composition of the arable mosaic, as well as the occurrence of permanent landscape elements (e.g. hedgerows, field margins, natural woodlands and grasslands), affect carabid beetle assemblages (<xref ref-type="bibr" rid="B277">Kinnunen et al. 1996</xref>, <xref ref-type="bibr" rid="B278">2001</xref>; <xref ref-type="bibr" rid="B82">Burel et al. 1998</xref>; <xref ref-type="bibr" rid="B461">Petit and Usher 1998</xref>; <xref ref-type="bibr" rid="B197">Fournier and Loreau 2001</xref>; <xref ref-type="bibr" rid="B378">Millán de la Peña et al. 2003</xref>; <xref ref-type="bibr" rid="B30">Aviron et al. 2005</xref>; <xref ref-type="bibr" rid="B471">Purtauf et al. 2005</xref>; <xref ref-type="bibr" rid="B216">Griffiths et al. 2007</xref>; <xref ref-type="bibr" rid="B233">Hendrickx et al. 2007</xref>; <xref ref-type="bibr" rid="B503">Saska et al. 2007</xref>).</p><p>In forest ecosystems, natural and anthropogenic disturbances create a dynamic mosaic of successional habitat patches for carabids (e.g. <xref ref-type="bibr" rid="B60">Bouget and Duelli 2004</xref> for windstorm disturbance). Each forest successional stage is characterised by a specific carabid assemblage, in terms of species composition as well as ecological group composition, with the greatest differences between early and advanced stages (e.g. <xref ref-type="bibr" rid="B535">Szyszko 1990</xref>; <xref ref-type="bibr" rid="B411">Niemelä et al. 1996</xref>; <xref ref-type="bibr" rid="B84">Butterfield 1997</xref>; <xref ref-type="bibr" rid="B291">Koivula et al. 2002</xref>; <xref ref-type="bibr" rid="B156">Du Bus de Warnaffe and Lebrun 2004</xref>; <xref ref-type="bibr" rid="B479">Richard et al. 2004</xref>; <xref ref-type="bibr" rid="B355">Magura et al. 2006</xref>; <xref ref-type="bibr" rid="B544">Taboada et al. 2008</xref>). Changes in population dynamics and morphological traits also take place through succession (e.g. Szysko et al. 1996 for <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic>, <xref ref-type="table" rid="T4">Table 4</xref>). Changes in the carabid fauna are possibly correlated with the amount of carbon accumulation in the forest system, i.e. in the wood, litter and mineral soil (<xref ref-type="bibr" rid="B537">Szyszko 2010</xref>; <xref ref-type="bibr" rid="B540">Szyszko et al. 2011</xref>). The increase of carbon in the mineral soil is related to the decomposition of litter by the macrofauna. For pine stands in Poland, <xref ref-type="bibr" rid="B533">Szyszko (1986a)</xref> demonstrated that biomass of the macrofauna is correlated with parameters of the carabid fauna, such as species number and Mean Individual Biomass (MIB). MIB increases as succession progresses (<xref ref-type="bibr" rid="B534">Szyszko 1986b</xref>; <xref ref-type="bibr" rid="B541">Szyszko et al. 2000</xref>; Szyszko<xref ref-type="bibr" rid="B205"> 2004</xref>), suggesting that this measure functions as a good indicator of the state of succession (see <italic>Bioindicators</italic> above). The rate at which species composition changes during succession and the successional trajectory followed by the carabid assemblages depends on environmental conditions, such as soil properties (<xref ref-type="bibr" rid="B534">Szyszko 1986b</xref>, 1990; <xref ref-type="bibr" rid="B511">Schwerk 2008</xref>), dominant tree species (<xref ref-type="bibr" rid="B156">Du Bus de Warnaffe and Lebrun 2004</xref>), and the type of disturbance that initiated the succession (<xref ref-type="bibr" rid="B156">Du Bus de Warnaffe and Lebrun 2004</xref>). Indeed, the larger the newly-created gap is and the fewer trees retained, the more severe the perturbation for carabid assemblages (<xref ref-type="bibr" rid="B286">Koivula 2002b</xref> following timber harvest; <xref ref-type="bibr" rid="B59">Bouget 2005</xref> and <xref ref-type="bibr" rid="B523">Skłodowski and Garbalińska 2010</xref> following windthrow gap). As a consequence, the maintenance of a variety of successional phases of the forest cycle results in increased heterogeneity at the landscape level and, therefore greater regional carabid diversity (e.g. <xref ref-type="bibr" rid="B390">Mullen et al. 2008</xref>; <xref ref-type="bibr" rid="B544">Taboada et al. 2008</xref>). Thus, the effects of forest landscape features on the carabid fauna have also been extensively addressed as regards to landscape heterogeneity, the occurrence, composition and spatial configuration of either natural or human-modified habitats (e.g. proportion of deciduous vs. coniferous forests, age and extent of exotic plantations, forest edge density and permeability), the role of particular landscape elements (e.g. retention tree groups), and the landscape context <pmc-comment>PageBreak</pmc-comment>resulting from historical and/or recent management practices (<xref ref-type="bibr" rid="B291">Koivula et al. 2002</xref>; <xref ref-type="bibr" rid="B58">Bouget 2004</xref>; <xref ref-type="bibr" rid="B40">Barbaro et al. 2005</xref>; <xref ref-type="bibr" rid="B372">Matveinen-Huju et al. 2006</xref>; <xref ref-type="bibr" rid="B543">Taboada et al. 2006b</xref>; <xref ref-type="bibr" rid="B41">Barbaro et al. 2007</xref>; <xref ref-type="bibr" rid="B413">Niemelä et al. 2007</xref>; <xref ref-type="bibr" rid="B202">Fuller et al. 2008</xref>; <xref ref-type="bibr" rid="B453">Pawson et al. 2008</xref>; <xref ref-type="bibr" rid="B42">Barbaro and van Halder 2009</xref>).</p><table-wrap id="T4" orientation="portrait" position="float"><label>Table 4.</label><caption><p>Changes in carabid fauna, interaction groups and populations of <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic> with changes in habitat (according to <xref ref-type="bibr" rid="B539">Szyszko et al. 1996</xref>, reprinted and modified with permission from Aarhus University Press).</p></caption><table frame="hsides" rules="groups"><tbody><tr><td rowspan="1" colspan="1"><italic>Comparatively early stage of succession</italic></td><td rowspan="1" colspan="3"></td><td rowspan="1" colspan="1"><italic>Comparatively late stage of succession</italic></td></tr><tr><td rowspan="1" colspan="5"><named-content content-type="taxon-name">Carabidae</named-content> fauna</td></tr><tr><td rowspan="1" colspan="1">low state of development of fauna</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">high state of development of fauna</td></tr><tr><td rowspan="1" colspan="1">high number of species</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">low number of species</td></tr><tr><td rowspan="1" colspan="1">small individuals</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">large individuals</td></tr><tr><td rowspan="1" colspan="1">low mean individual biomass (MIB)</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">high mean individual biomass (MIB)</td></tr><tr><td rowspan="1" colspan="5">Interaction group of <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">long period of activity</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">short period of activity</td></tr><tr><td rowspan="1" colspan="1">long survival of adults</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">short survival of adults</td></tr><tr><td rowspan="1" colspan="1">complicated age structure</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">simple age structure</td></tr><tr><td rowspan="1" colspan="1">small individuals (imago)</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">big individuals (imago)</td></tr><tr><td rowspan="1" colspan="1">high proportion of males</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">high proportion of females</td></tr><tr><td rowspan="1" colspan="1">low number of eggs in ovaries</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">high number of eggs in ovaries</td></tr><tr><td rowspan="1" colspan="1">high number of eggs laid?</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">low number of eggs laid?</td></tr><tr><td rowspan="1" colspan="1">good food situation for adults?</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">bad food situation for adults?</td></tr><tr><td rowspan="1" colspan="1">bad food situation for larvae?</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">good food situation for larvae?</td></tr><tr><td rowspan="1" colspan="1">unable to fly?</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">able to fly?</td></tr><tr><td rowspan="1" colspan="1">uneconomic life strategy</td><td rowspan="1" colspan="3">→</td><td rowspan="1" colspan="1">economic life strategy</td></tr><tr><td rowspan="1" colspan="5">Populations of <italic><named-content content-type="taxon-name">Pterostichus oblongopunctatus</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">asynchronously fluctuating interaction groups</td><td rowspan="1" colspan="1">→</td><td rowspan="1" colspan="1">synchronously fluctuating interaction groups</td><td rowspan="1" colspan="1">→</td><td rowspan="1" colspan="1">asynchronously fluctuating interaction groups</td></tr><tr><td rowspan="1" colspan="1">low probability of high fluctuations of numbers</td><td rowspan="1" colspan="1">→</td><td rowspan="1" colspan="1">high probability of high fluctuations of numbers</td><td rowspan="1" colspan="1">→</td><td rowspan="1" colspan="1">low probability of high fluctuations of numbers</td></tr><tr><td rowspan="1" colspan="1">resistant population</td><td rowspan="1" colspan="1">→</td><td rowspan="1" colspan="1">not very resistant population</td><td rowspan="1" colspan="1">→</td><td rowspan="1" colspan="1">resistant population</td></tr></tbody></table></table-wrap><p>Much effort has been devoted to investigating how carabid beetles are distributed in fragmented landscapes and insular environments (for reviews, see Niemelä<xref ref-type="bibr" rid="B204"> 2001</xref> and <xref ref-type="bibr" rid="B295">Kotze 2008</xref>, respectively). Carabid responses to fragmentation depend on the geographical context, are species specific and, to a great extent, relate to species’ life history traits and habitat associations (e.g. <xref ref-type="bibr" rid="B293">Koivula and Vermeulen 2005</xref>; <xref ref-type="bibr" rid="B211">Gaublomme et al. 2008</xref>). In a fragmented landscape context, mobility is crucial for persistence, especially for specialist and scarce species (<xref ref-type="bibr" rid="B145">De Vries 1994</xref>; <xref ref-type="bibr" rid="B146">De Vries et al. 1996</xref>). In general, good<pmc-comment>PageBreak</pmc-comment> dispersers and abundant species are expected to maintain populations in small and isolated patches through recolonisation of empty patches, whereas poor colonisers and scarce species may not be able to do so (<xref ref-type="bibr" rid="B113">Den Boer 1977</xref>; Niemelä<xref ref-type="bibr" rid="B204"> 2001</xref>). Hostile types of matrix or linear elements in the landscape can act as dispersal barriers for specialist species. For instance, some forest species are reluctant to cross highways (<xref ref-type="bibr" rid="B350">Mader 1984</xref>; <xref ref-type="bibr" rid="B293">Koivula and Vermeulen 2005</xref>) or open habitats (<xref ref-type="bibr" rid="B465">Plat et al. 1995</xref>; <xref ref-type="bibr" rid="B480">Riecken and Raths 1996</xref>), while other stenotopic species effectively move along hedgerows (<xref ref-type="bibr" rid="B81">Burel 1989</xref>; <xref ref-type="bibr" rid="B465">Plat et al. 1995</xref>; <xref ref-type="bibr" rid="B97">Charrier et al. 1997</xref>) and roadside verges (<xref ref-type="bibr" rid="B573">Vermeulen 1993</xref>, 1994; <xref ref-type="bibr" rid="B576">Vermeulen and Opdam 1995</xref>), which function as movement corridors for such species. Dirt roads in forested landscapes may serve as dispersal corridors for open habitat species (<xref ref-type="bibr" rid="B285">Koivula 2002a</xref>), and roadsides overgrown with poplars have been suggested to serve as corridors for forest species with low dispersal power (<xref ref-type="bibr" rid="B165">Dymitryszyn et al. 2003</xref>). Attempts to improve the connectivity of landscape elements by means of corridors may have contrasting effects on different carabid species according to their habitat requirements and, hence, new approaches regarding this matter are now under evaluation, such as semi-open corridors (<xref ref-type="bibr" rid="B166">Eggers et al. 2010</xref>) and innovative, small scale forest harvesting techniques (<xref ref-type="bibr" rid="B291">Koivula et al. 2002</xref>, see also <italic>Carabid conservation, protection and habitat management</italic> above).</p><p>Further studies have investigated the responses of ground beetles to anthropogenic or human-modified landscapes and urban environments (e.g. <xref ref-type="bibr" rid="B103">Czechowski 1982</xref>; <xref ref-type="bibr" rid="B281">Klausnitzer and Richter 1983</xref>; <xref ref-type="bibr" rid="B529">Šustek 1987</xref>, 1992; <xref ref-type="bibr" rid="B416">Niemelä et al. 2002</xref>). These investigations have identified distinct sets of species associated with the urban cores or city centres (but see <xref ref-type="bibr" rid="B416">Niemelä et al. 2002</xref>). However, for a considerable number of these species, urban populations may be dependent on recruitment from populations in the urban periphery (<xref ref-type="bibr" rid="B281">Klausnitzer and Richter 1983</xref>). The possible effects of urbanisation on carabid population genetics (i.e. genetic diversity and differentiation) remain unclear (<xref ref-type="bibr" rid="B140">Desender et al. 2005</xref>). In general, the overall abundance and species richness of carabids decrease with increasing urbanisation (<xref ref-type="bibr" rid="B414">Niemelä and Kotze 2009</xref>; but see <xref ref-type="bibr" rid="B354">Magura et al. 2010</xref>). Also, large species tend to be relatively scarce in urban habitats, resulting in a decline in average body size in urban areas compared to less disturbed ones, both for forest assemblages (<xref ref-type="bibr" rid="B416">Niemelä et al. 2002</xref>; <xref ref-type="bibr" rid="B259">Ishitani et al. 2003</xref>; <xref ref-type="bibr" rid="B486">Sadler et al. 2006</xref>; <xref ref-type="bibr" rid="B168">Elek and Lövei 2007</xref>) and those of open habitats (<xref ref-type="bibr" rid="B103">Czechowski 1982</xref>; <xref ref-type="bibr" rid="B529">Šustek 1987</xref>; <xref ref-type="bibr" rid="B569">Venn 2007</xref>). Flightless species also tend to be relatively scarce in urban assemblages (<xref ref-type="bibr" rid="B570">Venn et al. 2003</xref>; <xref ref-type="bibr" rid="B486">Sadler et al. 2006</xref>). Other responses detected in carabid assemblages (either in the proportion of species or the number of individuals) to urbanisation include (i) a decrease in species with restricted geographical ranges, along with the enhancement of those distributed over broad ranges; (ii) a decline in oligotopic, stenotopic and specialist species, whilst eurytopic, polytopic and generalist ones increase; (iii) a decrease in forest species and associated increase in open habitat species; (iv) an increase in xerophilic and mesohygrophilous species at the expense of more hygrophilous species; and (v) an increase in omnivorous species and a corresponding decrease in zoophagous species. Whilst stenotopic and specialist species generally decline with <pmc-comment>PageBreak</pmc-comment>increasing urbanisation, some extremely harsh urban habitats accommodate these species, such as populations of <italic><named-content content-type="taxon-name">Amara equestris</named-content></italic> in central reservations of a busy ring road in Helsinki, Finland (<xref ref-type="bibr" rid="B287">Koivula et al. 2005</xref>). In fact, <xref ref-type="bibr" rid="B182">Eversham et al. (1996)</xref> reported that more than 35% of Britain’s rare and scarce carabids are to be found from manmade sites, <italic><named-content content-type="taxon-name">Omophron limbatum</named-content></italic> and <italic><named-content content-type="taxon-name">Dyschirius obscurus</named-content></italic> exclusively so. Subsequent to numerous urbanisation studies from single cities, the GLOBENET project (<xref ref-type="bibr" rid="B415">Niemelä et al. 2000</xref>; <ext-link ext-link-type="uri" xlink:href="http://www.helsinki.fi/science/globenet">http://www.helsinki.fi/science/globenet</ext-link>) was established to apply a standard urbanisation gradient approach in cities across the globe (nine cities located in Europe, Japan and Canada). The main findings indicated that the carabid fauna of urban forested habitats display uniform patterns of response to the degree of urbanisation of the ‘concrete’ matrix (<xref ref-type="bibr" rid="B414">Niemelä and Kotze 2009</xref>; <xref ref-type="bibr" rid="B354">Magura et al. 2010</xref>).</p><p>In the BIOASSESS project (<ext-link ext-link-type="uri" xlink:href="http://www.nbu.ac.uk/bioassess">http://www.nbu.ac.uk/bioassess</ext-link>), global patterns in carabid responses to a land-use intensity gradient from old-growth or unmanaged forests to arable crop-dominated landscape across ten countries, have so far reported effects on overall species richness, number of individuals, ecological groups and species composition (<xref ref-type="bibr" rid="B214">Grandchamp et al. 2005</xref>; <xref ref-type="bibr" rid="B509">Schweiger et al. 2005</xref>; <xref ref-type="bibr" rid="B563">Vanbergen et al. 2005</xref>, <xref ref-type="bibr" rid="B564">2010</xref>; <xref ref-type="bibr" rid="B233">Hendrickx et al. 2007</xref>; <xref ref-type="bibr" rid="B363">Martins da Silva et al. 2008</xref>). Similarly, changes in landscape structure over time (i.e. landscape history) have been addressed when investigating carabid population declines or range-size modifications in human-altered landscapes (<xref ref-type="bibr" rid="B560">Turin and Den Boer 1988</xref>; <xref ref-type="bibr" rid="B137">Desender et al. 1994b</xref>; <xref ref-type="bibr" rid="B460">Petit and Burel 1998</xref>; <xref ref-type="bibr" rid="B298">Kotze and O’Hara 2003</xref>). Additionally, these investigations have related contemporary distribution patterns of carabid endemism, rarity and habitat specialisation across landscapes to landscape history.</p><p>Even now, the spatial scale at which carabid beetles relate to resources across landscapes is not completely understood. Future studies should accomplish multiscale approaches that consider a wide range of fine and coarse grains at which each carabid species may perceive the landscape, depending on its mobility and body size (e.g. <xref ref-type="bibr" rid="B83">Burel et al. 2004</xref>; <xref ref-type="bibr" rid="B30">Aviron et al. 2005</xref>; <xref ref-type="bibr" rid="B260">Janssen et al. 2009</xref>). The spatial distribution of carabid species in a given landscape is nearly always aggregated at some scale (see e.g. <xref ref-type="bibr" rid="B411">Niemelä et al. 1996</xref>; <xref ref-type="bibr" rid="B552">Thomas et al. 2002</xref>), which suggests that spatial autocorrelation should always be taken into account (<xref ref-type="bibr" rid="B45">Barton et al. 2009</xref>). Additionally, the use of multiple habitats by carabid species in mosaic heterogeneous landscapes (e.g. for feeding, reproducing and overwintering; <xref ref-type="bibr" rid="B567">van Huizen 1977</xref>), and the importance of particular habitat combinations for a species’ survival at the landscape level remain unclear (<xref ref-type="bibr" rid="B41">Barbaro et al. 2007</xref>). Moreover, since many of the reported carabid responses to landscape features are species specific, more attention should be devoted to the individual species level, and not only for species of present conservation concern but also for common and widely distributed species, as well as to the ecological group level. Nonetheless, sampling strategies that avoid confounding effects are needed to clearly assess the respective weights of local and landscape factors and, at the landscape scale, the respective importance of composition and configuration on a species’<pmc-comment>PageBreak</pmc-comment> survival. Indeed, experimental landscapes (<xref ref-type="bibr" rid="B107">Davies and Margules 1998</xref>) or mensurative experiments (<xref ref-type="bibr" rid="B254">Hurlbert 1984</xref>) would be useful to disentangle these gradients that tend to be naturally correlated (Niemelä<xref ref-type="bibr" rid="B204"> 2001</xref>; <xref ref-type="bibr" rid="B184">Fahrig 2003</xref>). Attention should also be paid to discrepancies between carabid species’ responses to landscape features across countries, possibly denoting that the impact of landscape structure on a particular species is likely to differ over its distribution range. Finally, in the current context of continuous landscape transformation, more emphasis should be given to the role of newly created habitats and abandoned areas across countries regarding carabid distribution, as well as to singular expanding elements and surrogate habitats, such as golf courses or private gardens in urban environments (<xref ref-type="bibr" rid="B545">Tanner and Gange 2005</xref>; <xref ref-type="bibr" rid="B485">Saarikivi et al. 2010</xref>), roads (<xref ref-type="bibr" rid="B293">Koivula and Vermeulen 2005</xref>; <xref ref-type="bibr" rid="B375">Melis et al. 2010</xref>; <xref ref-type="bibr" rid="B600">Yamada et al. 2010</xref>), power lines (<xref ref-type="bibr" rid="B244">Hollmen et al. 2008</xref>) and biomass crops (coppice with short and very-short rotation) in either open or forested landscapes. Eventually, potential mechanisms could be investigated by confronting the empirical data with models of dispersal and survival in heterogeneous landscapes (see e.g. <xref ref-type="bibr" rid="B575">Vermeulen and Opsteeg 1994</xref>; <xref ref-type="bibr" rid="B462">Pichancourt et al. 2006</xref>).</p></sec><sec><title>9 Concluding remarks</title><p>Carabids are among the most species-rich families of beetles, which has made them a natural focus of entomological research. Carabidologists are busy studying this evolutionarily successful group at several levels, from sub-cellular to supra-individual. Indeed, from the discovery of a pH receptor on the antennae of carabid beetles (<xref ref-type="bibr" rid="B376">Merivee et al. 2005</xref>; <xref ref-type="bibr" rid="B377">Milius et al. 2006</xref>) to cross-continental, landscape related research (<xref ref-type="bibr" rid="B414">Niemelä and Kotze 2009</xref>; <xref ref-type="bibr" rid="B354">Magura et al. 2010</xref>; <xref ref-type="bibr" rid="B564">Vanbergen et al. 2010</xref>) “carabidologists do it all”. They are helped by a reasonably solid taxonomy, even if evolutionary relationships are still undetermined.</p><p>Carabidology has contributed to several prominent ecological theories, including metapopulation theory (pioneering work by Piet den Boer and colleagues), and provides one of the best examples of a consistent, systematic study of the effects of urbanisation on biodiversity (<xref ref-type="bibr" rid="B416">Niemelä et al. 2002</xref>, and subsequent studies). These somewhat ad hoc examples are still powerful in the argumentation to encourage the use of carabids in ecological, evolutionary and behavioural studies.</p><p>Even from a subjective summary as this article admittedly is, it is obvious that carabids have contributed in a major way to our understanding of invertebrate adaptations, phylogeny and ecology. Accepting Hutchinson’s analogy that on the world stage an ecological play is being played out in the evolutionary theatre (<xref ref-type="bibr" rid="B250">Hutchinson 1965</xref>), watching and describing the peculiarities of one of the star players, ground beetles, will certainly advance our understanding of nature. In an age in which the earth is dominated by humans, this will provide important knowledge on how to maintain the richness of life on Earth, and with it, extend the lifespan of our own species.</p></sec></body><back><ack><title>Acknowledgements</title><p>We thank Thorsten Assmann and Piet den Boer for useful insights, Terry Erwin for information and reviewing this submission, and Ivailo Stoyanov and Lyubomir Penev for support and advice. For some authors, partial support was received from the Academy of Finland (JK, SV, project number 126915), the Czech Science Foundation (PS, grant # 206/09/1266), and the Ministerio de Ciencia e Innovación and the Fundación Séneca, Murcia (JS, respective project numbers: CGL2006–06706 and 08724PI08). The 14th ECM at Westerbork, the Netherlands received financial support from the Uyttenboogaart-Eliasen Foundation and the Province of Drente. 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