Lutra 53-2 - Nieuw in de Zoogdierwinkel
Lutra 53-2 - Nieuw in de Zoogdierwinkel
Lutra 53-2 - Nieuw in de Zoogdierwinkel
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Journal of the Dutch Mammal Society<br />
Volume <strong>53</strong> – Number 2 – December 2010
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
LUTRA Volume Deel <strong>53</strong> – Number Nummer 2 – December 2010<br />
<strong>Lutra</strong> is a scientific journal published by the Dutch Mammal Society (Zoogdier verenig<strong>in</strong>g). The society is <strong>de</strong>dicated<br />
to the study and protection of native mammals <strong>in</strong> Europe. <strong>Lutra</strong> publishes peer-reviewed scientific papers on mammals<br />
across all discipl<strong>in</strong>es, but tends to focus on ecology, biogeography, behaviour and morphology. Although<br />
exceptions are ma<strong>de</strong> <strong>in</strong> some cases, <strong>Lutra</strong> generally publishes articles on mammal species native to Europe,<br />
<strong>in</strong>clud<strong>in</strong>g mar<strong>in</strong>e mammals. <strong>Lutra</strong> publishes full articles as well as short notes which may <strong>in</strong>clu<strong>de</strong> novel research<br />
methods or remarkable observations of mammals. In addition <strong>Lutra</strong> publishes book reviews, and compilations of<br />
recent literature on mammals. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch. <strong>Lutra</strong> publishes two issues per<br />
year and <strong>Lutra</strong> is <strong>in</strong><strong>de</strong>xed <strong>in</strong> ‘Biological Abstracts’ and ‘Zoological Record’ and ‘Artik’.<br />
Editorial board Redactie Kees J. Canters (editor-<strong>in</strong>-chief), Jasja J.A. Dekker, Jan Piet Bekker,<br />
Kees (C.J.) Camphuysen, Bastiaan G. Meerburg (secretary),<br />
Johan B.M. Thissen, Koen Van Den Berge, Ben Verboom (manag<strong>in</strong>g editor)<br />
Editorial assistant Redactie-assistent Eric van Kaathoven<br />
Address Adres Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
Website <strong>Lutra</strong> is available from the <strong>in</strong>ternet: www.zoogdierverenig<strong>in</strong>g.nl<br />
Subscription Abonnement The annual fee for a subscription to <strong>Lutra</strong> is € 22,– (€ 25,50 for non-Euro<br />
countries)<br />
The annual fee for a <strong>Lutra</strong> subscription and Zoogdierverenig<strong>in</strong>g membership<br />
is € 34,– (€ 37,50 for non-Euro countries)<br />
This also <strong>in</strong>clu<strong>de</strong>s a subscription to Zoogdier, a journal that publishes <strong>in</strong><br />
Dutch only. Stu<strong>de</strong>nts are entitled to a discount of € 4,50 for the first two<br />
years of their Zoogdierverenig<strong>in</strong>g-membership.<br />
Send payment to one of the follow<strong>in</strong>g accounts:<br />
the Netherlands: account 203737 of the ‘ING’,<br />
addressed to Zoogdier verenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
(IBAN: NL26INGB0000203737; BIC: INGBNL2A).<br />
Belgium: account 000-1486269-35 of the ‘Bank van <strong>de</strong> Post’,<br />
addressed to Zoogdierverenig<strong>in</strong>g, Nijmegen, the Netherlands.<br />
Sticht<strong>in</strong>g Publicatiefonds <strong>Lutra</strong> Hans (G.J.) Bekker, Bauke Hoekstra & Ben Verboom<br />
Information: Bornsestraat 118, NL-7601 GK Almelo, the Netherlands<br />
DUTCH MAMMAL SOCIETY<br />
Information Secretariaat Zoogdierverenig<strong>in</strong>g, Toernooiveld 1, NL-6525 ED Nijmegen, the Netherlands<br />
tel. +31 (0)24 74 10 500; fax +31 (0)24 74 10 501<br />
e-mail: lutra@zoogdierverenig<strong>in</strong>g.nl<br />
website: http://www.zoogdierverenig<strong>in</strong>g.nl<br />
English language edit<strong>in</strong>g by: textualheal<strong>in</strong>g.nl<br />
Photograph cover by: Ben Verboom<br />
Art: Peter Twisk (p. 104, soprano pipistrelle), Elias Nauta (p. 108, badger)<br />
© 2010 Zoogdierverenig<strong>in</strong>g ISSN 0024-7634
When you prepare a manuscript for <strong>Lutra</strong> please comply<br />
with the gui<strong>de</strong>l<strong>in</strong>es set out below. For additional <strong>in</strong>formation<br />
on style and format please consult a recent issue and<br />
the “How to submit a paper to <strong>Lutra</strong>” document that is<br />
available <strong>in</strong> English as well as Dutch from the <strong>in</strong>ternet<br />
(http://www.zoogdierverenig<strong>in</strong>g.nl) or Zoogdierverenig<strong>in</strong>g<br />
office (address see 18).<br />
1. <strong>Lutra</strong> publishes <strong>in</strong> British English as well as Dutch.<br />
2. Choose an <strong>in</strong>formative title for the manuscript that<br />
does not exceed 15 words.<br />
3. Use 200-250 words for the abstract of a full article.<br />
For short notes an abstract is not required, but if<br />
<strong>de</strong>sired it should be 100-150 words. Add a Dutch<br />
summary (400 words maximum for a full article, 250<br />
for a short note).<br />
4. Use up to ten different key words or short phrases<br />
that best i<strong>de</strong>ntify the manuscript.<br />
5. A manuscript should generally follow a standard<br />
scientific format. When appropriate you may subdivi<strong>de</strong><br />
the text with second or even third level head<strong>in</strong>gs.<br />
6. When you name a species for the first time <strong>in</strong> the text<br />
use the English or Dutch name, <strong>de</strong>pend<strong>in</strong>g on the language<br />
of the article, followed by the scientific name<br />
between brackets and <strong>in</strong> italics: p<strong>in</strong>e marten (Martes<br />
martes).<br />
7. When you report the f<strong>in</strong>d<strong>in</strong>gs <strong>in</strong> the results section<br />
avoid repetition from tables or figures, but do <strong>in</strong>tegrate<br />
the most important or <strong>in</strong>terest<strong>in</strong>g aspects <strong>in</strong> the text.<br />
8. Use digits for numbers from 13 and up, and use<br />
words for numbers up to twelve. If a number is part<br />
of a measurement unit use digits: 7 g or 2.5 km.<br />
9. In general, refer to tables, figures and photographs at<br />
the end of a sentence between brackets: (figure 1) or<br />
(table 1) or (photo 1). With maps use a scale bar to<br />
<strong>de</strong>pict the units of measurement or distance. For figures<br />
use simple symbols or patterns, all <strong>in</strong> black and<br />
white. Contact the editorial board if you need to have<br />
a figure pr<strong>in</strong>ted <strong>in</strong> colour. When you construct a table<br />
use tabs or the table function and use only horizontal<br />
l<strong>in</strong>es.<br />
10. Tables, figures and photographs are either one or two<br />
columns wi<strong>de</strong>. Please prepare your tables and figures<br />
<strong>in</strong> such a way that the axis titles rema<strong>in</strong> readable. Use<br />
the “times new roman” font for the axis titles, legend<br />
and possible footnote. Axis titles start with a capital<br />
letter and have no po<strong>in</strong>t at the end.<br />
11. Reference citations <strong>in</strong> the text are normally placed at<br />
the end of a sentence. Use “et al.” for three authors or<br />
more. The references are first arranged <strong>in</strong> chronological<br />
or<strong>de</strong>r and then <strong>in</strong> alphabetical or<strong>de</strong>r: (Reeve &<br />
Huijser 1999, Broekhuizen et al. 2000, Jansman &<br />
Broekhuizen 2000).<br />
12. List the references <strong>in</strong> alphabetical or<strong>de</strong>r and then by<br />
year of publication. Do not prepare your manuscript<br />
with tabs here. Please provi<strong>de</strong> the full name of a journal.<br />
For books only name the first location if a publisher<br />
has offices <strong>in</strong> more than one city. Use the full<br />
name of the country, but abbreviate United K<strong>in</strong>gdom<br />
and United States of America: UK and USA. Only<br />
refer to a site on the <strong>in</strong>ternet if you are reasonably<br />
sure the site concerned has a relatively long life span.<br />
Instructions to authors<br />
Please review the follow<strong>in</strong>g examples<br />
Barrett, G.W. & J.D. Peles (eds.) 1999. Landscape<br />
ecology of small mammals. Spr<strong>in</strong>ger, New York,<br />
USA.<br />
Bergers, P.J.M. 1997. Versnipper<strong>in</strong>g door rail<strong>in</strong>fra<br />
structuur. Een verkennen<strong>de</strong> studie. Report 262.<br />
Instituut voor Bos- en Natuuron<strong>de</strong>rzoek, Wagen<strong>in</strong>gen,<br />
The Netherlands.<br />
Clarke, G.P., P.C.L. White & S. Harris 1998. Effects<br />
of roads on badger Meles meles populations <strong>in</strong><br />
southwest England. Biological Conservation 86:<br />
117-124.<br />
Clevenger, A.P. 1998. Permeability of the Trans-<br />
Canada Highway to wildlife <strong>in</strong> Banff National<br />
Park: importance of cross<strong>in</strong>g structures and factors<br />
<strong>in</strong>fluenc<strong>in</strong>g their effectiveness. In: G.L. Ev<strong>in</strong>k, P.<br />
Garrett, D. Zeigler & J. Berry (eds.). Proceed<strong>in</strong>gs<br />
of the <strong>in</strong>ternational conference on wildlife ecology<br />
and transportation: 109-119. FL-ER-69-98. Florida<br />
Department of Transportation, Tallahassee, USA.<br />
Shkedy, Y. & B. Shalmon 1997. Evaluat<strong>in</strong>g open<br />
landscapes <strong>in</strong> the Negev Desert, and the implications<br />
on military activity. Nature Reserves<br />
Authority, Jerusalem, Israel. (In Hebrew with<br />
English summary).<br />
13. Use the entire width of the page; do not prepare your<br />
manuscript <strong>in</strong> two columns. Do not hyphenate words<br />
at the right marg<strong>in</strong> and use only left justified text.<br />
Prepare your manuscript on A4 sized pages with<br />
2.5 cm marg<strong>in</strong>s and double space between the l<strong>in</strong>es.<br />
14. The first page should show the title, authors and authors’<br />
addresses. The page that follows starts with the<br />
abstract, immediately followed by the key words and<br />
the body of the manuscript. Tables and accompany<strong>in</strong>g<br />
text should be placed at the end of the manuscript,<br />
followed by the text accompany<strong>in</strong>g figures and photographs.<br />
The figures and photographs should be on<br />
separate pages that also show the figure number and<br />
the name of the first author.<br />
15. Book reviews of Dutch publications may be written<br />
<strong>in</strong> either Dutch or English. Reviews of publications <strong>in</strong><br />
any other language should be written <strong>in</strong> English.<br />
16. We encourage you to submit your manuscript digitally<br />
(preferably <strong>in</strong> Word) on CD-Rom or through<br />
e-mail as this speeds up the review<strong>in</strong>g process. If<br />
you submit your manuscript on paper please <strong>in</strong>clu<strong>de</strong><br />
a total of three copies (laser pr<strong>in</strong>ter, figures with 600<br />
x 600 dpi). Please <strong>in</strong>clu<strong>de</strong> the telephone number,<br />
fax number and e-mail address of the correspond<strong>in</strong>g<br />
author.<br />
17. There are no page charges for <strong>Lutra</strong>, but contact the<br />
correspond<strong>in</strong>g editor for <strong>in</strong>formation concern<strong>in</strong>g the<br />
costs of colour illustrations.<br />
18. Manuscripts should be sent to the editorial board:<br />
Zoogdierverenig<strong>in</strong>g, Redactie <strong>Lutra</strong>, Tournooiveld 1,<br />
NL-6525 ED Nijmegen, The Netherlands. The e-mail<br />
address of the editorial board is: lutra@zoogdierverenig<strong>in</strong>g.nl.<br />
Close to or shortly after publication<br />
you will receive 25 repr<strong>in</strong>ts free of charge.<br />
19. The editorial board strongly advises you to have your<br />
manuscript critically reviewed by colleagues before<br />
you submit your manuscript.
LUTRA<br />
Volume Deel <strong>53</strong> - Number Nummer 2 – December 2010<br />
Contents Inhoud<br />
Editorial / Redactioneel<br />
49 The bat, the weevil and other ‘species’<br />
Ben Verboom<br />
Research Papers / Artikelen<br />
51 A comparison of the hibernation patterns of seven bat species <strong>in</strong> Estonia<br />
Terhi Wermundsen & Yrjö Siivonen<br />
63 The <strong>in</strong>fluence of treel<strong>in</strong>e structure and w<strong>in</strong>d protection on commut<strong>in</strong>g and forag<strong>in</strong>g<br />
common pipistrelles (Pipistrellus pipistrellus)<br />
Ben Verboom & Hans H. Huitema<br />
81 Badger (Meles meles) road mortality <strong>in</strong> the Netherlands: the characteristics of<br />
victims and the effects of mitigation measures<br />
Jasja J.A. Dekker & Hans (G.J.) Bekker<br />
93 The first historic record of a humpback whale (Megaptera novaeangliae) from the<br />
Low Countries (Southern Bight of the North Sea)<br />
Jan Haelters, Francis Kerckhof & Kees (C. J.) Camphuysen<br />
Short Notes / Korte Berichten<br />
101 A killer whale (Orc<strong>in</strong>us orca) <strong>in</strong> the castle: first f<strong>in</strong>d of the species <strong>in</strong> a Dutch<br />
archaeological context<br />
Jørn T. Zeiler & Erw<strong>in</strong> J.O. Kompanje<br />
105 First record of soprano pipistrelle (Pipistrellus pygmaeus Leach, 1825; Chiroptera:<br />
Vespertilionidae) <strong>in</strong> Wallonia (Belgium)<br />
Daan Dekeukeleire<br />
Book Reviews / Boekbesprek<strong>in</strong>gen<br />
109 Zoogdieren <strong>in</strong> Europa<br />
V<strong>in</strong>cent van Laar<br />
In<strong>de</strong>x<br />
112 Contents of Volume <strong>53</strong> (2010)<br />
Pr<strong>in</strong>ted by Drukkerij Wilco, Amersfoort<br />
Layout by Image Realize, Ommeren ISSN 0024-7634