Invasive alien species in Switzerland - Schweizer ...
Invasive alien species in Switzerland - Schweizer ...
Invasive alien species in Switzerland - Schweizer ...
Create successful ePaper yourself
Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.
29<br />
06<br />
> Environmental studies<br />
> Organisms<br />
> <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong><br />
<strong>in</strong> <strong>Switzerland</strong><br />
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity<br />
and economy <strong>in</strong> <strong>Switzerland</strong><br />
&%&&&&&0?0&&000&?0?/^/&0&&&%%&&&&&&&%&&&&&%&&&0???00???0//??/&&&&&&&&&&&&&0&0&&0&&&&&00?000?0??//0&&&&00&&&%0??00??000?000???&0000&&&&<br />
&&%%%&&00/?0000???//&0&&&&&&&&%&%&&&0&&&&&%&&&00???0???????0&0&&&&&&&&&&&%&&&&&0&&00000?/0???///??0&&&&0&&0000?0??000?00000?000000&0&&<br />
&00&?&0&??/?&00?/^&&0&&%%&&&&&%%%&0&00&&&&00&00????00??///&&&&&&%&&%%&&&&&&&&&0&&&00?0&0/000?////?0&&&0???????0&&&&000&%%%&0&000000000<br />
&&&&0&00???/?//^?0&&&&%%&%&&&&&&&%0&&0&&&&&0?00???/&??//&&0&&&&&&&&&&&&&&&&&&&0?&&0??000??0??/////0???//?0&0&&000&&&00000&000??0000000<br />
0&&0000&00?&//&0&&&&&&&%&%&&&%&&&&&0&0??0&000?0/?0??//&00&&&&&&&&&&&&&%&&%&&&&0???000??00/???/?//?&&00&&&00000&000&&&00&000?0?/??00000<br />
&&0000&0000?%&0&&&&%&%%&0&%%&&%0&&&0&&0??/0&&?????//%&&&&&%&&&&&&&%&&0&&@@%&&00?/?00??/??0/?////0&&&%&&&&00000&&00&&&&&&&0000??????0??<br />
&&0&?&00&?/%?&&&&&%&@%%%&&%&&&&0&&&0?0&?0?000???//?&&&&&&&&%&&%&&&&&&&0&&&&&000?//0???/?00?///?000&&&&&&&&00?&&&&&&&&&&&&00000??/??0?0<br />
&0&00&&??^&0&%%%&&&&%%%&&0&&&&0&000000&??0?????/?&0&&%&0&&%&%%%&%&&&&&0&&&&00?0????0??///?//?00000&&%&&&0&00?00&&&00&&&&&&0?0???//?&00<br />
&&&00?///%0&&%%&&%&&&&&&&&&&%&00&??000?/?//??^^00&&&%&^,““›&%%&&&&0&0&0&&&&0&0??///???/?//?&&@%0000&&&&&&0&&0?000000?&&&&&00?0???//0&0<br />
&0?&0/?/&0&&&%&%%&&%&&&&&&0&&0&0&00??0????//^&0&0&&%&%000/›;&&&&//;&&&0?0&000???////??//?&&&@@&&?000&%&%&&000??&0&000000&&0????////000<br />
&00???/%0%%%%%%%&&%%%%&&&&&?&0&??0&&?//&0?^%0&&&&%&&&&&&›››^^&%/›“/&&00?&&&0???//??/?/?&00&&&&&?00000&&%%&&000??&&00000&&&00?/??///?0&<br />
0?0?/?&&&0&%&%%%&&@&&&&&%%&?&0%0&??/?///^/&0&&%%&%&%&,;;“““%&››;›/?&000???&???^/////?&00&&0&&&00??0000&&%&&0000?&00000?0&&&0?//?/^/?0&<br />
??&/?00&&%&&0&&%&&&%%&&00&00000&?????/^0&0&&&&&&&&&&›“^››^,/;/&@^;^?^;&@???????//^/&&&&&&&&&0&&&000&0&&&&&&&&00?0&0000??000?0//?/^/?00<br />
0?/^0&&%&&@%%&&&%%&&%&0&%00%/0?0&0?//^0?&%&&&&&&%%&?/“““““;,/““^/%^››^?0?/////?//0&&&&&&&&&&&000?000&&&&%%&&&&0??00000???000?//?///?0?<br />
//?00&&&%&&%%&&0&%&&0&0&%0?//?0/&?/^^&0&%%&&&%&&&%%??,““““““^^“““^@@,@%@?/?/?//?&&&%%&0&&&&&0&&0&0?000&0&%%&&&0???0&&00???00///////?00<br />
^0?&&%&&&@%%%?&00&&&&?0???0?0?/^/^^?0&&&%%&%%&%&&&&?/?;“““^“““““““›0&›^^@?@?^/&00&&&&&&00&&&&&&00000000?&&&&&&&????0&0?0?/?0?////??&&0<br />
%/&%&%@@%%%%&0&&&%0?0???///???//^?&0&&&&%&&%%&%&&&;““^/›“,“›@“““,;,@0;^;“›//&0&&&00&&&&&&0&&&%&&00&000000&%%&00?0?/????0?//??//?&00000<br />
?0&&&%@%@%&&%&&&&?0%0&?0/??0?//^%?&%&&%%&&&%&&%@&0&/^,^““/›““““““/@@0,““^,,@@%&&%&&0&&&&&&00&&&&&0&&&0&???&&&&&00?/?????/?///?00000000<br />
&%&&&%%%&&&0&&?0?&&&0&?/0/?&&^^%0&0%%%%%%%%%&@%%&&&00&?^››/›“““@““@@@^?,“,““%&%&&%&%&0&&&&&0&&&&&0000000??0&&&00??/?????//?&00000??000<br />
@%%%&&%&&%&&%&&?0??&0/^//?0^/^00&%&&&&%&&%&&&&%%@%00&&&??^,“““““““^?@@?“““^,;%%&&&&&&&&&&&&0&&&&&&&00000&??&&&&?????/?/?&&000?00000?0?<br />
@%%@%&&&0%&&0%0&?&?&^??/?&//%?&&%%&%&&%%%%%@&&%%&%&&&0%%?/“““›,““““““@@“&““;“,%0&&&&&&&&%&&&0&0&&&00000000?0000?/???/&&&&000???00000??<br />
&%@%%&&&&0%&0%%&&?0?//^0?/^%0&@%%%%@&&%%0&%%@&%%%%0&@%&&/^““/››“““““,“&?@,““;,›@&0&&&&&&&&&&&0&&&&???0??0??????//^/&&&&&0???/?/00000//<br />
&&&%%%&0&00&&&??0//?///?/^?0&%@%%%%%&%&&%%@%%&&0&&?0&%00/›^/›^››“““?“““›@?%“›@“››//0&&&0&&0000?00?000?0????/?//^0&&&0&&&0?//?//??00??/<br />
@%@%@%&&000%&0?0/0^^?/^^%0&0&&&&&%%%@@%&&%&&%%&%&&&00&?0/0/^^?››““/,“““,“0@,“,“;,“??&%%&0&&0&??00?00???????//000000000000/?/?//^??0?//<br />
@&&%%@@&&?/&%???0//??/0&0&%%&%%%%%&@%@@&&%@&%@%&@0/0&?///?/?^^^;/,//^“““““@%““““;,;0&0&&0000?00??/????????^?&&&000000??????//////???//<br />
&&@&@%0&&?0/?&/&0?^/^?&&&&%%@&&%%%&&%%%&%%%@@%&@%%/%&??^^///^›0???//?^,“““““““,““““?/0&000?00??????//?//^?0&&@&?000???/?0???/??///?/?/<br />
@%%%&0&0&000?^/?^/^^&?&%@%%%@%%&@@&%@%&&0&@%&&&0&0?&?0/////^›%??0&0/?^^^;““““““““““??00/00????//?/??/^^0&&&&&0&0000????/???/////^?/?//<br />
&%&%&0%0?&00///@/^^@?&@@@@%%%@&%&@%&@@%&?%&%&0&0??^///?//^^^@?0&&&&@%/^^^“›“““““““›?00?????/?/?/^/?^^&&0&&&&&&&??0??0?///??///???/?//?<br />
0?&%%&&00&//?0//^/%&00&%@@@%@@%%&&&%%%@%%&&&00/^?????///^^%0000&%%&000?/^›››“,““›/^///?0/??////?/^/&%%&&0&&&%&%0??&??///^?0/??/?/////?<br />
%0??&&0?00^/?/0/%?&&&&&&%@&@@%%%%%&&%%0&&?/?&^^›//^^?^^^@?&&%&0%&0&&000?^/›^^›^,›^/???/?/^/?^/^^^&0&&&&&00&&@0&&00/???0//?????////////<br />
%%00///&/??/0/^%?0%%@%@&%0&%&%%%@%%&0%&?&??^//^0?%00^?@0&0&&%&0&&&000&&?^/^^//////?///?/?^^?/^/&0&0&&&&&000&%%%0&?^0?///^/??//??//?///<br />
&&&&?0/^?///^^&?0&%&%@@@@%&&%&%&&&%&?0/??^^&?00??&/^0&&%%&&%&&&&%&0000?&??0/?0?/?/??0/^^?^/^^%&@@%&0&&000????&0/?//0//^////??/////////<br />
0??0??0/0////&0%%@@%&%&%%%&%%&&&&%&&?0/?/?//?&&//^^%0&%%%@&0&&%&0??&&?0?00?//0?/??///^//^^^&0&@@%&0&00&&00????//?/^//^?^^^/?/////^//^/<br />
0&?//^^/^/^00&%&@@@@@%@&@&&&&00?&0%////0^?^^^/0&^000&%@&%%@&&%&0?0000?/?0?&?////?/^?///^^%000&&%&&&&?0&0?0?/&??/?/^//^/^/^^?/////^//^^<br />
/0?&?//?^^%000%&%@%@@&@&@%@&%00&??/?0//^&^^?^&^^&00&&&&%&%&%000/00?0^????&0/?///^^?^//^?0&%&%%0&&&0&00&&000/0???///^//^//////////^///^<br />
&&0&^//^^0?&&00%%%%%@@&%%%&&%0????^0/›0/^/^?^/@0&&&&&&%%%&%&&&0&?????00?&??/??///0//^0%?0&&%&@&000&0?&????//?////^/^?/0//^^/?/^^?^//^^<br />
^^?&?0^^?0&%%%%%&@@@&?0&&??%0^/&?&/?^^?^^^^^&0&0%&%%&&00&00&?&??00??/?/?^?/??/^///^^%&&&&%%%&&0&&0??&??&//^??//^^/^^/^/0/^^^//^^/^^^^/<br />
%^??/^%??&%%@@%@%@%%%0&&&&?0/^/^/^?/^^^^^//&&0%&&&&%&%%@0&0&?00&00&/&?/?0/^/?^^/^^%00&&@@&&&&00&&?&0000??/////^?^^^^/////^^^//^^^^0&00<br />
^0^//@?&%0%%%&@@%&&0%&@&&//////^//^0^/?^^&00%%%%0%%&%&%&&&?0000?0?/0??//^^/^/^/^/&0000&%%%%0&&00000%&?/?/^?////^^^^^///^^//^//^^/000??<br />
/&^^@?%%%%%@@@%@/&0&?&?&&%^^0?^^/^^/^››?000&%%@@0&00&&%%&%/0?/00//???^//^/?^^^^@&00%@&&&&%%&?0????0&&0?//////^^/^^^^//^/0?0^^/000000??<br />
^^?00?&&@%&@@&@&&??&?/?^^/^?^//?››^/›?%00%@@@%%%%&&%&&&&&0000?/&//??//›^/&^^^&?&&&&&@0&&&0&000?&?/0/^/?/?^^^^/^^^^^^^///?/^&&000?00??/<br />
^&/&&&00&&%%0?%&&00&^/^?^//›^&///^^›@0&0%%@@@%&&&&&%0&&000?/?????//?^^/^^^^^000&&%%0&&&&%@^&?/0///?;›/?/?^^//^^^^//^^/^^?&000000????/^<br />
&0&@@&&0@%&&&&?&/^&&^/??^^›?^/^^^^&&0&&%0%@@%&&@&0&&&??&0/0^/?0/??//?^^/^/@00&&&&&%00^00%/?“/??//?^“›?//^^//^^^^^^^^^?&0&00?00?0????/^<br />
0@0%%%0?000/?0&/%/?0//??^0/^?//››&00%%%%%&@&&&00?00&&&00?&/0^0/^^›^///^^?0&00&&&%0????“,?0&/;“?//›““^›;^^^/^/^^^^^^?&00000?000???/?//^<br />
%0&&0&&?0&?0??//^^^//^0^/^›/^^›@00&&%%%%&&&@&&000????%???/0/?^?^^^///^^@00000&&&&&&&0/^^^;^›/“,%““;^%;“›^^^^^^^^^?&000??&00??????//&?^<br />
&&/&&&0&?00?00/0/&00?/›?^?^^›/000%0%%%&&&&@&@@&0%&0%?00?/?//??//^^0^›?0@0&?&&&&?&0?%0??0/››%““,“›^››%0^“›^?/^^^/00000?00%&&???????//?/<br />
&?&?&%?00???^//^&//?/^^^›^››@?&%%%@%&%0@0&@@%“;@&^??/00&?^??^^^?/^^^&0&0&00?&%&&&&000&,??@@@@&&,›/^“0^%,;^/^^00000&&0?0&&&0??/?0?/^/^^<br />
@@&&&/&0??//^???/^/›^^//^›@?00%%@@/%@@@@?%@@&›@@@@@%?/0?//^/^^^^^?%000&0&&0&?&&&00&›0/&@@@@@/@%“/›0@@^/“›^^^&0&%000&&0?0?0?0?/??///^^^<br />
/?0?&??//?^0??0^/^^^^?^›/&?&&%@&%0/^“;0›;@@@0›?@@@@@%›&//^///^/^00&0&?&0%&&?0???/&?0@@@%@›^,^““&@@/0“;“^“›&?00000???0?0?000?////?^^^//<br />
/&%&?00????00?//^^0/0^^@?0&%&&@&%@?@^,@@/@@@@@@@@@&@@@%%/?^/^››%0&0&&&0&%%?0?^?/^@@@@@;›^/““,0,;,“,^“,““?/000&?0?&0?0?0/00??/?/^/^^^^^<br />
/&&///?///?0^^›^0&%%^›%00&@@@0@?,&?““@@@?;^^@@@@&@?“@@@%%“?//0&00&%&%%%0&&&&/›;@@@@^›?›““,?›““,?“““““““^,^&0000?00&/??0????///^^//^^^^<br />
/&/&????//?^&?0&?/?^%0?0%@@@&/&&››,““%@/%&@@@&›%&@@›%@@@“/0›??&&&&%&&%00&^,›““^“^^?“&“/^;“,;““&““,“““0^››,;0&&?0/0?//0?????//^^/^^^^^^<br />
^00?/0?/0^//0/%0^?›@0&0%%@%0&0/^›@@@“““&%@@@@@@%%@@%&@@%““&&&00%%&%@@››;/;“››““^““,^“,,›““,/“&›““““,^“^//^0&000??/?????0////^^^^^^^^^›<br />
›^^›^/??&^^^&?/^›@&?%%&&??&/?^^/^@@@@;/&%›“%%%%“““&%%?%&&“&0?%&&%?&^““,,,^›^^;;;›,“;“&“›?;@^“@““““?;^//?0??00??////??/^?/^^^/^^^^^^^››<br />
@%^&0?/^›››››››/@&??/?/^^?/^›^^0?›@@?/%&/“““&%&;“““%%@%%^&&0?&&&%&&›“““““,“,›“““;“;“/“%^?@“,“““““?^?0000/^??00?///???/??//^/^^^^^^^›^›<br />
00&0??&&@@0&&%/@@%&&?/^››/››››/^^^“››/^&&??00&&&/&&&&&››??›?/000?0&/;“““““““““““,“““/%“,“?“““,““;^?0000&^^/??/////^?^///?/^^^^^^^/^›^›<br />
&&&&%&0&&&&@%00???????0@@@%%&0/››;,“››““““??????000&0,“““/^^^0%///?//››“;““““““““““;“^“““““,,“0^/0&&&000^^/?0//??//?/^//^^^^^^^^^^›/^›<br />
%00&&&&&&&&0@&&&&0&&&&%&?0&00?^/&&^;“›“““,›^^//^›////““,/^^;^&?^//^/^›^›?›“““,›“““,“,,,››,^›00^0000?/?&??/0^//?//^//^^/^^^^^/^^^›^›^^?<br />
%%&%@@%&&&&0&@?%&%&&@%&%&%&&&&0^?//^;“›,“,,,,;;““^^^›››››;^““^^/?/&/››^//^^››,^“›;;›^0^›^^›0/?&@@&??//??^?^^^/^^^/^///^/^^^^^^^^›››^??<br />
%&%@%%%%&%&&&?0/&@@@@%%%&&@@@@%?&?^^/›““^,“““““,,,,“““,,›;/0^›^››^›^^^^^^^^0/›^^^^^0&/^›^^%?????0??^^//^^^^^^^//^^^/^^^^^^^/^^››››//??<br />
&&%@%%%%%%&%&&%?/%%@@@@@%%@@@@@%@&0?//^›““““““““““““““““›››/?^?%%%&&?0^››^›^›^^^^^%0?^››%?/?/@0???/^?^^^///^/^/^^^^/^^^^^^^^^^›››?/???<br />
@%%%%&@%&&%&&0?@?%@@%@@@@@@@%@&0@@&%0&0/^^;“/“““““““““;^?/&%&&00??00?///&@@%&%/^›^››^^^%?////?&&/&^?^///^/›^^^^/^^//^^^^^^^^^^››???/??<br />
%@@%@@%&&@@@&&0?%^?&@&&@@%%@@@@@%@@@@%%&0^?/^&/%%@,0?^?0&0/0&&0&?&0&&@@@00?00/???&%&%%&?/^/^/^^&^^?^/^0//^/^^^^^^^^^^^^^^^^^^^^????/?/<br />
%%@%%%0%@@@@&0&&0/0@&&%@@@%@0%%%%%&%%&&%%%0??›&^@@@@&@%&&&0/?&0?&%%0%0%%&%%&0//&0???/&@%@%&0/^›››^^^^^^^^^///^/^›/^^^^^/^^^^^›????///?<br />
%@&0%%%&@%%&%&&&0%^??%%%@%%%&&&@%@@%&&@@%&@%0?^@^0?&^^&&&%%^???&^/%&@?000&0&000?%00^/0?^%?0????0&%%&00/›^/^^^^^^/^^^?^^^^^^^^/????////<br />
&0&@%@%%@@@%@%0&?&@^??&&%&%&&&%@@&&%&&@%%%%&0/?/%^00&0&%%&%%&&0&??&&%@&&0%0?000/??00/?00?/?///0??????0????0&&&??/››^^^/^^^^^&//?/////?<br />
&%%%@%&@%@%&&&0?&&/%^&&&&%%&&%&&&&&0%%%%&%&&&^0&?%^?%?%0/0?^?@@@?^^0&&0&&&?%0&?//??&?0??/&//?////??%/0????@??00???&&&&&0???&//?////?//<br />
%&%%%%%@%%%&%0&&&&0??//&%??&%&%@%&&&%%%%&&%0/00%&^%^/?00^^^/^›^?^^^/%&&&&0?&?00/0??00&0?/^0000?0//??^??//&&/?00/%????00?0?0&&0?//////^<br />
@%%%%%&%&%&%&&%%%&&???0??&@@@%&%%00&&&&&&&&?&0%%0?/%?//0//&/??%?/?^/?/0???00?&??0?/%@?0&^?/?00//^/?0/^???0?^?/?^/??/^?/^/???//??0&&?^^<br />
00&&%&%&%%@&%&@%000?0?%/?0?%&0@%&&%%&&00&&?%%&?&0???0/^/?/?/?%//??/&?0&0?/00^?&/0?0&%&??&/?^?///?0/??^/0/0?^?0/^^/?/^///??//?0^?//???0<br />
&&&%%%%&%%%&%%&&&0?&&??/^%?0@&?/&&0&?%%%&?&&@&&/0/0/0?^?/^^?//0^00/?&?0?&^^/0?^^@?0@&0???//^^?/////?/0?////^0?/^^///^?/?&?^//^/?&&0/^/<br />
%&%%%@%%%&0%%@&%?0&&&0?%^0?&&/&&&&%%%&%00&&&&%0&0???&?^0^^/&?^/0/^/^?/?/?0&0/^0?0%%@0???&&0&^^/00?^0?/^^?//?^/^^//?/^//???^^////^0&/^/<br />
&&%%%%&%%0%%%&&000&&&%0?%^^//&@&&0&&&&&?%&0%&&00&/0%@00/%/?^/?^^^//^???^0?00?/????%&0?0?&0%&^^^^/0///^/??/?^^0//^^/^^/^?^/^^////?/0?^^<br />
&%@%&%%&%%&&%&&?&%&0&00/0%/?@0?0&&&0??/&@&&0&&0?/?00?0??/&^^?0^/^^^/^//?^??^0/?^???%0&0?/??%?^^?//?/^?^^^//^//^^^//^^^^?/^^^^////^??^/<br />
?%&0%&&&&%%&0?0&&%&&&0&0%?&^/?&0&%000?&&&&0&00??^0&?0?0???&^/^??^^?^/^//?//&^^//??0&???0?0/?%^^^?//^?^^^//^////^^^^^^/^/^^^/^??////?^?
Environmental studies > Organisms<br />
><br />
<strong>Invasive</strong> <strong>alien</strong> <strong>species</strong><br />
<strong>in</strong> <strong>Switzerland</strong><br />
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity<br />
and economy <strong>in</strong> <strong>Switzerland</strong><br />
Mit deutscher Zusammenfassung – Avec résumé en français<br />
Published by the Federal Office for the Environment FOEN<br />
Bern, 2006
Impressum<br />
Editor<br />
Federal Office for the Environment (FOEN)<br />
FOEN is an office of the Federal Department of Environment,<br />
Transport, Energy and Communications (DETEC).<br />
Authors<br />
Rüdiger Wittenberg, CABI Bioscience <strong>Switzerland</strong> Centre,<br />
CH–2800 Delémont<br />
Marc Kenis, CABI Bioscience <strong>Switzerland</strong> Centre, CH–2800 Delémont<br />
Theo Blick, D–95503 Hummeltal<br />
Ambros Hänggi, Naturhistorisches Museum, CH–4001 Basel<br />
André Gassmann, CABI Bioscience <strong>Switzerland</strong> Centre,<br />
CH–2800 Delémont<br />
Ewald Weber, Geobotanical Institute, Swiss Federal Institute of<br />
Technology, CH–8044 Zürich<br />
FOEN consultant<br />
Hans Hosbach, Head of Section, Section Biotechnology<br />
Suggested form of citation<br />
Wittenberg, R. (ed.) (2005) An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their<br />
threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong>. CABI Bioscience<br />
<strong>Switzerland</strong> Centre report to the Swiss Agency for Environment,<br />
Forests and Landscape. The environment <strong>in</strong> practice no. 0629.<br />
Federal Office for the Environment, Bern. 155 pp.<br />
Design<br />
Ursula Nöthiger-Koch, 4813 Uerkheim<br />
Fact sheets<br />
The fact sheets are available at<br />
www.environment-switzerland.ch/uw-0629-e<br />
Pictures<br />
Cover picture:<br />
Harmonia axyridis<br />
Photo Marc Kenis, CABI Bioscience, Delémont.<br />
Orders<br />
FOEN<br />
Documentation<br />
CH-3003 Bern<br />
Fax +41 (0)31 324 02 16<br />
docu@bafu.adm<strong>in</strong>.ch<br />
www.environment-switzerland.ch/uw-0629-e<br />
Order number and price:<br />
UW-0629-E / CHF 20.– (<strong>in</strong>cl. VAT)<br />
© FOEN 2006
Table of contents 3<br />
Table of contents<br />
Abstracts 5<br />
Vorwort 7<br />
Summary 8<br />
Zusammenfassung 12<br />
Résumé 17<br />
1 Introduction 22<br />
1.1 Def<strong>in</strong>itions 23<br />
1.2 <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong> – a global overview 24<br />
1.3 Status of <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> 28<br />
1.4 Pathways 28<br />
1.5 Impacts of <strong>in</strong>vasive <strong>alien</strong> <strong>species</strong> 29<br />
1.6 Discussion 31<br />
1.7 Recommendations 32<br />
1.8 Acknowledgements 34<br />
2 Vertebrates – Vertebrata 36<br />
2.1 Mammals – Mammalia 36<br />
2.2 Birds – Aves 44<br />
2.3 Reptiles – Reptilia 52<br />
2.4 Amphibians – Amphibia 54<br />
2.5 Fish – Pisces 55<br />
3 Crustaceans – Crustacea 65<br />
4 Insects – Insecta 71<br />
4.1 Introduction 71<br />
4.2 Coleoptera 73<br />
4.3 Lepidoptera 75<br />
4.4 Hymenoptera 77<br />
4.5 Diptera 79<br />
4.6 Hemiptera 80<br />
4.7 Orthoptera 82<br />
4.8 Dictyoptera 83<br />
4.9 Isoptera 83<br />
4.10 Thysanoptera 83<br />
4.11 Psocoptera 84<br />
4.12 Ectoparasites 84<br />
5 Spiders and Allies – Arachnida 101<br />
5.1 Introduction 101<br />
5.2 List of <strong>species</strong> 102<br />
5.3 Species of natural habitats 103<br />
5.4 Species <strong>in</strong>, and <strong>in</strong> close proximity to,<br />
human build<strong>in</strong>gs 106<br />
5.5 Greenhouse-<strong>in</strong>habit<strong>in</strong>g <strong>species</strong> 108<br />
5.6 «Banana spiders» and terrarium <strong>species</strong> 109<br />
5.7 Discussion and recommendations 109<br />
6 Molluscs – Mollusca 113<br />
6.1 Snails and slugs (Gastropoda) 113<br />
6.2 Bivalves (Bivalvia) 115<br />
7 Other selected <strong>in</strong>vertebrate groups 121<br />
7.1 Nematodes – Nemathelm<strong>in</strong>thes 121<br />
7.2 Flatworms – Turbellaria, Plathelm<strong>in</strong>thes 122<br />
7.3 Segmented worms – Annelida 122<br />
7.4 Centipedes and millipedes – Myriapoda 123<br />
8 Lichens (Lichen-form<strong>in</strong>g fungi) 124<br />
9 Fungi and a selected bacterium 125<br />
10 Plants – Planta 128<br />
10.1 Introduction and term<strong>in</strong>ology 128<br />
10.2 The native and <strong>alien</strong> flora of <strong>Switzerland</strong> 130<br />
10.3 The geographic orig<strong>in</strong> of <strong>alien</strong><br />
and naturalized <strong>species</strong> 131<br />
10.4 Status of the <strong>alien</strong> <strong>species</strong> of <strong>Switzerland</strong> 133<br />
10.5 Naturalized <strong>species</strong> of <strong>Switzerland</strong> 134<br />
10.6 Life form 135<br />
10.7 The habitats of <strong>alien</strong> plants <strong>in</strong> <strong>Switzerland</strong> 136<br />
10.8 <strong>Invasive</strong> plant <strong>species</strong> <strong>in</strong> Europe 138<br />
10.9 Discussion 139<br />
Fact sheets 155
Abstracts 5<br />
> Abstracts<br />
Globalization <strong>in</strong>creases trade, travel and transport and is lead<strong>in</strong>g to an unprecedented<br />
homogenization of the world’s biota by transport and subsequent establishment of<br />
organisms beyond their natural barriers. Some of these <strong>alien</strong> <strong>species</strong> become <strong>in</strong>vasive<br />
and pose threats to the environment and human economics and health. This report on<br />
<strong>alien</strong> biota <strong>in</strong> <strong>Switzerland</strong> lists about 800 established <strong>alien</strong> <strong>species</strong> and characterises<br />
107 <strong>in</strong>vasive <strong>alien</strong> <strong>species</strong> (IAS) <strong>in</strong> Fact Sheets: five mammals, four birds, one reptile,<br />
three amphibians, seven fish, four molluscs, 16 <strong>in</strong>sects, six crustaceans, three spiders,<br />
two «worms», seven fungi, one bacteria, and 48 plants. A general chapter expla<strong>in</strong>s<br />
some common patterns <strong>in</strong> pathways, impacts and management, and gives recommendations<br />
for the management of <strong>alien</strong> <strong>species</strong>. The ma<strong>in</strong> body of the report is organised<br />
<strong>in</strong>to taxonomic groups, and <strong>in</strong>cludes an overview, lists of <strong>alien</strong> <strong>species</strong>, Fact Sheets on<br />
the <strong>in</strong>vasive <strong>species</strong>, and an evaluation of the status, impacts, pathways, control options<br />
and recommendations. The Fact Sheets summarize <strong>in</strong>formation on the <strong>in</strong>vasive <strong>species</strong><br />
under the head<strong>in</strong>gs taxonomy, description, ecology, orig<strong>in</strong>, <strong>in</strong>troduction, distribution,<br />
impact, management and references.<br />
Mit der zunehmenden Globalisierung nimmt auch der Handel, Verkehr und das Reisen<br />
zu und führt zu e<strong>in</strong>er noch nie dagewesenen Homogenisierung der Biodiversität;<br />
Organismen werden über die natürlichen Grenzen h<strong>in</strong>aus transportiert. E<strong>in</strong>ige dieser<br />
Neuankömml<strong>in</strong>ge können sich etablieren, und wiederum e<strong>in</strong>ige von diesen werden<br />
<strong>in</strong>vasiv und bedrohen die e<strong>in</strong>heimische Vielfalt, richten wirtschaftlichen Schaden an<br />
oder schädigen die menschliche Gesundheit. Dieser Bericht über die gebietsfremden<br />
Arten der Schweiz listet über 800 etablierte gebietsfremde Arten auf und stellt die 107<br />
Problemarten <strong>in</strong> Datenblättern vor: fünf Säugetiere, vier Vögel, e<strong>in</strong> Reptil, drei Amphibien,<br />
sieben Fische, vier Weichtiere, 16 Insekten, sechs Krebstiere, drei Sp<strong>in</strong>nen, zwei<br />
«Würmer», sieben Pilze, e<strong>in</strong> Bakterium und 48 Pflanzen. Das erste Kapitel erläutert<br />
e<strong>in</strong>ige allgeme<strong>in</strong>e E<strong>in</strong>führungswege, negative E<strong>in</strong>flüsse und Gegenmassnahmen und<br />
gibt Vorschläge für den Umgang mit gebietsfremden Arten. Der Hauptteil besteht aus<br />
den Kapiteln zu den e<strong>in</strong>zelnen taxonomischen Gruppen. Die Listen werden begleitet<br />
durch e<strong>in</strong>en erläuternden Text, die Datenblätter stellen die Problemarten vor und<br />
schliesslich wird e<strong>in</strong>e Auswertung der Situation, der Auswirkungen, der E<strong>in</strong>führungswege,<br />
mögliche Gegensteuerungsmassnahmen und Empfehlungen zu den jeweiligen<br />
taxonomischen Gruppen gegeben. Die Datenblätter bieten Information zu Taxonomie,<br />
Beschreibung, Ökologie, Herkunft, E<strong>in</strong>führungswege, Verbreitung, Auswirkungen,<br />
Ansätze zur Gegensteuerung und e<strong>in</strong> Literaturverzeichnis.<br />
Keywords:<br />
harmful organisms,<br />
<strong>alien</strong> <strong>species</strong>,<br />
<strong>in</strong>vasive <strong>species</strong>,<br />
biodiversity<br />
Stichwörter:<br />
Schadorganismen,<br />
gebietsfremde Organismen,<br />
<strong>in</strong>vasive Organismen,<br />
Biodiversität,<br />
Neobiota,<br />
Neophyten,<br />
Neozooa
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 6<br />
La mondialisation implique une augmentation du commerce et des transports et en-<br />
traîne une uniformisation sans précédant des biomes par le transfert et l’implantation<br />
des organismes vivants au delà de leurs barrières naturelles. Certa<strong>in</strong>es de ces espèces<br />
exotiques deviennent envahissantes et représentent une menace pour l’environnement,<br />
l’économie et la santé publique. Ce rapport sur les organismes biologiques exotiques en<br />
Suisse <strong>in</strong>ventorie environ 800 espèces non-<strong>in</strong>digènes établies dans le pays et détaille<br />
107 espèces envahissantes sous forme de fiches d’<strong>in</strong>formation : c<strong>in</strong>q mammifères,<br />
quatre oiseaux, un reptile, trois amphibiens, sept poissons, quatre mollusques, 16<br />
<strong>in</strong>sectes, six crustacés, trois araignées, deux « vers », sept champignons, une bactérie et<br />
48 plantes. Un chapitre général explique les modes d’<strong>in</strong>troduction pr<strong>in</strong>cipaux des<br />
espèces exotiques et leur impact sur le milieu. Il donne également des recommandations<br />
sur la gestion et la lutte contre les organismes envahissants. Le corps pr<strong>in</strong>cipal du<br />
rapport est présenté par groupe taxonomique pour chacun desquels sont proposés une<br />
discussion générale, la liste des espèces non-<strong>in</strong>digènes, les fiches d’<strong>in</strong>formation sur les<br />
pr<strong>in</strong>cipales espèces envahissantes et une évaluation du statut, de l’impact, des modes<br />
d’<strong>in</strong>troduction, les méthodes de lutte et des recommandations générales. Les fiches<br />
résument pour des espèces particulièrement envahissantes ou potentiellement dangereuses<br />
les <strong>in</strong>formations sur la taxonomie, la description, l’écologie, l’orig<strong>in</strong>e, l’<strong>in</strong>troduction<br />
en Suisse et en Europe, la distribution, l’impact, la gestion et les références<br />
bibliographiques.<br />
La crescente globalizzazione implica un aumento del commercio, dei viaggi e dei<br />
trasporti e determ<strong>in</strong>a un’omogeneizzazione senza precedenti della biodiversità a seguito<br />
del trasferimento e del successivo <strong>in</strong>sediamento di organismi viventi oltre le loro<br />
barriere naturali. Alcune di queste specie <strong>alien</strong>e diventano <strong>in</strong>vasive, m<strong>in</strong>acciano la<br />
biodiversità locale, causano danni economici o sono nocive per l’uomo. Il presente<br />
rapporto elenca le oltre 800 specie <strong>alien</strong>e presenti <strong>in</strong> Svizzera e propone delle schede<br />
<strong>in</strong>formative per le 107 specie diventate <strong>in</strong>vasive. Si tratta di c<strong>in</strong>que mammiferi, quattro<br />
uccelli, un rettile, tre anfibi, sette pesci, quattro molluschi, 16 <strong>in</strong>setti, sei crostacei, tre<br />
aracnidi, due «vermi», sette funghi, un batterio e 48 piante. Il primo capitolo illustra<br />
alcune delle vie di penetrazione più comuni di tali specie nonché il loro impatto negativo<br />
sul nostro ambiente. Inoltre, propone possibili contromisure e raccomandazioni per<br />
la gestione delle specie <strong>alien</strong>e. La parte centrale del rapporto è suddivisa per gruppi<br />
tassonomici. Le liste sono corredate di un testo esplicativo, mentre le schede trattano le<br />
specie problematiche. Inf<strong>in</strong>e, il rapporto presenta una valutazione della situazione,<br />
dell’impatto e delle vie di penetrazione, alcune contromisure e delle raccomandazioni<br />
concernenti i s<strong>in</strong>goli gruppi tassonomici. Le schede contengono <strong>in</strong>formazioni relative a<br />
tassonomia, descrizione, ecologia, provenienza, vie di penetrazione, diffusione, impatto,<br />
eventuali misure di gestione e <strong>in</strong>dicazioni bibliografiche.<br />
Mots-clés :<br />
organismes nuisibles,<br />
organismes exotique,<br />
organismes envahissants,<br />
diversité biologique,<br />
néophytes,<br />
animaux envahissants,<br />
plantes envahissantes<br />
Parole chiave:<br />
organismi nocivi,<br />
organismi allogeni,<br />
organismi <strong>in</strong>vasivi,<br />
biodiversità,<br />
neofite,<br />
animale <strong>in</strong>vasivi,<br />
piante <strong>in</strong>vasive
Vorwort 7<br />
> Vorwort<br />
Die weitgehend durch Klima und Geologie bestimmte Verteilung der Tier- und Pflanzenarten<br />
auf der Erde wurde lange Zeit durch natürliche Barrieren, wie Meere, Gebirge,<br />
Wüsten und Flüsse, aufrechterhalten. Mit der Überw<strong>in</strong>dung dieser Barrieren durch<br />
den Menschen ist, namentlich <strong>in</strong> den letzten hundert Jahren durch zunehmenden Handel<br />
und Tourismus, e<strong>in</strong>e neue Situation entstanden. Die Erde ist kle<strong>in</strong> geworden.<br />
Der Mensch reiste und reist aber nicht alle<strong>in</strong>e. Im «Gepäck» hat er – beabsichtigt oder<br />
unbeabsichtigt – Pflanzen- und Tierarten mitgeschleppt, von denen e<strong>in</strong>ige <strong>in</strong> der neuen<br />
Heimat zu massiven Problemen geführt haben. Bekannte Beispiele s<strong>in</strong>d die Ziegen auf<br />
den Galapagos- Inseln oder die Ratten und Katzen <strong>in</strong> Neuseeland, die zum Aussterben<br />
von Arten geführt haben, die e<strong>in</strong>malig auf der Welt waren.<br />
Im Gegensatz zu Inseln, die mit ihren spezifisch angepassten Arten e<strong>in</strong>zigartige Ökosysteme<br />
darstellen, ist Europa bislang weitgehend verschont geblieben von Problemen<br />
mit gebietsfremden Arten. Über die Ursachen wird spekuliert. Es kann daran liegen,<br />
dass Europa nie E<strong>in</strong>wanderungen erlebt hat wie Nord-Amerika oder Austr<strong>alien</strong>, wo die<br />
neuen Siedler mit ihren mitgebrachten Haustieren und Nutzpflanzen e<strong>in</strong>en massiven<br />
E<strong>in</strong>fluss auf die vorhandene Flora und Fauna ausgeübt haben. Vielleicht s<strong>in</strong>d aber auch<br />
unsere Ökosysteme robuster, so dass neue Arten es schwerer gehabt haben, Fuss zu<br />
fassen und die e<strong>in</strong>heimischen Arten zu verdrängen.<br />
Allerd<strong>in</strong>gs mehren sich auch <strong>in</strong> Europa und bei uns <strong>in</strong> der Schweiz heute die Anzeichen<br />
für Invasionen: Kanadische Goldrute, Riesenbärenklau und Ambrosia s<strong>in</strong>d Beispiele<br />
aus dem Pflanzenreich, die aktuell durch die Tagespresse gehen. Aus dem<br />
Tierreich s<strong>in</strong>d es das Grauhörnchen, die Schwarzkopfruderente oder der Amerikanische<br />
Flusskrebs, die den Naturschützern und Behörden zunehmend Kopfzerbrechen<br />
bereiten. Und selbst Insekten fallen vermehrt negativ auf, z.B. der Maiswurzelbohrer<br />
oder der Asiatische Marienkäfer, die unsere Nutzpflanzen direkt oder <strong>in</strong>direkt bedrohen.<br />
Die Folgen dieser Entwicklung s<strong>in</strong>d heute noch nicht abschätzbar.<br />
Nach der Biodiversitätskonvention ist die Schweiz verpflichtet, Massnahmen gegen<br />
<strong>in</strong>vasive gebietsfremde Arten zu ergreifen und deren Verbreitung e<strong>in</strong>zudämmen oder<br />
zu verh<strong>in</strong>dern. Nach dem Motto «Gefahr erkannt – Gefahr gebannt» ist es für die<br />
Schweiz von zentraler Bedeutung, potenziell gefährliche Arten zu erkennen. Das<br />
vorliegende Kompendium ist hierfür gedacht. Es beschreibt <strong>in</strong> umfassender Weise von<br />
den Flechten bis h<strong>in</strong> zu Säugetieren gebietsfremde Arten mit Schadenpotenzial, die<br />
schon hier s<strong>in</strong>d oder die vor den Toren der Schweiz stehen.<br />
Georg Karlaganis<br />
Head of Substances, Soil and Biotechnology Division<br />
Federal Office for the Environment (FOEN)
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 8<br />
> Summary<br />
Globalization is <strong>in</strong>creas<strong>in</strong>g trade and travel on an unprecedented scale, and has <strong>in</strong>advertently<br />
led to the <strong>in</strong>creased transport and <strong>in</strong>troduction of <strong>alien</strong> <strong>species</strong>, break<strong>in</strong>g down<br />
the natural barriers between countries and cont<strong>in</strong>ents. Alien <strong>species</strong> are not bad per se,<br />
<strong>in</strong> fact many <strong>species</strong> are beneficial for humans, e.g. most crop <strong>species</strong>. However, some<br />
<strong>alien</strong> <strong>species</strong> have become harmful, and pose threats to the environment and humans.<br />
<strong>Invasive</strong> <strong>alien</strong> <strong>species</strong> (IAS) are <strong>in</strong>creas<strong>in</strong>gly recognized as one of the major threats to<br />
biodiversity.<br />
All signatories to the Convention on Biological Diversity (CBD), <strong>in</strong>clud<strong>in</strong>g <strong>Switzerland</strong>,<br />
have agreed to prevent the <strong>in</strong>troduction of, control or eradicate those <strong>alien</strong> <strong>species</strong><br />
which threaten ecosystems, habitats or <strong>species</strong>.<br />
There is a widespread view that IAS are of less concern <strong>in</strong> Central Europe than on<br />
other cont<strong>in</strong>ents (and more especially on islands). Possible reasons for this <strong>in</strong>clude the<br />
small size of nature reserves, the high human impact on all ‘natural’ environments, and<br />
the long association of <strong>alien</strong> <strong>species</strong> and humans lead<strong>in</strong>g to familiarisation and adaptation.<br />
However, the number of cases of dramatic impact is <strong>in</strong>creas<strong>in</strong>g and awareness<br />
among scientists and the public is steadily grow<strong>in</strong>g. Thus, the threats from IAS should<br />
not be underrated. One of the major consequences, which is undoubtedly unfold<strong>in</strong>g<br />
before our eyes, is global homogenization, with the unique character of places such as<br />
<strong>Switzerland</strong> be<strong>in</strong>g lost, the characteristic flora and fauna <strong>in</strong>vaded by organisms which<br />
reproduce to eventually form the largest proportion of biomass <strong>in</strong> certa<strong>in</strong> ecosystems.<br />
Time lags, i.e. the gap between establishment and <strong>in</strong>vasion, makes prediction of <strong>in</strong>vasiveness<br />
of <strong>alien</strong> <strong>species</strong> very difficult. Well-established <strong>species</strong> show<strong>in</strong>g no h<strong>in</strong>t of any<br />
harm to the environment may still become <strong>in</strong>vasive <strong>in</strong> the future. There are three major<br />
categories of factors that determ<strong>in</strong>e the ability of a <strong>species</strong> to become <strong>in</strong>vasive: <strong>in</strong>tr<strong>in</strong>sic<br />
factors or <strong>species</strong> traits; extr<strong>in</strong>sic factors or relationships between the <strong>species</strong> and<br />
abiotic and biotic factors; and the human dimension, <strong>in</strong>corporat<strong>in</strong>g the importance of<br />
<strong>species</strong> to humans.<br />
This report compiles <strong>in</strong>formation about <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> from published<br />
sources and experts <strong>in</strong> <strong>Switzerland</strong> and abroad. Imm<strong>in</strong>ent future bio<strong>in</strong>vasions are also<br />
<strong>in</strong>cluded. The availability of national lists varies greatly between taxonomic groups.<br />
Thus, unfortunately, it is not possible to list all the <strong>alien</strong> <strong>species</strong> of <strong>Switzerland</strong>, s<strong>in</strong>ce<br />
not all resident <strong>species</strong> are known yet. However, for groups that are well known,<br />
complete lists have been compiled. For some groups only <strong>in</strong>vasive <strong>species</strong> rather than<br />
<strong>alien</strong> <strong>species</strong> are treated, and other groups could not be covered at all.<br />
The broad taxonomic group<strong>in</strong>gs used are vertebrates, crustaceans, <strong>in</strong>sects, arachnids,<br />
molluscs, other animals, fungi and plants. For each group, we present an overview, a<br />
list of <strong>alien</strong> <strong>species</strong>, Fact Sheets for the <strong>in</strong>vasive <strong>species</strong>, an evaluation of the status,
Summary 9<br />
impacts, pathways and control options for the group, and recommendations. The Fact<br />
Sheets summarize <strong>in</strong>formation on the <strong>in</strong>vasive <strong>species</strong> under the head<strong>in</strong>gs taxonomy,<br />
description, ecology, orig<strong>in</strong>, <strong>in</strong>troduction, distribution, impact, management and references.<br />
Def<strong>in</strong>itions of the important terms used <strong>in</strong> the report are given, s<strong>in</strong>ce frequently-used<br />
terms such as ‘<strong>in</strong>vasive’ are often used <strong>in</strong> different ways.<br />
The situation regard<strong>in</strong>g IAS <strong>in</strong> <strong>Switzerland</strong> is similar to that <strong>in</strong> other Central European<br />
countries, <strong>in</strong> particular Austria, which is also a land-locked country conta<strong>in</strong><strong>in</strong>g part of<br />
the Alps. This report lists about 800 <strong>alien</strong> <strong>species</strong> and characterises 107 IAS <strong>in</strong> Fact<br />
Sheets: five mammals, four birds, one reptile, three amphibians, seven fish, four molluscs,<br />
16 <strong>in</strong>sects, six crustaceans, three spiders, two ‘worms’, seven fungi, one bacteria,<br />
and 48 plants.<br />
Pathways can be divided <strong>in</strong>to those for <strong>species</strong> deliberately <strong>in</strong>troduced and those for<br />
<strong>species</strong> accidentally <strong>in</strong>troduced. Pathways for deliberate <strong>in</strong>troductions <strong>in</strong>clude the trade<br />
of <strong>species</strong> used <strong>in</strong> aquaculture, for fisheries, as forest trees, for agricultural purposes,<br />
for hunt<strong>in</strong>g, for soil improvement and solely to please humans as ornamentals. Most of<br />
these can also transport hitchhik<strong>in</strong>g <strong>species</strong> and people can accidentally <strong>in</strong>troduce<br />
<strong>species</strong> while travell<strong>in</strong>g. In general, most aquatics and terrestrial <strong>in</strong>vertebratesand<br />
diseases are accidental arrivals, whereas most plants and vertebrates are deliberately<br />
<strong>in</strong>troduced. The global trend for the latter groups also holds true for <strong>Switzerland</strong>, e.g.<br />
75 % of the 20 Black List plants were <strong>in</strong>troduced pr<strong>in</strong>cipally as ornamentals, and 35 of<br />
the 37 vertebrates were deliberately <strong>in</strong>troduced. Thus, many damag<strong>in</strong>g <strong>in</strong>vaders were<br />
deliberately <strong>in</strong>troduced, often with little justification beyond the wish to “improve” the<br />
landscape, e.g. ornamental plants and waterfowl.<br />
The impacts of IAS are often considerable, <strong>in</strong> particular when ecosystem function<strong>in</strong>g is<br />
altered or <strong>species</strong> are pushed to ext<strong>in</strong>ction, as has been shown for many bird <strong>species</strong> on<br />
islands. The environmental impacts can be divided <strong>in</strong>to four major factors: competition,<br />
predation, hybridization and transmission of diseases. The most obvious examples<br />
for competition are between <strong>in</strong>troduced and native plants for nutrients and exposure to<br />
sunlight. Resource competition has also led to the replacement of the native red squirrel<br />
(Sciurus vulgaris) by the <strong>in</strong>troduced American grey squirrel (S. carol<strong>in</strong>ensis) <strong>in</strong><br />
almost all of Great Brita<strong>in</strong> and it is predicted that this trend will cont<strong>in</strong>ue on the cont<strong>in</strong>ent.<br />
The musk rat (Ondatra zibethicus) causes decl<strong>in</strong>es of native mussels population<br />
(Unionidae) by predation and the amphipod Dikerogammarus villosus is a serious<br />
predator of native freshwater <strong>in</strong>vertebrates. A well-known example of hybridization<br />
from Europe is the ruddy duck (Oxyura jamaicensis), which hybridizes with the endangered<br />
native white-headed duck (O. leucocephala). In some cases IAS can harbour<br />
diseases and act as a vector for those diseases to native <strong>species</strong>. This is the case with<br />
American <strong>in</strong>troduced crayfish <strong>species</strong> to Europe, which are almost asymptomatic<br />
carriers of the <strong>alien</strong> crayfish plague (Aphanomyces astaci), but the native noble crayfish<br />
(Astacus astacus) is highly susceptible to the disease, and thus is struggl<strong>in</strong>g to<br />
coexist with populations of the American crayfish <strong>species</strong>.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 10<br />
In addition to impacts on biodiversity, many IAS cause enormous economic costs.<br />
These costs can arise through direct losses of agricultural and forestry products and<br />
through <strong>in</strong>creased production costs associated with control measures. A North American<br />
study calculated costs of US$ 138 billion per annum to the USA from IAS. A<br />
recently released report estimates that weeds are cost<strong>in</strong>g agriculture <strong>in</strong> Australia about<br />
Aus$ 4 billion a year, whereas weed control <strong>in</strong> natural environments cost about Aus$<br />
20 million <strong>in</strong> the year from mid 2001 to mid 2002. In Europe, the costs of giant hogweed<br />
(Heracleum mantegazzianum) <strong>in</strong> Germany are estimated at € 10 million; € 1<br />
million relat<strong>in</strong>g to each of the environment and health sectors, and the rema<strong>in</strong>der<br />
represents costs to the agricultural and forestry sectors. Economic damage by the<br />
western corn rootworm (Diabrotica virgifera) is <strong>in</strong>creas<strong>in</strong>g as it spreads through<br />
Europe. Some IAS also have implications for human health, e.g. giant hogweed produces<br />
copious amounts of a sap conta<strong>in</strong><strong>in</strong>g phototoxic substances (furanocoumar<strong>in</strong>s),<br />
which can lead to severe burns to the sk<strong>in</strong>. The raccoon dog (Nyctereutes procyonoides),<br />
<strong>in</strong>troduced as a fur animal, can, like the native red fox (Vulpes vulpes), act as a<br />
vector of the most dangerous parasitic disease vectored by mammals to humans <strong>in</strong><br />
Central Europe, i.e. the fox tapeworm (Ech<strong>in</strong>ococcus multilocularis). Known impacts<br />
of <strong>species</strong> <strong>in</strong>troduced <strong>in</strong>to <strong>Switzerland</strong> are presented, although some recent <strong>in</strong>vaders<br />
have not yet been reported to have impact <strong>in</strong> <strong>Switzerland</strong>; <strong>in</strong> these cases impacts assessed<br />
<strong>in</strong> other countries are given. Demonstration of environmental impacts is often<br />
difficult because of the complexity of ecosystems, but <strong>alien</strong> <strong>species</strong> occurr<strong>in</strong>g <strong>in</strong> high<br />
numbers, such as Japanese knotweed (Reynoutria japonica) totally cover<strong>in</strong>g riversides,<br />
or an animal biomass of <strong>alien</strong> <strong>species</strong> of up to 95 % <strong>in</strong> the Rh<strong>in</strong>e near Basel, must have<br />
impacts on the native ecosystem. All <strong>species</strong> use resources and are resources to other<br />
creatures and so they alter the web and nutrient flow of the ecosystems they are liv<strong>in</strong>g<br />
<strong>in</strong>.<br />
Recommendations for the management – <strong>in</strong> its widest sense – of groups of <strong>in</strong>vasives<br />
and <strong>in</strong>dividual <strong>species</strong> are given <strong>in</strong> the respective chapters and Fact Sheets. Prepar<strong>in</strong>g a<br />
national strategy aga<strong>in</strong>st IAS is recommended to deal with IAS <strong>in</strong> an appropriate way<br />
and as anticipated by the CBD. This action plan should identify the agency responsible<br />
for assess<strong>in</strong>g the risks posed by <strong>in</strong>troductions, provide fund<strong>in</strong>g mechanisms and technical<br />
advice and support for control options. Prevention measures aga<strong>in</strong>st further bio<strong>in</strong>vasions<br />
need to be put <strong>in</strong> place to stem the tide of <strong>in</strong>com<strong>in</strong>g new <strong>species</strong> arriv<strong>in</strong>g<br />
accidentally with trade and travel or <strong>in</strong>troduced deliberately for various purposes. New<br />
deliberate <strong>in</strong>troductions must be assessed as to the threat they may present and only<br />
<strong>in</strong>troduced on the basis of a risk analysis and environmental impact assessment. This<br />
report <strong>in</strong>dicates some important pathways for consideration and shows that most<br />
<strong>in</strong>vasive <strong>species</strong> are deliberately <strong>in</strong>troduced. The use of native plants and non-<strong>in</strong>vasive<br />
<strong>alien</strong> plants for garden<strong>in</strong>g and other purposes should be promoted. Laws regulat<strong>in</strong>g<br />
trade <strong>in</strong> plant <strong>species</strong> on the Black List would be a first step <strong>in</strong> the right direction to<br />
reduce the impact of these <strong>species</strong>. However, restrictions for <strong>species</strong> already widely<br />
distributed with<strong>in</strong> <strong>Switzerland</strong> will not drastically change the situation unless the<br />
populations already present are eradicated or controlled. Fish <strong>in</strong>troductions are regulated<br />
by the Fisheries Act, which names those <strong>species</strong> for which an authorization for<br />
release is needed, and <strong>species</strong> for which release is prohibited altogether. This is a good<br />
basis, although the law could be better adapted to the current situation, as described <strong>in</strong>
Summary 11<br />
the fish section. The aquarium and terrarium trade is another important sector that<br />
could be more strictly regulated to stop releases of pets <strong>in</strong>to the wild. A major problem<br />
with <strong>in</strong>troductions of IAS is that the costs when they <strong>in</strong>vasive are borne by the public,<br />
while the f<strong>in</strong>ancial <strong>in</strong>centives for <strong>in</strong>troduc<strong>in</strong>g them lie with <strong>in</strong>dividuals or specific<br />
bus<strong>in</strong>esses. Development of economic tools that shift the burden of IAS to those who<br />
benefit from <strong>in</strong>ternational trade and travel is a neglected approach (also called the<br />
‘polluter pays’ pr<strong>in</strong>ciple). Appropriate tools would be fees and taxes, <strong>in</strong>clud<strong>in</strong>g fees<br />
levied on those who import the organisms or goods. Awareness rais<strong>in</strong>g is a significant<br />
tool <strong>in</strong> the prevention and management of IAS, s<strong>in</strong>ce some member of the public would<br />
adhere to advice if they knew about its importance and the reason for it. Scientists and<br />
decision-makers also need better access to <strong>in</strong>formation about <strong>in</strong>vasive <strong>species</strong>, their<br />
impacts, and management options. To address the impacts of those <strong>in</strong>vasive <strong>species</strong><br />
already present <strong>in</strong> <strong>Switzerland</strong>, their populations need to be managed: either eradicated<br />
or controlled. Possible targets for eradication are the sika deer (Cervus nippon), the<br />
mouflon (Ovis orientalis), and the ruddy shelduck (Tadorna ferrug<strong>in</strong>ea), which will<br />
otherwise <strong>in</strong>crease its range and spread to neighbour<strong>in</strong>g countries. A pilot national<br />
eradication / control programme aga<strong>in</strong>st a prom<strong>in</strong>ent <strong>in</strong>vader, e.g. a Black List plant<br />
<strong>species</strong>, is recommended as a case study. Monitor<strong>in</strong>g populations of some <strong>alien</strong> <strong>species</strong><br />
is recommended to detect any sudden <strong>in</strong>creases <strong>in</strong>dicat<strong>in</strong>g potential <strong>in</strong>vasiveness. By<br />
do<strong>in</strong>g this, control or even eradication efforts can be employed before the populations<br />
become unmanageable. While compil<strong>in</strong>g the <strong>in</strong>formation for this report it became clear<br />
that more <strong>in</strong>formation about the status of IAS <strong>in</strong> <strong>Switzerland</strong> is needed. More studies<br />
on <strong>alien</strong> <strong>species</strong> are highly recommended to assess the importance of IAS and to demonstrate<br />
their significance to policy-makers and politicians.<br />
Limited resources dictate the need for sett<strong>in</strong>g priorities and allocat<strong>in</strong>g funds where it<br />
will have the greatest impact <strong>in</strong> combat<strong>in</strong>g IAS. Important po<strong>in</strong>ts, for example, are to<br />
critically assess the feasibility of different approaches, and to target <strong>species</strong> for which<br />
there is no conflict of <strong>in</strong>terest. Opposition to action aga<strong>in</strong>st less-important ornamentals<br />
on the Black List and <strong>species</strong> of direct human health concern (giant hogweed) should<br />
be negligible.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 12<br />
> Zusammenfassung<br />
Mit der zunehmenden Globilisierung ist e<strong>in</strong> starker Anstieg des Warentransportes,<br />
Verkehrs und Tourismus zu verzeichnen. Dies führt zu ungewollten wie beabsichtigten<br />
E<strong>in</strong>führungen von gebietsfremden Arten <strong>in</strong> e<strong>in</strong>em noch nie dagewesenen Umfang, und<br />
der Verschmelzung von Biodiversitäten der unterschiedlichen Länder und Kont<strong>in</strong>ente,<br />
so dass nur schwer zu überbrückende natürliche Ausbreitungsschranken plötzlich<br />
überwunden werden. Nicht alle gebietsfremden Arten s<strong>in</strong>d automatisch als negativ zu<br />
bewerten. Tatsächlich s<strong>in</strong>d viele Arten wichtige Bestandteile der Ökonomie e<strong>in</strong>es<br />
Landes, man denke nur an die zahlreichen gebietsfremden Kulturpflanzen. E<strong>in</strong>ige<br />
Arten entwickeln sich allerd<strong>in</strong>gs zu Problemarten und bedrohen die e<strong>in</strong>heimische<br />
Biodiversität, richten wirtschaftlichen Schaden an oder stellen e<strong>in</strong>e Gefahr für die<br />
Gesundheit dar.<br />
Gebietsfremde Problemarten (<strong>in</strong>vasive <strong>alien</strong> <strong>species</strong>) werden heute als e<strong>in</strong>e Hauptbedrohung<br />
für die Biodiversität angesehen. Die Biodiversitätskonvention (CBD) verpflichtet<br />
die <strong>in</strong>ternationale Staatengeme<strong>in</strong>schaft Vorsorge gegen diese <strong>in</strong>vasiven Arten<br />
zu treffen und diese gegebenenfalls zu bekämpfen.<br />
Gebietsfremde Arten <strong>in</strong> Zentraleuropa werden oft als ger<strong>in</strong>ges Problem e<strong>in</strong>gestuft, im<br />
Vergleich zu anderen Kont<strong>in</strong>enten und vor allem Inseln. Mögliche Gründe für diese<br />
Unterschiede s<strong>in</strong>d die relativ kle<strong>in</strong>en Schutzgebiete, was die Möglichkeit für e<strong>in</strong>e<br />
<strong>in</strong>tensive Pflege eröffnet, die stark vom Menschen bee<strong>in</strong>flussten ‘Naturräume’ und das<br />
lange Zusammenleben von vielen gebietsfremden Arten mit dem Menschen, das zu<br />
vielfältigen Anpassungen geführt hat. Trotzdem nehmen die Fälle von dramatischen<br />
Auswirkungen von gebietsfremden Arten und das Bewusstse<strong>in</strong> <strong>in</strong> der Bevölkerung und<br />
bei den Wissenschaftlern zu. Zweifellos ist die globale Homogenisierung <strong>in</strong> vollem<br />
Gange und der e<strong>in</strong>zigartige Charakter von lokalen Ökosystemen, wie zum Beispiel <strong>in</strong><br />
der Schweiz, gehen für immer verloren, da die charakteristische Pflanzen- und Tierwelt<br />
von gebietsfremden Arten verändert wird und e<strong>in</strong>ige dieser Arten die grössten Anteile<br />
an der Biomasse von e<strong>in</strong>igen Ökosystemen erreichen.<br />
Die Zeitdifferenz, die zwischen der Ankunft e<strong>in</strong>er Art und ihrer starken Ausbreitung<br />
auftreten kann, macht Voraussagungen der Invasivität von Arten aussserordentlich<br />
schwierig. E<strong>in</strong>ige schon lange etablierte Arten können plötzlich und unerwartet <strong>in</strong>vasiv<br />
werden. Drei Kategorien von Faktoren bestimmen die Invasivität von Arten: 1. die<br />
biologischen Merkmale e<strong>in</strong>er Art, 2. das Zusammenspiel e<strong>in</strong>er Art mit ihrer abiotischen<br />
und biotischen Umwelt und 3. die Beziehungen der Menschen zu dieser Art.<br />
In diesem Bericht ist Information über gebietsfremde Arten der Schweiz sowohl von<br />
publizierten Dokumenten als auch von direktem Austausch mit Experten <strong>in</strong> der<br />
Schweiz und des Auslandes zusammengetragen. Bevorstehende E<strong>in</strong>wanderungen s<strong>in</strong>d<br />
ebenfalls erfasst worden. Die Verfügbarkeit von Artenlisten <strong>in</strong> den e<strong>in</strong>zelnen taxonomischen<br />
Gruppen ist sehr unterschiedlich, so dass es nicht möglich ist alle gebiets-
Zusammenfassung 13<br />
fremden Arten zu benennen. In e<strong>in</strong>igen Gruppen ist das Wissen sogar der e<strong>in</strong>heimische<br />
Arten so rudimentär, dass ke<strong>in</strong> Versuch gemacht wurde, sie zu bearbeiten, und bei<br />
anderen Gruppen wurden nur Problemarten aufgenommen. Die Listen der gebietsfremden<br />
Arten wieder anderer Gruppen dagegen s<strong>in</strong>d vollständig.<br />
Die gebietsfremden Organismen wurden <strong>in</strong> folgende Gruppen aufgeteilt: Wirbeltiere,<br />
Krebstiere, Insekten, Sp<strong>in</strong>nentiere, Weichtiere, andere Tiere, Pilze und Pflanzen. In<br />
jedem Kapitel bef<strong>in</strong>den sich die Listen der gebietsfremden Arten, e<strong>in</strong> erläuternder Text,<br />
Datenblätter der Problemarten und e<strong>in</strong>e Auswertung der Situation, der negativen<br />
Auswirkungen, der E<strong>in</strong>führungswege, der möglichen Gegensteuerungsmassnahmen<br />
und Empfehlungen für den Umgang mit diesen Arten. Die Datenblätter bieten Information<br />
zu Taxonomie, Beschreibung, Ökologie, Herkunft, E<strong>in</strong>führungswege, Verbreitung,<br />
Auswirkungen, Ansätze zur Gegensteuerung und e<strong>in</strong> Literaturverzeichnis.<br />
Def<strong>in</strong>itionen der wichtigsten Begriffe, wie sie <strong>in</strong> diesem Dokument benutzt werden,<br />
werden ebenfalls gegeben, da sie oftmals unterschiedlich gebraucht werden.<br />
Die Situation der gebietsfremden Arten der Schweiz ist ähnlich wie <strong>in</strong> anderen mitteleuropäischen<br />
Ländern, vor allem Österreich, das e<strong>in</strong>e ähnliche Topographie besitzt.<br />
Dieser Bericht über die gebietsfremden Arten der Schweiz listet über 800 etablierte<br />
gebietsfremde Arten auf und stellt die 107 Problemarten <strong>in</strong> Datenblättern vor: fünf<br />
Säugetiere, vier Vögel, e<strong>in</strong> Reptil, drei Amphibien, sieben Fische, vier Weichtiere, 16<br />
Insekten, sechs Krebstiere, drei Sp<strong>in</strong>nen, zwei ‘Würmer’, sieben Pilze, e<strong>in</strong> Bakterium<br />
und 48 Pflanzen.<br />
Es können versehentlich e<strong>in</strong>geschleppte Arten und bewusst e<strong>in</strong>geführte Arten unterschieden<br />
werden. E<strong>in</strong>geführt s<strong>in</strong>d zum Beispiel Arten der Aquakulturen, der Fischerei,<br />
der Waldwirtschaft, der Landwirtschaft, der Jagd, zur Bodenverbesserung und e<strong>in</strong>fach<br />
zur Bereicherung der Landschaft, wie Zierpflanzen. Viele der e<strong>in</strong>geführten Arten<br />
können allerd<strong>in</strong>gs andere Arten auf und <strong>in</strong> sich tragen und so e<strong>in</strong>schleppen, und der<br />
reisende Mensch transportiert ebenfalls oftmals gebietsfremde Arten. Die meisten<br />
aquatischen und terrestrischen Wirbellosen und Krankheiten wurden versehentlich<br />
e<strong>in</strong>geschleppt, während Pflanzen und Wirbeltiere meist e<strong>in</strong>geführt worden s<strong>in</strong>d. Dieser<br />
globale Trend f<strong>in</strong>det sich auch bei den gebietsfremden Arten der Schweiz wieder, denn<br />
75 % der 20 Arten auf der ‘Schwarzen Liste’ wurden als Zierpflanzen e<strong>in</strong>geführt und<br />
35 der 37 Wirbeltiere wurde zu e<strong>in</strong>em bestimmten Zweck importiert. Das heisst, dass<br />
viele der Problemarten bewusst e<strong>in</strong>geführt wurden, oftmals mit e<strong>in</strong>er ger<strong>in</strong>gfügigen<br />
Rechtfertigung, z.B. um die Landschaft mit Zierpflanzen und Wasservögeln zu ‘bereichern’.<br />
Die Auswirkungen, die gebietsfremde Arten auslösen können, s<strong>in</strong>d oft beträchtlich, vor<br />
allem wenn die Funktion e<strong>in</strong>es Ökosystems gestört wird, e<strong>in</strong>heimische Arten verdrängt<br />
werden oder sogar aussterben, wie es bei Vogelarten auf Inseln dokumentiert worden<br />
ist. Vier Faktoren können zu solchen Problemen führen: 1. Konkurrenz zu e<strong>in</strong>heimischen<br />
Arten, 2. e<strong>in</strong> gebietsfremder Räuber, 3. die Hybridisierung mit e<strong>in</strong>heimischen<br />
Arten und 4. die Ausbreitung von Krankheiten durch e<strong>in</strong>en gebietsfremden Vektor.<br />
Offensichtliche Beispiele für Konkurrenz s<strong>in</strong>d der Kampf um Licht und Nährstoffe
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 14<br />
zwischen gebietsfremden und e<strong>in</strong>heimischen Pflanzenarten. Der Konkurrenzkampf um<br />
Nahrung hat <strong>in</strong> Grossbrittanien zur fast völligen Verdrängung des Eichhörnchens<br />
(Sciurus vulgaris) durch das e<strong>in</strong>geführte Grauhörnchen (S. carol<strong>in</strong>ensis) geführt und es<br />
ist zu befürchten, dass dieser Trend auch auf dem Festland weitergehen wird. Der<br />
Bisam (Ondatra zibethicus) hat als Räuber der e<strong>in</strong>heimischen Muscheln (Unionidae)<br />
zu ihrem Rückgang beigetragen und der Amphipode Dikerogammarus villosus ist e<strong>in</strong><br />
grosser Fe<strong>in</strong>d der e<strong>in</strong>heimischen Wirbellosen der Gewässer. E<strong>in</strong> bekanntes Beispiel für<br />
e<strong>in</strong>e Hybridisierung ist die e<strong>in</strong>geführte Schwarzkopf-Ruderente (Oxyura jamaicensis),<br />
die sich mit der stark gefährdeten Weisskopf-Ruderente (O. leucocephala) verpaart. In<br />
e<strong>in</strong>igen Fällen können gebietsfremde Arten Krankheiten unter e<strong>in</strong>heimischen Arten<br />
verbreiten. Dies ist der Fall bei der berüchtigten Krebspest (Aphanomyces astaci), die<br />
von ebenfalls e<strong>in</strong>geführten nordamerikanischen Flusskrebsen, die fast ke<strong>in</strong>e Symptome<br />
zeigen, auf den e<strong>in</strong>heimischen Flusskrebs (Astacus astacus), der dramatisch mit e<strong>in</strong>em<br />
sofortigen Zusammenbruch der Population reagiert, übertragen werden.<br />
Neben diesen Auswirkungen auf die Umwelt, können gebietsfremde Arten auch enorme<br />
ökonomische Schäden verursachen. Die Kosten können durch den Verlust von<br />
land- und forstwirtschaftlichen Produkten und durch erhöhte Produktionskosten durch<br />
Bekämpfungsmassnahmen entstehen. E<strong>in</strong>e nordamerikanische Studie hat die jährlichen<br />
Kosten von gebietsfremden Arten <strong>in</strong> der USA auf 13,8 Milliarden US Dollar berechnet.<br />
E<strong>in</strong> anderer Bericht schätzt die Kosten durch Unkräuter für die australische Landwirtschaft<br />
auf 4 Milliarden Australische Dollar, und 20 Millionen Aus. $ wurden während<br />
e<strong>in</strong>es Jahres zwischen Mitte 2001 und Mitte 2002 für die Unkrautbekämpfung auf<br />
naturnahen Flächen ausgegeben. Die Kosten durch den Riesenbärenklau (Heracleum<br />
mantegazzianum) <strong>in</strong> Deutschland werden auf 10 Millionen € geschätzt, wobei je e<strong>in</strong>e<br />
Millionen im Umweltbereich und Gesundheitswesen anfallen und der Rest <strong>in</strong> Landwirtschaft<br />
und Forst. Der Westliche Maiswurzelbohrer (Diabrotica virgifera) dehnt<br />
se<strong>in</strong>e Verbreitung weiter nach Nordwesten aus und bereitet grosse Schäden an den<br />
Maiskulturen. E<strong>in</strong>ige gebietsfremde Arten schaden der menschlichen Gesundheit, so<br />
produziert der Riesenbärenklau grosse Mengen e<strong>in</strong>es Saftes der phototoxische Substanzen<br />
(Furanocumar<strong>in</strong>e), die zu starken Verbrennungen der Haut führen können,<br />
enthält. Der Marderhund (Nyctereutes procyonoides), als Pelztier e<strong>in</strong>geführt, kann, wie<br />
der e<strong>in</strong>heimische Rotfuchs (Vulpes vulpes), als Vektor des Fuchsbandwurmes (Ech<strong>in</strong>ococcus<br />
multilocularis), der gefährlichsten Krankheit, die <strong>in</strong> Zentraleuropa von Säugetieren<br />
auf den Menschen übertragen wird, fungieren. Für diesem Bericht wurden<br />
bekannte Auswirkungen von gebietsfremden Arten <strong>in</strong> der Schweiz zusammengetragen.<br />
Für Arten, die noch nicht lange <strong>in</strong> der Schweiz vorkommen, wurde auf Berichten von<br />
Auswirkungen <strong>in</strong> anderen Ländern zurückgegriffen. Es muss erwähnt werden, dass<br />
Nachweise von Auswirkungen e<strong>in</strong>er gebietsfremden Art <strong>in</strong> e<strong>in</strong>em komplexen Ökosystem<br />
oft schwierig zu führen s<strong>in</strong>d. Andererseits ist es offensichtlich, dass Arten wie der<br />
Japanische Staudenknöterich (Reynoutria japonica), der oft ganze Flussufer säumt,<br />
oder e<strong>in</strong>e tierische Biomasse von gebietsfremden Arten von 95 % im Rhe<strong>in</strong> bei Basel,<br />
e<strong>in</strong>e Auswirkung auf das Ökosystem haben müssen. Alle Arten verbrauchen Nährstoffe<br />
und dienen als Nährstoff für andere Organismen und ändern so das Nahrungsnetz und<br />
den Nährstofffluss der Ökosysteme, die sie besiedeln.
Zusammenfassung 15<br />
In den Texten der jeweiligen Kapitel und den Datenblättern s<strong>in</strong>d Empfehlungen zur<br />
Gegensteuerung (Prävention und Kontrolle) für die Gruppen und e<strong>in</strong>zelnen Arten<br />
gegeben. Allgeme<strong>in</strong> ist die Erstellung e<strong>in</strong>er Nationalen Strategie im H<strong>in</strong>blick auf<br />
gebietsfremde Arten zu empfehlen, um angemessene Schritte ergreifen zu können, und<br />
es von der Biodiversitätskonvention gefordert ist. Dieser Plan sollte e<strong>in</strong>e zuständige<br />
Behörde identifizieren, die die Risiken von E<strong>in</strong>führungen und E<strong>in</strong>schleppungen beurteilt,<br />
für f<strong>in</strong>anzielle Mittel sorgt und technische Unterstützung zur Bekämpfung bereitstellt.<br />
Massnahmen zur Prävention um weitere Bio<strong>in</strong>vasionen zu stoppen oder zu<br />
verm<strong>in</strong>dern müssen ausgearbeitet werden. E<strong>in</strong>führungen von neuen Organismen sollten<br />
vorher auf ihre möglichen Gefahren für die Umwelt untersucht werden und nur auf der<br />
Basis e<strong>in</strong>er Risikoanalyse e<strong>in</strong>geführt werden. Die Analyse der wichtigsten E<strong>in</strong>führungswege<br />
zeigt unmissverständlich, dass die meisten Problemarten bewusst e<strong>in</strong>geführt<br />
wurden (und werden). Die Nutzung von e<strong>in</strong>heimischen Arten und fremden Arten ohne<br />
Potential zur Invasivität zum Beispiel <strong>in</strong> Gärten, Parks und Forsten sollte mehr gefördert<br />
werden. Gesetze, die den Handel mit Pflanzenarten der ‘Schwarzen Liste’ regeln,<br />
wären e<strong>in</strong> konsequenter nächster Schritt, um die Auswirkungen dieser Arten zu reduzieren.<br />
Wenn die Arten allerd<strong>in</strong>gs schon e<strong>in</strong>e weite Verbreitung <strong>in</strong> der Schweiz besitzen,<br />
können nur Kontrollmassnahmen oder e<strong>in</strong>e erfolgreiche Ausrottung Abhilfe<br />
schaffen. Die Fischereiverordnung reguliert Fischaussetzungen, <strong>in</strong>dem sie Arten benennt<br />
für die e<strong>in</strong>e Bewilligung nötig ist und Arten, deren Aussetzung verboten ist.<br />
Diese solide Basis könnte noch verbessert werden, um der Situation besser zu entsprechen,<br />
wie <strong>in</strong> dem Teil über Fische beschrieben. E<strong>in</strong> weiterer Sektor, der mehr reguliert<br />
werden sollte, ist der Handel mit Haustieren (vor allem Aquarium and Terrarium), der<br />
immer wieder zu Aussetzungen führt. E<strong>in</strong> Grundproblem der E<strong>in</strong>führungen ist, dass die<br />
Kosten von Problemarten von der Öffentlichkeit getragen werden, während der f<strong>in</strong>anzielle<br />
Nutzen der E<strong>in</strong>führung e<strong>in</strong>zelnen Importeuren oder bestimmten Wirtschaftszweigen<br />
zugute kommt. Die Entwicklung von ökonomischen Programmen, die die Last<br />
auf die verteilt, die auch den Nutzen aus der E<strong>in</strong>fuhr haben, ist e<strong>in</strong> vernachlässigter<br />
Denkansatz (Verursacherpr<strong>in</strong>zip genannt). Möglichkeiten wären gegeben durch die<br />
Erhebung von Gebühren und Steuern, die für den Importeur zu bezahahlen wären. E<strong>in</strong>e<br />
wichtige Vorgehensweise, um die Probleme mit gebietsfremden Arten unter Kontrolle<br />
zu kriegen, ist die Schaffung e<strong>in</strong>es geschärftes Bewusstse<strong>in</strong>s der Problematik <strong>in</strong> der<br />
Bevölkerung. Wissenschaftler und Entscheidungsträger benötigen ebenfalls mehr<br />
Information über gebietsfremde Problemarten, deren Auswirkungen und den Möglichkeiten<br />
für e<strong>in</strong>e Gegensteuerung. E<strong>in</strong>ige Problemarten müssten bekämpft oder ausgerottet<br />
werde, um ihre Auswirkungen wirkungsvoll zu m<strong>in</strong>imieren. Mögliche Zielarten für<br />
e<strong>in</strong>e Ausrottung s<strong>in</strong>d der Sikahirsch (Cervus nippon), das Mufflon (Ovis orientalis)<br />
oder die Rostgans (Tadorna ferrug<strong>in</strong>ea), die sonst ihre Verbreitung weiter ausdehnt<br />
und die Nachbarländer erreichen wird. Für e<strong>in</strong>e erste grossangelegte Ausrottung oder<br />
Bekämpfung ist ebenfalls e<strong>in</strong>e Pflanzenart der ‘Schwarzen Liste’ zu empfehlen. Ausserdem<br />
wäre die Beobachtung der Populationen von gebietsfremden Arten empfehlenswert,<br />
um etwaige starke Zunahmen früh zu erkennen. In diesem Fall könnten<br />
Gegenmassnahmen ergriffen werden, bevor die Populationen zu gross werden. Beim<br />
Zusammentragen der Informationen wurde schnell klar, dass viel mehr Information<br />
über gebietsfremde Arten benotigt wird. Daher s<strong>in</strong>d mehr Studien zur Bedeutung von<br />
gebietsfremden Arten nötig, um Entscheidungsträger und Politiker auf die Lage aufmerksam<br />
zu machen.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 16<br />
Die limitierten Ressourcen, die zur Verfügung stehen, zw<strong>in</strong>gen Prioritäten zu setzen,<br />
um die f<strong>in</strong>anziellen Mittel dort e<strong>in</strong>zusetzen, wo sie die meiste Wirkung zeigen im<br />
Kampf gegen Problemarten. Dabei müssen wichtige Punkte berücksichtigt werden,<br />
etwa, welche Methode den grössten Nutzen br<strong>in</strong>gt, oder welche Arten für Bekämpfungsmassnahmen<br />
zuerst <strong>in</strong> Betracht gezogen werden sollten. Arten mit e<strong>in</strong>em hohen<br />
Potenzial für Konflikte versprechen weniger Erfolg. Wenn Arten der ‘Schwarzen<br />
Liste’, welche ke<strong>in</strong>e grosse Wichtigkeit als Zierpflanzen besitzen, oder Arten die den<br />
Menschen gefährden, als Ziele ausgewählt werden, ist der zu erwartende Widerstand<br />
gegen Massnahmen eher ger<strong>in</strong>g e<strong>in</strong>zuschätzen.
Résumé 17<br />
> Résumé<br />
La globalisation a pour effet une augmentation sans précédant du commerce et des<br />
transports, dont une des conséquences est l’accroissement des déplacements et <strong>in</strong>troductions<br />
d’espèces exotiques. Les espèces exotiques ne sont pas toutes nuisibles. En<br />
fait un grande nombre d’entre elles sont bénéfiques, comme par exemple les nombreuses<br />
plantes cultivées d’orig<strong>in</strong>e étrangère. Cependant, certa<strong>in</strong>es espèces exotiques deviennent<br />
nuisibles et posent des problèmes à l’environnement et à l’homme en général.<br />
Les Espèces Exotiques Envahissantes (EEE) sont de plus en plus reconnues comme<br />
une des menaces les plus sérieuses posées à la biodiversité.<br />
Tous les pays signataires de la convention sur la diversité biologique (CDB), dont la<br />
Suisse, se sont engagés à prévenir l’<strong>in</strong>troduction, à contrôler ou éradiquer les espèces<br />
exotiques menaçant les écosystèmes, les habitats ou les espèces.<br />
Il est communément avancé que les EEE causent mo<strong>in</strong>s de problèmes en Europe<br />
Centrale que dans d’autres cont<strong>in</strong>ents ou régions. Les raisons possibles sont, entre<br />
autres, la taille limitée des réserves naturelles, l’impact huma<strong>in</strong> important dans tous les<br />
milieux « naturels » et la longue association, en Europe, entre les espèces exotiques et<br />
l’homme, ayant conduit à une familiarisation de ces espèces et une adaptation à<br />
l’environnement huma<strong>in</strong>. Cependant, le nombre de cas d’espèces exotiques causant des<br />
dégâts importants est en augmentation en Europe, un phénomène dont les chercheurs,<br />
mais également le public, ont de plus en plus conscience. De fait, la menace des espèces<br />
envahissantes ne doit pas être sous-estimée. Une des conséquences les plus visibles<br />
est le phénomène d’uniformisation, menant à la perte de paysages uniques, y compris<br />
en Suisse. La flore et la faune caractéristiques sont de plus en plus envahies par les<br />
organismes exotiques qui se reproduisent, pour f<strong>in</strong>alement composer la plus grande<br />
partie de la biomasse de certa<strong>in</strong>s écosystèmes.<br />
Le délai qui s’écoule entre la phase d’établissement et d’<strong>in</strong>vasion d’une espèce exotique<br />
(‘time lag’), rend la prédiction du phénomène d’<strong>in</strong>vasion très difficile. Des espèces<br />
bien établies qui n’ont actuellement aucun impact reconnu sur l’environnement peuvent<br />
malgré tout devenir envahissantes dans le futur. Trois catégories de facteurs<br />
déterm<strong>in</strong>ent la capacité d’une espèce à devenir envahissante: les facteurs <strong>in</strong>tr<strong>in</strong>sèques<br />
liés à l’espèce, les facteurs extr<strong>in</strong>sèques, c.-à-d. les relations entre l’espèce et les facteurs<br />
biotiques ou abiotiques, et la dimension huma<strong>in</strong>e, par exemple l’importance de<br />
l’espèce pour l’homme.<br />
Ce rapport est une compilation des connaissances sur les espèces exotiques en Suisse,<br />
rassemblées à partir de publications et d’avis d’experts suisses et étrangers. Des <strong>in</strong>formations<br />
sur les <strong>in</strong>vasions biologiques imm<strong>in</strong>entes sont également <strong>in</strong>clues. La connaissance<br />
des espèces présentes en Suisse variant fortement d’un groupe taxonomique à<br />
l’autre (pour certa<strong>in</strong>s taxa, même les espèces <strong>in</strong>digènes sont lo<strong>in</strong> d’être toutes connues),<br />
il n’a malheureusement pas été possible d’établir une liste exhaustive d’espèces exoti-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 18<br />
ques pour tous les groupes. Une liste complète a été établie seulement pour les groupes<br />
taxonomiques bien connus. Pour certa<strong>in</strong>s groupes, seules les espèces envahissantes ont<br />
été compilées alors que quelques groupes n’ont pas pu être traités.<br />
Les grands groupes taxonomiques traités sont les vertébrés, les crustacés, les <strong>in</strong>sectes,<br />
les arachnides, les mollusques, les autres animaux, les champignons et les plantes. Pour<br />
chaque groupe, nous présentons une discussion générale, la liste des espèces non<strong>in</strong>digènes,<br />
les fiches d’<strong>in</strong>formation sur les espèces envahissantes et une évaluation du<br />
statut, de l’impact, des modes d’<strong>in</strong>troduction, les méthodes de lutte et des recommandations<br />
générales. Les fiches d’<strong>in</strong>formation résument, pour des espèces particulièrement<br />
envahissantes ou potentiellement dangereuses, les <strong>in</strong>formations sur la taxonomie, la<br />
description, l’écologie, l’orig<strong>in</strong>e, l’<strong>in</strong>troduction en Suisse et en Europe, la distribution,<br />
l’impact, la gestion et les références bibliographiques.<br />
Les déf<strong>in</strong>itions des termes anglais les plus importants utilisés dans ce rapport sont<br />
données, parce que les mots fréquemment utilisés comme « <strong>in</strong>vasive » sont parfois<br />
utilisés dans des sens différents.<br />
La situation concernant les EEE en Suisse est similaire à celle d’autres pays d’Europe<br />
Centrale, en particulier l’Autriche, un pays également enclavé et Alp<strong>in</strong>. Ce rapport<br />
<strong>in</strong>ventorie environ 800 espèces non-<strong>in</strong>digènes et détaille 107 espèces envahissantes<br />
sous forme de fiches d’<strong>in</strong>formation : c<strong>in</strong>q mammifères, quatre oiseaux, un reptile, trois<br />
amphibiens, sept poissons, quatre mollusques, 16 <strong>in</strong>sectes, six crustacés, trois araignées,<br />
deux « vers », sept champignons, une bactérie et 48 plantes.<br />
Les modes d’<strong>in</strong>troduction des espèces exotiques sont différents selon qu’il s’agit<br />
d’espèces délibérément <strong>in</strong>troduites ou d’espèces <strong>in</strong>troduites accidentellement. Les<br />
<strong>in</strong>troductions délibérées concernent pr<strong>in</strong>cipalement les espèces importées pour l’aquaculture,<br />
la pêche, la chasse, la sylviculture, l’agriculture, l’horticulture et la protection<br />
des sols. Les organismes <strong>in</strong>troduits <strong>in</strong>volontairement sont souvent transportés par<br />
<strong>in</strong>advertance avec d’autres importations ou par des voyageurs. En général, la plupart<br />
des <strong>in</strong>vertébrés et des pathogènes ont été <strong>in</strong>troduits accidentellement, alors que les<br />
plantes et les vertébrés l’ont souvent été <strong>in</strong>tentionnellement. Cette tendance est également<br />
valable pour la Suisse. Sur les 20 plantes envahissantes de la liste noire, 75 % ont<br />
été <strong>in</strong>troduites pr<strong>in</strong>cipalement en tant que plantes ornementales et 35 des 37 vertébrés<br />
exotiques établis en Suisse ont été <strong>in</strong>troduits délibérément. Il est donc important de<br />
constater que beaucoup d’envahisseurs, dont certa<strong>in</strong>s parmi les plus nuisibles, ont été<br />
<strong>in</strong>troduits <strong>in</strong>tentionnellement, souvent sans autre soucis que d’améliorer le paysage,<br />
comme c’est le cas pour les plantes et animaux d’ornement.<br />
L’impact des EEE est parfois considérable, en particulier quand l’envahisseur altère le<br />
fonctionnement d’un écosystème ou pousse les espèces <strong>in</strong>digènes vers l’ext<strong>in</strong>ction,<br />
comme cela a été souvent observé avec les oiseaux en milieu <strong>in</strong>sulaire. Les impacts<br />
écologiques peuvent être causés par quatre mécanismes majeurs : la compétition, la<br />
prédation, l’hybridation et la transmission de maladies. Parmi les exemples les plus<br />
significatifs de compétition, nous pouvons citer la compétition entre les plantes <strong>in</strong>digènes<br />
et exotiques pour les nutriments et la lumière. La compétition pour les ressources a
Résumé 19<br />
également conduit au remplacement de l’écureuil roux <strong>in</strong>digène (Sciurus vulgaris) par<br />
l’écureuil gris américa<strong>in</strong> (S. carol<strong>in</strong>ensis) dans la plus grande partie de la Grande-<br />
Bretagne, et on s’attend à un même phénomène sur le cont<strong>in</strong>ent. Le rat musqué (Ondatra<br />
zibethicus) est responsable du décl<strong>in</strong> des populations de moules <strong>in</strong>digènes par<br />
prédation, et l’amphipode Dikerogammarus villosus est un sérieux prédateur des<br />
<strong>in</strong>vertébrés aquatiques <strong>in</strong>digènes. Un exemple bien connu d’hybridation en Europe est<br />
illustré par l’érismature rousse (Oxyura jamaicensis), un canard américa<strong>in</strong> qui<br />
s’hybride avec l’érismature à tête blanche (O. leucocephala), une espèce européenne en<br />
danger d’ext<strong>in</strong>ction. Dans certa<strong>in</strong>s cas, les EEE peuvent abriter des maladies et agir<br />
comme vecteur d’<strong>in</strong>fection pour les espèces <strong>in</strong>digènes. C’est le cas des espèces américa<strong>in</strong>es<br />
d’écrevisse <strong>in</strong>troduites en Europe, porteuses résistantes de la peste de l’écrevisse<br />
(Aphanomyces astaci), alors que l’écrevisse européenne (Astacus astacus) est très<br />
sensible à la maladie et a du mal à survivre au côté des espèces américa<strong>in</strong>es <strong>in</strong>troduites.<br />
En plus des impacts sur la biodiversité, les EEE ont des impacts considérables pour<br />
l’économie. Ces impacts économiques peuvent être liés aux pertes directes de produits<br />
agricoles ou forestiers ou à l’augmentation des coûts de production associés à la lutte<br />
contre les envahisseurs. Une étude américa<strong>in</strong>e a calculé que les EEE coûtait aux Etats-<br />
Unis la somme de 138 milliards de dollars par an. De même, un récent rapport austr<strong>alien</strong><br />
estime que les mauvaises herbes coûtent à l’agriculture austr<strong>alien</strong>ne environ 4<br />
milliards de dollars austr<strong>alien</strong>s par an et que le coût de la lutte contre les plantes envahissantes<br />
dans les milieux naturels atte<strong>in</strong>t 20 millions de dollars par an. En Europe, le<br />
coût de la berce du Caucase (Heracleum mantegazzianum) est estimé à € 10 millions<br />
en Allemagne, un million chacun pour les secteurs de l’environnement et de la santé<br />
publique, le reste représentant le coût pour les secteurs agricoles et forestiers. L’impact<br />
économique de la chrysomèle des rac<strong>in</strong>es du maïs (Diabrotica virgifera) augmente en<br />
même temps que sa dissém<strong>in</strong>ation en Europe. Certa<strong>in</strong>es EEE ont un impact sur la santé<br />
publique. Par exemple, la berce du Caucase contient des substances phototoxiques<br />
(furanocoumar<strong>in</strong>es) qui peuvent causer des brûlures sérieuses. Le chien viverr<strong>in</strong> (Nyctereutes<br />
procyonoides), <strong>in</strong>troduit pour sa fourrure, peut, comme le renard <strong>in</strong>digène<br />
(Vulpes vulpes), être vecteur le l’éch<strong>in</strong>ococcose du renard (Ech<strong>in</strong>ococcus multilocularis<br />
), une dangereuse maladie parasitaire pouvant être transmise à l’homme. Les impacts<br />
connus des espèces <strong>in</strong>troduites en Suisse sont présentés dans ce rapport. Comme<br />
plusieurs envahisseurs récents n’ont pas encore montré d’impacts écologiques en<br />
Suisse, les impacts observés dans d’autres pays sont également présentés. Mesurer les<br />
impacts écologiques des EEE est souvent une tâche difficile à cause de la complexité<br />
des écosystèmes. Cependant, les espèces exotiques présentes en très grand nombre,<br />
comme la renouée du Japon qui couvre totalement les bords de certa<strong>in</strong>es rivières, ou<br />
une faune aquatique exotique composant 95 % de la biomasse animale dans le Rh<strong>in</strong><br />
près de Bâle ont forcément un impact important sur les écosystèmes <strong>in</strong>digènes. Toutes<br />
les espèces utilisent des ressources et sont elle-mêmes ressources d’autres organismes<br />
vivants et, de ce fait, les espèces exotiques altèrent les chaînes alimentaires des écosystèmes<br />
dans lesquelles elles ont été <strong>in</strong>troduites.<br />
Des recommandations pour la gestion (au sens le plus large) des espèces envahissantes<br />
sont données dans les chapitres respectifs et les fiches d’<strong>in</strong>formation. Une stratégie<br />
nationale appropriée contre les EEE est recommandée af<strong>in</strong> de gérer le problème de la
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 20<br />
manière la plus appropriée, selon les exigences de la CDB. Le plan d’action devrait<br />
identifier l’agence responsable pour l’évaluation des risques posés par les <strong>in</strong>troductions<br />
et proposer les mécanismes de f<strong>in</strong>ancement a<strong>in</strong>si que le support technique pour les<br />
moyens de lutte. Des mesures de prévention contre les <strong>in</strong>vasions biologiques futures<br />
doivent être mises en place pour contenir l’implantation de nouvelles espèces, qu’elles<br />
soient accidentellement <strong>in</strong>troduites avec le commerce ou les voyages ou importées de<br />
façon <strong>in</strong>tentionnelle pour des raisons diverses. Les nouvelles <strong>in</strong>troductions délibérées<br />
doivent être évaluées pour le danger qu’elles représentent et les espèces <strong>in</strong>troduites<br />
uniquement après une analyse de risques et une étude d’impact écologique. Ce rapport<br />
mentionne les modes d’<strong>in</strong>troduction les plus importants et montre qu’un grand nombre<br />
d’EEE ont été délibérément <strong>in</strong>troduites. L’utilisation pour le jard<strong>in</strong>age de plantes<br />
<strong>in</strong>digènes et de plantes exotiques non-envahissantes devrait être promue. Une législation<br />
réglementant le commerce des plantes de la liste noire serait un premier pas dans<br />
la bonne direction pour réduire l’impact de ces espèces. Cependant, des restrictions<br />
d’utilisation pour les espèces déjà largement présentes en Suisse ne changeront pas<br />
grand-chose à la situation, à mo<strong>in</strong>s d’éradiquer ou de contrôler les populations déjà<br />
présentes. Les importations et implantations de poissons exotiques sont régulées par la<br />
loi fédérale sur la pêche, qui cite les espèces pour lesquelles une autorisation<br />
d’<strong>in</strong>troduction est nécessaire, et celles dont l’<strong>in</strong>troduction est prohibée. C’est une bonne<br />
base, bien que la loi pourrait être mieux adaptée à la situation actuelle, comme suggéré<br />
dans le chapitre relatif aux poissons. Le commerce des animaux d’aquarium et terrarium<br />
est un autre secteur important qui devrait être mieux régulé pour limiter les lâchers<br />
d’animaux de compagnie dans la nature. Un problème majeur lié aux EEE est<br />
que le coût de ces <strong>in</strong>vasions est payé par le public alors que ces <strong>in</strong>troductions sont<br />
motivées par des <strong>in</strong>térêts f<strong>in</strong>anciers privés. Le développement d’outils économiques<br />
transférant le coût des EEE aux bénéficiaires des échanges commerciaux impliquant<br />
ces organismes exotiques est une approche pour l’<strong>in</strong>stant négligée (pr<strong>in</strong>cipe du pollueur-payeur).<br />
Par exemple, des contributions ou taxes pourraient être imposés aux<br />
importateurs d’organismes vivants ou de marchandises. La sensibilisation du public est<br />
également un outil important dans la prévention et la gestion des EEE. Une partie du<br />
public adhèrerait sans aucun doute aux recommandations s’il était au courant de leur<br />
importance et de leur raison d’être.<br />
Les chercheurs et décisionnaires ont également beso<strong>in</strong> d’un meilleur accès à l’<strong>in</strong>formation<br />
concernant les espèces envahissantes, leur impact et les moyens de lutte. Pour<br />
limiter l’impact des espèces envahissantes déjà présentes en Suisse, leurs populations<br />
doivent être gérées, c.-à-d. éradiquées ou contrôlées. Parmi les espèces pouvant être<br />
éradiquées, on peut citer le cerf sika (Cervus nippon), le mouflon (Ovis orientalis) et la<br />
tadorne casarca (Tadorna ferrug<strong>in</strong>ea) qui, dans le cas contraire, risquent d’élargir leur<br />
distribution et d’envahir des pays vois<strong>in</strong>s. Un programme national pilote d’éradication<br />
ou de lutte, par exemple contre une plante de la liste noire, est recommandé comme<br />
étude de cas. Il est également recommandé de surveiller les populations de certa<strong>in</strong>es<br />
espèces exotiques pour détecter les hausses souda<strong>in</strong>es de population pouvant <strong>in</strong>diquer<br />
un caractère envahissant. En faisant cela, des programmes de contrôle, ou même<br />
d’éradication, pourraient être décidés avant que les populations ne deviennent totalement<br />
<strong>in</strong>gérables. En accumulant l’<strong>in</strong>formation nécessaire à la rédaction de ce rapport, il<br />
est apparu clairement que trop peu d’<strong>in</strong>formation était disponible sur les EEE en
Résumé 21<br />
Suisse. Il est fortement recommandé d’étudier plus en détails ces espèces, en particulier<br />
leur impact, af<strong>in</strong> de démontrer leur importance aux décisionnaires et politiques.<br />
Les ressources f<strong>in</strong>ancières étant limitées, il est nécessaire de dicter des priorités dans la<br />
lutte contre les EEE et d’allouer ces ressources là où elles auront l’impact le plus<br />
important. Il est nécessaire d’évaluer de manière critique la faisabilité de différentes<br />
méthodes de lutte, et de cibler en priorité les espèces pour lesquelles il n’y a pas de<br />
conflit d’<strong>in</strong>térêt. Par exemple, parmi les plantes de la liste noire, une action contre la<br />
berce du Caucase, une plante de faible <strong>in</strong>térêt ornemental et posant de sérieux problèmes<br />
de santé publique, ne devrait pas susciter d’opposition.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 22<br />
1 > Introduction<br />
The values of societies and the views on issues such as <strong>alien</strong> <strong>species</strong> change over time,<br />
and today the seriousness of <strong>alien</strong> <strong>species</strong> with their detrimental impacts on biodiversity,<br />
economics and human health is widely accepted and management options for<br />
<strong>in</strong>vasive <strong>alien</strong> <strong>species</strong> (IAS) are sought. In the past people did not have the knowledge<br />
we have today and took many familiar <strong>species</strong> with them to new areas they were<br />
settl<strong>in</strong>g. Acclimatization societies were established with the goal of <strong>in</strong>troduc<strong>in</strong>g European<br />
<strong>species</strong> to the new colonies and the newly discovered <strong>species</strong> of exotic lands <strong>in</strong>to<br />
Europe. Thus, people cheered the arrival of the English sparrow <strong>in</strong> North America. The<br />
poet William Cullen Bryant wrote <strong>in</strong> ‘The Old World Sparrow’:<br />
“A w<strong>in</strong>ged settler has taken his place<br />
With Teutons and men of the Celtic race.<br />
He has followed their path to our hemisphere;<br />
The Old World Sparrow at last is here.”<br />
This report compiles published <strong>in</strong>formation about <strong>alien</strong> <strong>species</strong> and other <strong>in</strong>formation<br />
from experts <strong>in</strong> <strong>Switzerland</strong> and abroad on the <strong>alien</strong> flora and fauna of <strong>Switzerland</strong>.<br />
Species occurr<strong>in</strong>g <strong>in</strong> Europe but not (yet) <strong>in</strong> <strong>Switzerland</strong> but known to have <strong>in</strong>vasive<br />
characteristics are also <strong>in</strong>cluded <strong>in</strong> this publication. The fact that they are <strong>in</strong>vasive and<br />
occur <strong>in</strong> neighbour<strong>in</strong>g countries puts them on a Watch List for which monitor<strong>in</strong>g<br />
and/or prevention measures would be advisable.<br />
Names of scientists who wrote chapters are given at the beg<strong>in</strong>n<strong>in</strong>g of each one, but the<br />
many other people whose help was vital <strong>in</strong> prepar<strong>in</strong>g this report are acknowledged at<br />
the end of this chapter.<br />
Availability of lists varies tremendously between taxonomic groups, thus, unfortunately,<br />
it is not possible to list all the <strong>alien</strong> <strong>species</strong> of <strong>Switzerland</strong>. However, for groups<br />
that are well known, complete lists could be assembled. For some other groups only<br />
<strong>in</strong>vasive <strong>species</strong> are given, and some groups are not covered at all. For these latter<br />
groups, lists of native <strong>species</strong> are not even available and the <strong>in</strong>troduced or native status<br />
of many <strong>species</strong> discovered <strong>in</strong> <strong>Switzerland</strong> cannot be demonstrated with certa<strong>in</strong>ty.<br />
These <strong>species</strong> are called cryptogenic. It has to be emphasized that a report on the <strong>alien</strong><br />
<strong>species</strong> of a region can only give a snapshot of the current situation, s<strong>in</strong>ce more <strong>species</strong><br />
will <strong>in</strong>vade and be <strong>in</strong>troduced as time goes by. As long as knowledge about the diversity<br />
and distribution of native taxa is <strong>in</strong>complete, only <strong>in</strong>vasions of taxonomically<br />
uncommon or high-impact <strong>species</strong> will be detected. Invasions of cryptic taxa often go<br />
unnoticed (Müller and Griebeler, 2002). In conclusion, a consistent approach between<br />
taxonomic groups is not possible due to gaps <strong>in</strong> knowledge. The groups are discussed
Résumé 23<br />
<strong>in</strong> this report on the basis of the available knowledge, which reflects difficulties <strong>in</strong><br />
monitor<strong>in</strong>g and taxonomy and also the importance of the different groups.<br />
The taxonomic groups chosen, i.e. vertebrates, <strong>in</strong>sects, crustaceans, arachnids, molluscs,<br />
other animals, plants and fungi, are the basis of separate chapters present<strong>in</strong>g lists<br />
of <strong>alien</strong> <strong>species</strong> and additional <strong>in</strong>formation about them, Fact Sheets for the <strong>in</strong>vasive<br />
<strong>species</strong>, and f<strong>in</strong>ally an evaluation of the status, impacts, pathways, control options and<br />
recommendations for the <strong>alien</strong> <strong>species</strong> of each group. The Fact Sheets summarize<br />
<strong>in</strong>formation on the <strong>in</strong>vasive <strong>species</strong> under the head<strong>in</strong>gs taxonomy, description, ecology,<br />
orig<strong>in</strong>, <strong>in</strong>troduction, distribution, impact, management and references. A special focus<br />
is given to <strong>in</strong>formation on impacts of the <strong>species</strong> on the environment and economy to<br />
<strong>in</strong>dicate their detrimental nature and the importance of manag<strong>in</strong>g them. This <strong>in</strong>formation<br />
is often difficult to f<strong>in</strong>d but crucial to show the true potential threat to <strong>Switzerland</strong><br />
without be<strong>in</strong>g hysterical about it.<br />
All signatories to the Convention on Biological Diversity (CBD), <strong>in</strong>clud<strong>in</strong>g <strong>Switzerland</strong>,<br />
are obliged to prevent the <strong>in</strong>troduction of, control or eradicate those <strong>alien</strong> <strong>species</strong><br />
which threaten ecosystems, habitats or <strong>species</strong>. There are other <strong>in</strong>ternational <strong>in</strong>struments,<br />
ratified by <strong>Switzerland</strong>, call<strong>in</strong>g for management of <strong>in</strong>vasive <strong>species</strong>, e.g. the<br />
Convention on the Conservation of Migratory Species of Wild Animals and the Convention<br />
on the Conservation of European Wildlife and Natural Habitats. This report<br />
can also be used <strong>in</strong> response to the call from the CBD Secretariat for countries to<br />
produce lists of <strong>alien</strong> <strong>species</strong> and can provide <strong>in</strong>formation for a national strategy on<br />
IAS.<br />
The importance of IAS is <strong>in</strong>creas<strong>in</strong>gly recognized <strong>in</strong> Central Europe, <strong>in</strong>dicated by the<br />
recently published book Neobiota <strong>in</strong> Österreich (Essl and Rabitsch, 2002) and the very<br />
recently produced Gebietsfremde Arten; Positionspapier des Bundesamtes für Naturschutz<br />
<strong>in</strong> Germany (Kl<strong>in</strong>genste<strong>in</strong> et al., 2005).<br />
The follow<strong>in</strong>g sections of this chapter expla<strong>in</strong> the def<strong>in</strong>itions used <strong>in</strong> the report, then<br />
present a predom<strong>in</strong>antly global perspective of the IAS issue, with examples from<br />
<strong>Switzerland</strong>. Evaluations of pathways, orig<strong>in</strong>s of the <strong>species</strong>, number of <strong>alien</strong> <strong>species</strong> <strong>in</strong><br />
comparison to native <strong>species</strong>, their impacts, management options and other trends are<br />
given <strong>in</strong> the chapters on the taxonomic groups. An overall evaluation of these factors<br />
was attempted, but it became obvious that that was not a constructive exercise, s<strong>in</strong>ce<br />
the clear trends <strong>in</strong> some taxonomic groups were diluted and comb<strong>in</strong><strong>in</strong>g groups with<br />
very different knowledge bases made it difficult to compare the results. However,<br />
with<strong>in</strong> the groups, trends are often very obvious and for these readers are referred to<br />
the chapters on the taxonomic groups.<br />
1.1 Def<strong>in</strong>itions<br />
Def<strong>in</strong>itions used <strong>in</strong> relation to IAS are as varied as the <strong>species</strong> themselves. The term<br />
‘<strong>in</strong>vasive’, for <strong>in</strong>stance, is used <strong>in</strong> one extreme, to describe a population that is expand-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 24<br />
<strong>in</strong>g (<strong>in</strong>vad<strong>in</strong>g somewhere) to another, for <strong>species</strong> which have a negative impact on<br />
native <strong>species</strong>, ecosystems and habitats. A set of widely used def<strong>in</strong>itions is provided by<br />
the Convention on Biological Diversity, available on their websites (see Article 2, Use<br />
of Terms at: http://www.biodiv.org/convention/articles.asp?lg=0&a+cbd–02 and Annex 2<br />
of the item 8 at: http://www.biodiv.org/doc/meet<strong>in</strong>g.aspx?mtg=sbstta–06&tab=1).<br />
As long as the terms are not harmonized, it is crucial to give the def<strong>in</strong>itions used <strong>in</strong> a<br />
document. Thus, short def<strong>in</strong>itions used <strong>in</strong> this report are presented below. In some<br />
chapters, separate def<strong>in</strong>itions were used for reasons of clarity, and these are given at<br />
the beg<strong>in</strong>n<strong>in</strong>g of the relevant chapters.<br />
> Introduction: the movement, by direct or <strong>in</strong>direct human agency, of a <strong>species</strong> or<br />
lower taxon <strong>in</strong>to a new area. Indirect agency is, for <strong>in</strong>stance, the build<strong>in</strong>g of a canal,<br />
while direct movement <strong>in</strong>cludes deliberate and accidental <strong>in</strong>troductions. This can be<br />
either with<strong>in</strong> <strong>Switzerland</strong> or from another country <strong>in</strong>to <strong>Switzerland</strong>, breach<strong>in</strong>g formerly<br />
<strong>in</strong>surmountable barriers for the <strong>species</strong>. The most obvious natural barriers for<br />
a <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> are the Alps between the Tic<strong>in</strong>o and the rest of the country,<br />
but also the different watersheds, towards the North Sea, the Black Sea and the<br />
Mediterranean Sea.<br />
> Deliberate <strong>in</strong>troduction: the purposeful movement by humans of a <strong>species</strong> <strong>in</strong>to a<br />
new area. This <strong>in</strong>cludes also <strong>species</strong> <strong>in</strong>troduced <strong>in</strong>to conf<strong>in</strong>ement, e.g. aquariums<br />
and zoos. Thus, the <strong>in</strong>troduction is def<strong>in</strong>ed here as the <strong>in</strong>itial movement <strong>in</strong>to a new<br />
area. They can subsequently escape or be released <strong>in</strong>to the environment.<br />
> Accidental <strong>in</strong>troduction: a <strong>species</strong> utiliz<strong>in</strong>g humans or human delivery systems as<br />
vectors. This also <strong>in</strong>cludes contam<strong>in</strong>ants and diseases of deliberately <strong>in</strong>troduced<br />
<strong>species</strong>.<br />
> Native <strong>species</strong>: a <strong>species</strong> liv<strong>in</strong>g with<strong>in</strong> its natural range.<br />
> Alien <strong>species</strong>: a <strong>species</strong> <strong>in</strong>troduced outside its natural distribution.<br />
> <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong> (IAS): an <strong>alien</strong> <strong>species</strong> which threatens ecosystems, habitats<br />
and <strong>species</strong>. These are addressed under Article 8(h) of the CBD.<br />
Some <strong>species</strong> are difficult to handle under these def<strong>in</strong>itions because of a lack of <strong>in</strong>formation,<br />
e.g. it is often not possible to determ<strong>in</strong>e whether a European <strong>species</strong> arrived<br />
with human help or on its own w<strong>in</strong>gs or legs. Thus, these def<strong>in</strong>itions should not be<br />
treated as cast <strong>in</strong> stone but are flexibly used for the purpose of this report.<br />
1.2 <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong> – a global overview<br />
This report can only give a glimpse of the complex issue of IAS, to set the stage for an<br />
assessment of their importance. A good place to start brows<strong>in</strong>g through the IAS literature<br />
is <strong>in</strong> books published on the topic, e.g. the classic Elton (1958), Drake et al.<br />
(1989), Di Castri et al. (1990), Williamson (1996), Mooney and Hobbs (2000), Sh<strong>in</strong>e et<br />
al. (2000), Low (2001), McNeely (2001), Wittenberg and Cock (2001), Bask<strong>in</strong> (2002),<br />
Kowarik and Starf<strong>in</strong>ger (2002), Leppäkoski et al. (2002), Pimentel (2002), Kowarik<br />
(2003), Mooney et al. (2005) and others.
Résumé 25<br />
Changes <strong>in</strong> distribution of <strong>species</strong> are a natural phenomenon; ranges expand and retract<br />
and <strong>species</strong> colonize new areas outside their natural range by long-distance dispersal,<br />
for example reptiles on float<strong>in</strong>g wood to new islands. However, these events are rare<br />
and mostly restricted by natural barriers. The relatively recent globalization of trade<br />
and travel has <strong>in</strong>advertently led to the <strong>in</strong>creased transport of organisms and <strong>in</strong>troduction<br />
of <strong>alien</strong> <strong>species</strong>, tear<strong>in</strong>g down these natural barriers. Alien <strong>species</strong> are not bad per<br />
se, <strong>in</strong> fact many <strong>species</strong> are used for human consumption, e.g. most crop <strong>species</strong> are<br />
<strong>alien</strong>s <strong>in</strong> the majority of regions where they are grown. However, some of them may<br />
subsequently become harmful and pose threats to the environment and human populations.<br />
IAS are <strong>in</strong>creas<strong>in</strong>gly recognized as one of the major threats to biodiversity. These<br />
negative effects are best documented <strong>in</strong> bird ext<strong>in</strong>ctions on islands where the majority<br />
s<strong>in</strong>ce 1800 have been caused by IAS (BirdLife International, 2000). This global problem<br />
needs global and local responses, or, even better, proactive measures, and solutions.<br />
The complexity of the IAS topic stems from the very different <strong>species</strong> <strong>in</strong>volved, their<br />
diverse orig<strong>in</strong>s, the variety of pathways used, their varied impacts on new environments,<br />
their relationships with <strong>in</strong>digenous and other <strong>alien</strong> <strong>species</strong>, the ecosystem<br />
changes caused, their dependence on other factors such as global warm<strong>in</strong>g, their human<br />
dimensions <strong>in</strong>clud<strong>in</strong>g changes <strong>in</strong> political and ethical views, and their on-go<strong>in</strong>g evolution.<br />
<strong>Invasive</strong> <strong>alien</strong> <strong>species</strong> are found <strong>in</strong> virtually every taxonomic group. The follow<strong>in</strong>g<br />
examples will attest this statement.<br />
The West Nile virus caus<strong>in</strong>g encephalitis hitched a ride to the New World <strong>in</strong> an <strong>in</strong>fected<br />
bird, mosquito or human (Enser<strong>in</strong>k, 1999).<br />
The bacterium Vibrio cholerae (Pac<strong>in</strong>i), the causal organism of the human disease<br />
cholera, is a member of brackish water communities and is frequently found <strong>in</strong> ballast<br />
water of ships (McCarthy and Khambaty, 1994), by which means some new highly<br />
virulent stra<strong>in</strong>s have been redistributed lead<strong>in</strong>g to epidemic outbreaks of cholera.<br />
Another bacterium, Erw<strong>in</strong>ia amylovora (see Fact Sheet <strong>in</strong> chapter 9) is a serious threat<br />
to the fruit <strong>in</strong>dustry <strong>in</strong> <strong>Switzerland</strong>.<br />
Some fungal pathogens are amongst the IAS with the most disruptive impacts on<br />
ecosystems. Some well-known examples <strong>in</strong>clude fungi attack<strong>in</strong>g trees, e.g. chestnut<br />
blight (Cryphonectria parasitica (Murrill) Barr) was <strong>in</strong>troduced with <strong>alien</strong> chestnut<br />
trees to North America, where it virtually eradicated the American chestnut (Castanea<br />
dentata (Marsh.) Borkh.), which was a dom<strong>in</strong>ant tree <strong>in</strong> eastern forests, thereby chang<strong>in</strong>g<br />
the entire ecosystem and composition of the forests (Hendrickson, 2002) and is<br />
now, albeit with less severity, attack<strong>in</strong>g trees <strong>in</strong> <strong>Switzerland</strong>.<br />
Weeds are another predom<strong>in</strong>ant group of IAS known to cause economic problems as<br />
well as deleterious effects on the environment. The giant reed (Arundo donax L.) is<br />
used <strong>in</strong> many countries, for example as w<strong>in</strong>dbreaks, and is readily <strong>in</strong>vad<strong>in</strong>g natural<br />
areas; the small herb common ragweed (Ambrosia artemisiifolia L.) is swiftly expand-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 26<br />
<strong>in</strong>g its exotic range <strong>in</strong> Europe caus<strong>in</strong>g severe allergic problems for the human population.<br />
Many different k<strong>in</strong>ds of worms found their way to new areas with human assistance,<br />
especially parasitic worms from the Plathelm<strong>in</strong>thes and Nemathelm<strong>in</strong>thes, for example<br />
the nematode Anguillicola crassus, which attacks the native eel (Anguilla anguilla L.).<br />
A spectacular disaster caused by an <strong>in</strong>troduced snail was the <strong>in</strong>troduction of the carnivorous<br />
rosy wolfsnail (Eugland<strong>in</strong>a rosea (Férussac)) to many subtropical and tropical<br />
islands destroy<strong>in</strong>g the diverse endemic snail faunas. Another example of an <strong>in</strong>troduced<br />
mollusc <strong>in</strong> Europe is perhaps the most aggressive freshwater <strong>in</strong>vader worldwide,<br />
the zebra mussel (Dreissena polymorpha (Pallas)), <strong>in</strong>flict<strong>in</strong>g not only huge economic<br />
costs but also severe biotic changes as it functions as an ecosystem eng<strong>in</strong>eer <strong>species</strong><br />
(Karateyev et al., 2002).<br />
Small <strong>in</strong>troduced crustaceans dom<strong>in</strong>ate the fauna of many rivers and lakes worldwide<br />
due to the <strong>in</strong>creased ship traffic that transports organisms <strong>in</strong> ballast water to new areas,<br />
and also to the creation of canals connect<strong>in</strong>g formerly <strong>in</strong>surmountable natural barriers<br />
between watersheds. Thus, <strong>alien</strong> <strong>species</strong> (ma<strong>in</strong>ly crustaceans and molluscs) dom<strong>in</strong>ate<br />
the Rh<strong>in</strong>e <strong>in</strong> total abundance and produce a biomass of more than 80 % (Haas et al.,<br />
2002).<br />
Interest<strong>in</strong>gly, <strong>in</strong>troduced <strong>in</strong>sects, despite their diversity, have not shown a high potential<br />
for caus<strong>in</strong>g environmental problems, although they can be devastat<strong>in</strong>g pests <strong>in</strong><br />
agriculture and forestry. However, several ant <strong>species</strong> destroy native faunas, especially<br />
on islands – but also, e.g. the Argent<strong>in</strong>e ant (L<strong>in</strong>epithema humile (Mayr)), <strong>in</strong> southern<br />
Europe.<br />
The <strong>in</strong>famous cane toad (Bufo mar<strong>in</strong>us (L.)) is spread<strong>in</strong>g quickly through Australia<br />
feed<strong>in</strong>g on everyth<strong>in</strong>g smaller than itself and poison<strong>in</strong>g the bigger predators, such as<br />
quolls (Dasyurus spp. E. Geoffroy St.-Hilaire).<br />
One of the most devastat<strong>in</strong>g <strong>in</strong>troduced reptiles is the brown tree snake (Boiga irregularis<br />
(Merrem)) on Pacific islands; it arrived on Guam on military equipment and<br />
brought a silent spr<strong>in</strong>g to the island by feast<strong>in</strong>g on the bird <strong>species</strong>. Moreover, it causes<br />
frequent power cuts and is a danger to babies because of its venom.<br />
The Nile perch (Lates niloticus (L.)), <strong>in</strong>troduced <strong>in</strong>to Lake Victoria to improve fisheries,<br />
caused the ext<strong>in</strong>ction of more than 100 fish <strong>species</strong> of the cichlid family; most of<br />
them endemic to the lake – before the predator came it was called an evolutionary<br />
laboratory, afterwards an ecological disaster.<br />
The American ruddy duck (Oxyura jamaicensis (Gmel<strong>in</strong>)) was <strong>in</strong>troduced to England<br />
as an addition to the wildfowl fauna, where it ostensibly does no harm, but then it<br />
spread to Spa<strong>in</strong> where it has swiftly endangered the native close relative, the whiteheaded<br />
duck (O. leucocephala (Scopoli)) by hybridization.
Résumé 27<br />
Feral mammals <strong>in</strong>troduced to islands brought many bird <strong>species</strong> to the br<strong>in</strong>k of ext<strong>in</strong>ction<br />
or beyond by feed<strong>in</strong>g on their eggs and chicks (e.g. Long, 2003).<br />
However, some taxonomic groups seem to <strong>in</strong>clude more <strong>in</strong>vasive <strong>species</strong> than others<br />
do. Mammals are a major threat to island faunas and floras. Whereas rats, mongooses,<br />
mustelids and feral cats devastate the local bird and reptile fauna of islands, feral goats<br />
(Capra hircus L.) can dim<strong>in</strong>ish the native flora drastically. Weeds alter the vegetation<br />
on many archipelagos to the detriment of the entire ecosystem. Island ecosystems,<br />
<strong>in</strong>clud<strong>in</strong>g isolated lakes, are particularly vulnerable to these <strong>in</strong>vaders. However, all<br />
cont<strong>in</strong>ents and habitats seem to be vulnerable to <strong>in</strong>vasions, although there appear to be<br />
some differences between cont<strong>in</strong>ents. In the highly populated area of Central Europe,<br />
IAS seem to be of less importance to biodiversity than on other cont<strong>in</strong>ents with large<br />
tracts of more natural habitats. The smaller reserves <strong>in</strong> Central Europe are easier to<br />
manage and control of <strong>alien</strong> <strong>species</strong> <strong>in</strong> these areas is often more practical. The long<br />
association between <strong>in</strong>troduced <strong>species</strong> and the human population <strong>in</strong> Europe is a very<br />
different situation compared to other cont<strong>in</strong>ents, as all habitats are highly altered and<br />
human-made habitats dom<strong>in</strong>ate. These human-made habitats are often regarded as<br />
valuable heritage <strong>in</strong> Central Europe and some of them are dom<strong>in</strong>ated by <strong>alien</strong> <strong>species</strong><br />
<strong>in</strong>troduced centuries ago.<br />
Compar<strong>in</strong>g the numbers of <strong>species</strong> <strong>in</strong>troduced to <strong>species</strong> that have established and<br />
<strong>species</strong> that have become <strong>in</strong>vasive, one has to bear <strong>in</strong> m<strong>in</strong>d the long lag phases which<br />
often occur. Most <strong>in</strong>troduced <strong>species</strong> take some time before they become <strong>in</strong>vasive, i.e.<br />
enter the log phase. Kowarik (2003) shows that for woody <strong>alien</strong> plants <strong>in</strong> one part of<br />
Germany, the average time lag between first <strong>in</strong>troduction and expansion is about 147<br />
years. The occurrence of time lags makes predictions on <strong>in</strong>vasiveness of <strong>alien</strong> <strong>species</strong><br />
very difficult. A <strong>species</strong> do<strong>in</strong>g no harm today can still become an <strong>in</strong>vasive of tomorrow,<br />
especially <strong>in</strong> comb<strong>in</strong>ation with other global changes. There are three major factors<br />
that determ<strong>in</strong>e the ability of a <strong>species</strong> to become <strong>in</strong>vasive:<br />
1. Intr<strong>in</strong>sic factors or <strong>species</strong> traits, such as the ability to adapt to different conditions, a<br />
wide tolerance of abiotic factors, pre-adaptations to different climatic zones, and a<br />
high reproductive rate.<br />
2. Extr<strong>in</strong>sic factors or relationships between the <strong>species</strong> and abiotic and biotic factors,<br />
such as the number of natural enemies, the number of compet<strong>in</strong>g <strong>species</strong> (native and<br />
<strong>alien</strong>), other <strong>in</strong>teractions with native and <strong>alien</strong> <strong>species</strong> (poll<strong>in</strong>ation, dispersal, food<br />
source, ecosystem eng<strong>in</strong>eers), climatic conditions, soil conditions, degree of disturbance<br />
(natural and human-<strong>in</strong>duced), global climate change, change <strong>in</strong> land-use patterns,<br />
and control and eradication of other IAS.<br />
3. Human dimension. The attractiveness and importance of <strong>species</strong> to humans <strong>in</strong>fluence<br />
<strong>in</strong>troduction pathways, vectors, the number of specimens <strong>in</strong>troduced, the number<br />
of <strong>in</strong>troductions, and the potential for eradication or control.<br />
Predictions about the <strong>in</strong>vasive potential of a <strong>species</strong> prior to <strong>in</strong>troduction, as now made<br />
<strong>in</strong> Australia and New Zealand, rema<strong>in</strong> difficult, despite recent progress <strong>in</strong> science. The<br />
best <strong>in</strong>dicator is still whether a <strong>species</strong> has become <strong>in</strong>vasive <strong>in</strong> a similar area elsewhere<br />
(i.e. its <strong>in</strong>vasion history).
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 28<br />
1.3 Status of <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong><br />
As po<strong>in</strong>ted out above, the accelerat<strong>in</strong>g pace of movement of people and goods is<br />
<strong>in</strong>creas<strong>in</strong>g the number of <strong>in</strong>troductions of already established <strong>species</strong> and new arrivals.<br />
Therefore lists of <strong>alien</strong> <strong>species</strong> cannot be comprehensive, but give only a snapshot of<br />
the current knowledge. Moreover, the lack of both knowledge and taxonomists for<br />
some groups precludes lists for such groups be<strong>in</strong>g presented.<br />
The situation regard<strong>in</strong>g IAS <strong>in</strong> <strong>Switzerland</strong> is similar to that <strong>in</strong> other Central European<br />
countries, <strong>in</strong> particular Austria, which is also a land-locked country conta<strong>in</strong><strong>in</strong>g part of<br />
the Alps. Some of the taxonomic sections present numbers of <strong>alien</strong> <strong>species</strong> recorded <strong>in</strong><br />
other countries for comparison. A comparison between countries for all taxa cannot be<br />
given here, s<strong>in</strong>ce the def<strong>in</strong>itions and criteria used differ considerably between country<br />
reports and make it impossible to use the figures for comparison with neighbour<strong>in</strong>g<br />
countries. The different levels of knowledge for the different groups and countries is<br />
another source of <strong>in</strong>accuracy. Some features specific to <strong>Switzerland</strong> will allow a different<br />
guild of <strong>species</strong> to <strong>in</strong>vade. The deep, cold lakes, for example, are probably prevent<strong>in</strong>g<br />
or at least reduc<strong>in</strong>g <strong>in</strong>vasions by aquatic weeds that thrive <strong>in</strong> warm, shallow water.<br />
This report lists about 800 <strong>alien</strong> <strong>species</strong> and describes 107 IAS <strong>in</strong> Fact Sheets, i.e. five<br />
mammals, four birds, one reptile, three amphibians, seven fish, four molluscs, 16<br />
<strong>in</strong>sects, six crustaceans, three spiders, two ‘worms’, seven fungi, one bacteria, and 48<br />
plants. As mentioned above, some of these <strong>species</strong>, e.g. the grey squirrel (Sciurus<br />
carol<strong>in</strong>ensis Gmel<strong>in</strong>), have not yet entered <strong>Switzerland</strong>, but their arrival <strong>in</strong> the near<br />
future is likely.<br />
1.4 Pathways<br />
Pathways can be divided <strong>in</strong>to those for <strong>species</strong> deliberately <strong>in</strong>troduced and those for<br />
<strong>species</strong> accidentally moved around. Examples of <strong>species</strong> <strong>in</strong> the first category are<br />
<strong>species</strong> used <strong>in</strong> aquaculture, for fisheries, as forest trees, for agricultural purposes, for<br />
hunt<strong>in</strong>g, and plants used for soil improvement and solely to please humans as ornamentals.<br />
Most of these can also transport hitchhik<strong>in</strong>g <strong>species</strong> and people can accidentally<br />
<strong>in</strong>troduce <strong>species</strong> on cargo, their boots, etc. To summarize, most aquatics and <strong>in</strong>vertebrates<br />
<strong>in</strong> general, are accidental arrivals, whereas most plants and vertebrates are<br />
deliberately <strong>in</strong>troduced. M<strong>in</strong>ch<strong>in</strong> and Gollasch (2002) and Carlton and Ruiz (2005)<br />
give excellent accounts of pathways and vectors <strong>in</strong> more depth. The latter divide<br />
pathways <strong>in</strong>to cause (why a <strong>species</strong> is transported), route (the geographical path) and<br />
vector (how a <strong>species</strong> is transported).<br />
The global trend that vertebrates and plants are ma<strong>in</strong>ly deliberately <strong>in</strong>troduced also<br />
holds true for <strong>Switzerland</strong>. 75 % (15) of the 20 Black List plants were <strong>in</strong>troduced<br />
chiefly as ornamentals and 35 of the 37 vertebrates have been deliberately <strong>in</strong>troduced.<br />
In general, the <strong>in</strong>troduction pathways of <strong>species</strong> <strong>in</strong>to Europe, rather than <strong>in</strong>to <strong>Switzerland</strong>,<br />
have been analysed <strong>in</strong> this report, s<strong>in</strong>ce some <strong>species</strong> arrived <strong>in</strong> <strong>Switzerland</strong> by
Résumé 29<br />
expand<strong>in</strong>g their populations from neighbour<strong>in</strong>g countries. In conclusion, many serious<br />
<strong>in</strong>vaders were deliberately <strong>in</strong>troduced, often without much <strong>in</strong>centive beyond the wish<br />
to colour the landscape, e.g. with ornamental plants and waterfowl.<br />
Prediction and prevention of new <strong>in</strong>vaders is difficult s<strong>in</strong>ce trade <strong>in</strong> <strong>species</strong> can be<br />
changed by chang<strong>in</strong>g demands. A group not currently a problem may yet become<br />
problematic <strong>in</strong> the near future. If, for example, the pet trade <strong>in</strong> reptiles shifts from<br />
predom<strong>in</strong>antly tropical and subtropical <strong>species</strong> to <strong>species</strong> orig<strong>in</strong>at<strong>in</strong>g <strong>in</strong> regions <strong>in</strong> the<br />
same latitudes as Europe, such as North America and Ch<strong>in</strong>a, the situation could become<br />
much worse.<br />
1.5 Impacts of <strong>in</strong>vasive <strong>alien</strong> <strong>species</strong><br />
The impacts of IAS are often considerable, <strong>in</strong> particular where ecosystem function<strong>in</strong>g<br />
is be<strong>in</strong>g altered or <strong>species</strong> are be<strong>in</strong>g pushed to ext<strong>in</strong>ction, as has been shown for many<br />
bird <strong>species</strong>. Most ext<strong>in</strong>ctions are likely to be caused by a comb<strong>in</strong>ation of factors and<br />
are not attributable to a s<strong>in</strong>gle cause, but there is no doubt that IAS play an important<br />
role. The environmental impacts can be divided <strong>in</strong>to four major factors:<br />
> competition,<br />
> predation (<strong>in</strong>clud<strong>in</strong>g herbivory), and more subtle <strong>in</strong>teractions such as<br />
> hybridization and<br />
> transmission of diseases.<br />
All these factors alone or <strong>in</strong> concert with other factors can decrease biodiversity and<br />
cause ext<strong>in</strong>ction. The most obvious examples for competition are between <strong>in</strong>troduced<br />
and native plants for nutrients and exposure to sunlight. Resource competition has also<br />
led to the replacement of the native red squirrel (Sciurus vulgaris L.) by the <strong>in</strong>troduced<br />
American grey squirrel (S. carol<strong>in</strong>ensis) <strong>in</strong> almost all of Great Brita<strong>in</strong> and it is predicted<br />
that this trend will cont<strong>in</strong>ue on the cont<strong>in</strong>ent. The latter forages more efficiently<br />
for food and is stronger than the native <strong>species</strong> (Williamson, 1996). Impacts due to<br />
predation and herbivory are very extensive on island fauna and flora, as mentioned<br />
above. The brown tree snake elim<strong>in</strong>ated most of the bird <strong>species</strong> on Guam, and feral<br />
goats are a menace to native vegetation on islands, where they were often released as a<br />
liv<strong>in</strong>g food resource. A well-known example of hybridization from Europe is the ruddy<br />
duck, which hybridizes with the native white-headed duck, as mentioned above. In<br />
some cases IAS can harbour diseases and act as a vector for those diseases to native<br />
<strong>species</strong>. This is the case with American <strong>in</strong>troduced crayfish <strong>species</strong> to Europe, which<br />
are almost asymptomatic carriers of the crayfish plague (Aphanomyces astaci Schikora),<br />
but the native noble crayfish (Astacus astacus (L.)) is highly susceptible to the<br />
disease, and thus is struggl<strong>in</strong>g to coexist with populations of the American crayfish<br />
<strong>species</strong>. Introduced <strong>species</strong> can <strong>in</strong>teract with natives <strong>in</strong> a variety of ways and <strong>in</strong>direct<br />
effects can be very difficult to demonstrate. Direct and <strong>in</strong>direct effects can lead to very<br />
complex <strong>in</strong>teractions and a comb<strong>in</strong>ation of effects can cause complex impacts.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 30<br />
In addition to impacts on biodiversity, many IAS impose enormous economic costs.<br />
These costs can arise through direct losses of agricultural and forestry products and<br />
through <strong>in</strong>creased production costs associated with control measures (US Congress,<br />
1993; Pimentel et al., 2000). One often-cited example is the costs <strong>in</strong>flicted on water<br />
plants by the zebra mussel, which clogs water pipes and other structures <strong>in</strong> the Great<br />
Lakes <strong>in</strong> North America. Costs for environmental problems are more difficult to calculate<br />
than costs imposed <strong>in</strong> the agricultural and other economic sectors. A North American<br />
study calculated costs of US$ 138 billion per annum to the USA from IAS (Pimentel<br />
et al., 2000). Some of the costs given <strong>in</strong> this paper are estimates rather than actual,<br />
however, even give or take an order of magnitude, it is still an enormous figure and<br />
shows the importance of IAS. A recently released report (S<strong>in</strong>den et al., 2004) estimates<br />
that weeds are cost<strong>in</strong>g agriculture <strong>in</strong> Australia about Aus$ 4 billion a year, around 20 %<br />
of which is borne by the consumer, the other 80 % by the producer. Accord<strong>in</strong>g to this<br />
report, costs associated with lost production and controll<strong>in</strong>g weeds are equivalent to<br />
0.5 % of gross domestic product, or 14 % of the value of agricultural production <strong>in</strong><br />
Australia. The Aus$ 4 billion estimate is conservative, as it does not <strong>in</strong>clude the impact<br />
on the natural environment, the effect of pollen from the weeds on human health, and<br />
the cost of volunteer weed control. It also hides the comb<strong>in</strong>ed Aus$ 1.8 billion cost to<br />
the economy of sal<strong>in</strong>ity and soil acidity. The amount spent on weed control rose by<br />
Aus$ 68,000 for each additional native plant that was threatened by the <strong>in</strong>vasive weeds.<br />
Weed control <strong>in</strong> natural environments cost about Aus$ 20 million <strong>in</strong> the year from mid<br />
2001 to mid 2002. Turn<strong>in</strong>g to Europe, the costs of Heracleum mantegazzianum Sommier<br />
et Levier (giant hogweed) <strong>in</strong> Germany are estimated at € 10 million; € 1 million<br />
each to the environment and health sectors, and the rema<strong>in</strong>der represents costs to the<br />
economy.<br />
Some IAS also have implications for human health. Giant hogweed was <strong>in</strong>troduced<br />
from the Caucasus to Europe as an ornamental plant. It produces copious amounts of a<br />
sap conta<strong>in</strong><strong>in</strong>g phototoxic substances (furanocoumar<strong>in</strong>s), which can lead to severe<br />
burns to the sk<strong>in</strong>. Regularly children, <strong>in</strong> particular, are hospitalized after contact with<br />
the plant, especially when they have been play<strong>in</strong>g with the hollow stems and petioles,<br />
us<strong>in</strong>g them as ‘peashooters’. The raccoon dog (Nyctereutes procyonoides Gray), <strong>in</strong>troduced<br />
as a fur animal, can, like the native red fox (Vulpes vulpes (L.)), act as a vector<br />
of the most dangerous parasitic disease vectored by mammals to humans <strong>in</strong> Central<br />
Europe, i.e. the fox tapeworm (Ech<strong>in</strong>ococcus multilocularis Leuckart) (Thiess et al.,<br />
2001). Although the raccoon dog is only an additional vector, this can have effects on<br />
the population dynamics of the parasite and lead to an <strong>in</strong>crease <strong>in</strong> the disease <strong>in</strong> humans.<br />
Known impacts of <strong>species</strong> <strong>in</strong>troduced <strong>in</strong>to <strong>Switzerland</strong> are presented <strong>in</strong> the chapters on<br />
the taxonomic groups and their Fact Sheets. Some recent <strong>in</strong>vaders are without demonstrated<br />
impact <strong>in</strong> <strong>Switzerland</strong>; <strong>in</strong> these cases impacts assessed <strong>in</strong> other countries are<br />
given. These provide a glimpse of the possible future. An example is Procambarus<br />
clarkii Girard (an American crayfish <strong>species</strong>) which has caused considerable economic<br />
losses <strong>in</strong> Italy (Gherardi et al., 1999). Demonstration of impacts is often difficult<br />
because of the complexity of ecosystems, but <strong>alien</strong> <strong>species</strong> occurr<strong>in</strong>g <strong>in</strong> high numbers,<br />
such as Reynoutria japonica Houtt. (Japanese knotweed) totally cover<strong>in</strong>g the banks of
Résumé 31<br />
a stream <strong>in</strong> a valley, or an animal biomass of <strong>alien</strong> <strong>species</strong> of up to 95 % <strong>in</strong> the Rh<strong>in</strong>e<br />
near Basel, must have impacts on the native ecosystem. All <strong>species</strong> use resources and<br />
are resources to other creatures and so they alter the web and nutrient flow of the<br />
ecosystems they are liv<strong>in</strong>g <strong>in</strong>.<br />
1.6 Discussion<br />
This report compiles the available <strong>in</strong>formation about IAS <strong>in</strong> <strong>Switzerland</strong>. Dur<strong>in</strong>g the<br />
compilation it became clear that it is impossible to list all <strong>alien</strong> organisms, for the<br />
simple reason that there is a gap <strong>in</strong> taxonomic knowledge, not only <strong>in</strong> <strong>Switzerland</strong>, but<br />
Europe and the world, reflect<strong>in</strong>g a need for more taxonomic work. There are probably<br />
more than ten times the <strong>species</strong> described so far actually exist<strong>in</strong>g right now on the<br />
planet, and for the <strong>species</strong> that have been described we often know little more than<br />
their names. This gives a good <strong>in</strong>dication that there will be more <strong>species</strong> of both native<br />
and <strong>alien</strong> orig<strong>in</strong> found <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> the years to come. S<strong>in</strong>ce all knowledge and<br />
management are based on the concept of ‘<strong>species</strong>’, it seems obvious that taxonomic<br />
work is crucial.<br />
Despite the widespread view that IAS are of less concern <strong>in</strong> Central Europe than on<br />
other cont<strong>in</strong>ents (and more especially on islands) for various reasons, <strong>in</strong>clud<strong>in</strong>g the<br />
small size of nature reserves and the high human impact on all ‘natural’ environments,<br />
and the long association of <strong>alien</strong> <strong>species</strong> and humans lead<strong>in</strong>g to adaptation, however,<br />
the number of cases of dramatic impacts is <strong>in</strong>creas<strong>in</strong>g and the awareness among scientists<br />
and the public is steadily evolv<strong>in</strong>g. Thus, the threats from IAS should not be<br />
underrated. One of the major consequences, which is undoubtedly unfold<strong>in</strong>g before our<br />
eyes, is global homogenization (catchily called McDonaldization), with the unique<br />
character of places such as <strong>Switzerland</strong> be<strong>in</strong>g lost, the characteristic flora and fauna<br />
<strong>in</strong>vaded by organisms which often accomplish to form the largest biomasses <strong>in</strong> certa<strong>in</strong><br />
ecosystems. This is fact and cannot be argued about, while confirmation of impacts is<br />
difficult to obta<strong>in</strong> and can be controversial. The concept of ‘bad <strong>species</strong>’ and ‘good<br />
<strong>species</strong>’ <strong>in</strong> our ecosystems is a rather anthropomorphic view. Even if a population of a<br />
native <strong>species</strong> is enhanced, it is not necessarily good for the ecosystem but can disturb<br />
the natural balances and nutrient and energy flows <strong>in</strong> the ecosystem. After an era of<br />
exploitation, it is now time to address global warm<strong>in</strong>g and global swarm<strong>in</strong>g.<br />
IAS should not be seen as a specific topic but rather as a part of conservation, trade,<br />
and other activities. The ma<strong>in</strong> question to be answered is not how to deal with IAS, but<br />
what should <strong>Switzerland</strong> look like, what are the goals for conservation, what should a<br />
particular nature reserve or the national park look like? These goals need to be set and<br />
IAS management will be part of this bigger picture to conserve and re<strong>in</strong>state the unique<br />
ecosystems and habitats of <strong>Switzerland</strong>. The <strong>in</strong>tact ecosystems can deliver ecosystem<br />
services <strong>in</strong> a susta<strong>in</strong>able manner.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 32<br />
1.7 Recommendations<br />
This section gives some important advice and recommendations, which emerged while<br />
compil<strong>in</strong>g the report. Recommendations for groups of <strong>in</strong>vasives and specific <strong>species</strong><br />
are given <strong>in</strong> more detail <strong>in</strong> the respective chapters and Fact Sheets.<br />
Prepar<strong>in</strong>g a national strategy aga<strong>in</strong>st IAS is recommended to deal with IAS <strong>in</strong> an<br />
appropriate way and as demanded by the CBD. This action plan should identify the<br />
agency responsible for assess<strong>in</strong>g the risks posed by <strong>in</strong>troductions, provide fund<strong>in</strong>g<br />
mechanisms and technical advice and support for control options. This is necessary not<br />
only to fulfil <strong>in</strong>ternational commitments, but also to protect <strong>Switzerland</strong>’s ecosystems<br />
from the detrimental effects of future <strong>in</strong>vasions.<br />
Prevention measures aga<strong>in</strong>st further bio<strong>in</strong>vasions need to be put <strong>in</strong> place to stem the<br />
tide of <strong>in</strong>com<strong>in</strong>g new <strong>species</strong> arriv<strong>in</strong>g accidentally with trade and travel or <strong>in</strong>troduced<br />
deliberately for various purposes. New deliberate <strong>in</strong>troductions must be assessed as to<br />
the threat they may present and only <strong>in</strong>troduced on the basis of a risk analysis and<br />
environmental impact assessment. Scrut<strong>in</strong>iz<strong>in</strong>g imported goods can m<strong>in</strong>imize accidental<br />
<strong>in</strong>vasions. The report <strong>in</strong>dicates some important pathways for consideration. Most of<br />
the known <strong>in</strong>vasive <strong>species</strong>, apart from <strong>in</strong>sects, are deliberately <strong>in</strong>troduced, <strong>in</strong> particular<br />
as ornamentals and for fish<strong>in</strong>g or hunt<strong>in</strong>g. The use of native plants and non-<strong>in</strong>vasive<br />
<strong>alien</strong> plants for garden<strong>in</strong>g should be promoted. Every meadow ploughed, pond filled,<br />
or forest cut for roads, homes or <strong>in</strong>dustrial zones leaves less space for the native flora<br />
and fauna. Gardens and parks support<strong>in</strong>g native plants could be a great boost for native<br />
biodiversity.<br />
Laws regulat<strong>in</strong>g trade <strong>in</strong> plant <strong>species</strong> on the Black List would be a first step <strong>in</strong> the<br />
right direction to reduce the impact of these <strong>species</strong> on the environment, economics<br />
and human health. However, restrictions for <strong>species</strong> already widely distributed with<strong>in</strong><br />
<strong>Switzerland</strong> will not drastically change the situation unless the populations already<br />
present are eradicated or controlled. Arrivals of new <strong>species</strong> with potential for be<strong>in</strong>g<br />
<strong>in</strong>vasive need to be targeted well before they secure a beachhead and become unmanageable.<br />
Fish <strong>in</strong>troductions are regulated by the Fisheries Act, which names <strong>species</strong> for<br />
which an authorization for release is needed and <strong>species</strong> for which release is prohibited<br />
altogether. This is a good basis, although the law could be better adapted to the current<br />
situation. This is described <strong>in</strong> the chapter on fish. The aquarium and terrarium trade is<br />
another important sector that could be more strictly regulated to stop releases of pets<br />
<strong>in</strong>to the wild. With the <strong>in</strong>creas<strong>in</strong>g prohibition of the red-eared slider (Trachemys<br />
scripta Seidel) <strong>in</strong> Europe, other <strong>species</strong> will probably replace it <strong>in</strong> the pet trade. These<br />
<strong>species</strong> are not regulated and, if they are from a more temperate region, they would be<br />
of even more concern than the red-eared slider.<br />
A major problem with <strong>in</strong>troductions of IAS is that the costs when they establish and<br />
become <strong>in</strong>vasive are borne by the public, while the f<strong>in</strong>ancial <strong>in</strong>centives for <strong>in</strong>troduc<strong>in</strong>g<br />
them lie with <strong>in</strong>dividuals or specific bus<strong>in</strong>esses. Development of economic tools that<br />
shift the burden of IAS to those who benefit from <strong>in</strong>ternational trade and travel is a<br />
neglected approach. This is also called the ‘polluter pays’ pr<strong>in</strong>ciple. Appropriate tools
Résumé 33<br />
would be fees and taxes, <strong>in</strong>clud<strong>in</strong>g fees levied on those who import the organisms or<br />
goods. The money raised can be used for prevention and management of IAS.<br />
Awareness rais<strong>in</strong>g is a significant tool <strong>in</strong> the prevention and management of IAS, s<strong>in</strong>ce<br />
some people would adhere to advice if they knew about its importance and the reason<br />
for it, e.g. ‘free<strong>in</strong>g’ of pets <strong>in</strong>to the environment. Scientists and decision-makers also<br />
need better access to <strong>in</strong>formation about <strong>in</strong>vasive <strong>species</strong>, their impacts, and management<br />
options. This lack of organization of <strong>in</strong>formation can be addressed by databases<br />
or global compendia. This report is a basis for collat<strong>in</strong>g <strong>in</strong>formation on published<br />
eradication and management methods for harmful <strong>species</strong> occurr<strong>in</strong>g <strong>in</strong> <strong>Switzerland</strong><br />
(<strong>in</strong>clud<strong>in</strong>g key contacts).<br />
Some key <strong>in</strong>vasive groups to watch for <strong>in</strong> the future are plants, vertebrates, diseases<br />
and some <strong>in</strong>vertebrates, as the most <strong>in</strong>vasive <strong>species</strong> belong to these groups. Introductions<br />
of crayfish <strong>species</strong>, for <strong>in</strong>stance, should be regulated; thus the European native<br />
<strong>species</strong> are of concern, although so far most attention is focussed on the American<br />
<strong>species</strong>.<br />
To address the impacts of some <strong>in</strong>vasive <strong>species</strong> already present <strong>in</strong> <strong>Switzerland</strong>, their<br />
populations need to be managed: either eradicated or controlled.<br />
Monitor<strong>in</strong>g for populations of some <strong>alien</strong> <strong>species</strong> is recommended to detect any sudden<br />
explosion <strong>in</strong> their populations and to watch for their potential <strong>in</strong>vasiveness. By do<strong>in</strong>g<br />
this, control or even eradication efforts can be employed before the populations become<br />
unmanageable. In most cases, the transition from a ‘sleeper’ <strong>species</strong> to an <strong>in</strong>vasive<br />
<strong>species</strong> will go undetected without vigilant monitor<strong>in</strong>g.<br />
Despite the efforts <strong>in</strong> this report to document impacts of <strong>alien</strong> <strong>species</strong> which have<br />
<strong>in</strong>vasive characteristics, much <strong>in</strong>formation is still needed to assess the different impacts,<br />
direct and <strong>in</strong>direct, to the native biodiversity. More studies on <strong>alien</strong> <strong>species</strong> are<br />
highly recommended to ascerta<strong>in</strong> the importance of IAS and to demonstrate their<br />
significance to policy-makers and politicians.<br />
The first Swiss-wide efforts to combat the populations of selected IAS are recommended.<br />
The most efficient control options need to be explored and implemented.<br />
Some <strong>species</strong> of high concern, such as giant hogweed, seem to be good targets for<br />
eradication campaigns throughout <strong>Switzerland</strong>. Populations of giant hogweed are<br />
currently explod<strong>in</strong>g all over Europe (see, e.g., Kowarik (2003) for Germany and Czech<br />
Republic), they cause direct harm to people, and successful eradication has been<br />
achieved us<strong>in</strong>g mechanical and chemical control.<br />
Priority sett<strong>in</strong>g is always difficult. The limited resources need to be spent where they<br />
have the greatest impact <strong>in</strong> combat<strong>in</strong>g IAS. Important po<strong>in</strong>ts, for example, are to<br />
consider the feasibility of an approach (will the goal be reached?) and to target <strong>species</strong><br />
with no conflict of <strong>in</strong>terest. Opposition to action aga<strong>in</strong>st less-important ornamentals on<br />
the Black List and <strong>species</strong> of direct human health concern (giant hogweed) will be<br />
negligible. Other possible targets for eradication are a deer <strong>species</strong> (sika deer – Cervus<br />
nippon Temm<strong>in</strong>ck) or the mouflon (Ovis orientalis Gmel<strong>in</strong>), both <strong>species</strong> with a weak<br />
lobby.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 34<br />
1.8 Acknowledgements<br />
We would like to thank Sarah Brunel, Serge Buholzer, Laura Celesti-Grapow, Joao<br />
Dom<strong>in</strong>gues de Almeida, Helena Freitas, Jacques Maillet, Stefan Toepfer, Eduardo<br />
Sobr<strong>in</strong>o Vesper<strong>in</strong>as, Gerhard Baechli, Hannes Bauer, Claude Besuchet, Wolfgang<br />
Billen, Rudolf Bryner, Daniel Burckhardt, Anne Freitag, Yves Gonseth, Jean-Paul<br />
Haenni, Bernhard Merz, Ladislaus Reser, Bernhard Seifert, Uwe Starf<strong>in</strong>ger, Petr<br />
Pyšek, Baz Hughes, Ulrike Doyle, Simon Capt and Annette Otte for provid<strong>in</strong>g <strong>in</strong>formation<br />
on <strong>alien</strong> <strong>species</strong> <strong>in</strong> Europe and their help <strong>in</strong> the compilation of the list of <strong>alien</strong><br />
<strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>, as well as many more Swiss and European colleagues who<br />
helped us by send<strong>in</strong>g repr<strong>in</strong>ts or giv<strong>in</strong>g advice. We are grateful to the Swiss Commission<br />
for Wild Plant Conservation (CPS-SKEW) group for their <strong>in</strong>formation on <strong>in</strong>vasive<br />
<strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>. The work on <strong>alien</strong> <strong>in</strong>sects by Marc Kenis was partially supported<br />
by the ALARM project (EU 6 th Framework Programme – No. 506675). We<br />
thank also Tanja Re<strong>in</strong>hold, Beate Kiefer, Myriam Poll and Ghisla<strong>in</strong>e Cortat for compil<strong>in</strong>g<br />
<strong>in</strong>formation and the literature and Matthew Cock, Kurt Grossenbacher, Daniel<br />
Hefti, Robert Cowie, Christoph Vorburger, Christoph Scheidegger, Rudolf Müller,<br />
Piero Genovesi, Francis Cordillot, Harry Evans and Rebecca Murphy who greatly<br />
improved earlier drafts with their <strong>in</strong>valuable comments. F<strong>in</strong>ally, we want to express<br />
deep appreciation to Hans Hosbach from the Bundesamt für Umwelt, Wald und Landschaft<br />
(BUWAL) for his immense help and f<strong>in</strong>ancial support to prepare this report.
1 > Introduction 35<br />
References<br />
Bask<strong>in</strong>, Y. (2002) A plague of rats and rubberv<strong>in</strong>es. Island Press,<br />
Wash<strong>in</strong>gton, DC, 377 pp.<br />
BirdLife International (2000) Threatened birds of the world. Lynx Edicions<br />
and BirdLife International, Barcelona and Cambridge, UK, 852 pp.<br />
Carlton, J.T. and G.M. Ruiz (2005) Vector science and <strong>in</strong>tegrated vector<br />
management <strong>in</strong> bio<strong>in</strong>vasion ecology: Conceptual frameworks. In: Mooney,<br />
H.A., McNeely, J., Neville, L.E., Schei, P.J. and Waage, J.K. (eds) <strong>Invasive</strong><br />
<strong>alien</strong> <strong>species</strong>: search<strong>in</strong>g for solutions. Island Press, Covelo, California (<strong>in</strong><br />
press).<br />
Di Castri, F., Hansen, A.J. and M. Debusche (eds) Biological <strong>in</strong>vasions <strong>in</strong><br />
Europe and the Mediterranean bas<strong>in</strong>. Kluwer Academic Publishers,<br />
Dordrecht, Boston, London, 463 pp.<br />
Drake, J.A., Mooney, H.A., di Castri, F., Groves, R.H., Kruger, F.J.,<br />
Rejmánek, M. and M. Williamson (eds) Biological <strong>in</strong>vasions. A global<br />
perspective. John Wiley & Sons, Chichester, New York, Brisbane, Toronto,<br />
S<strong>in</strong>gapore, pp. 257–280.<br />
Elton, C.S. (1958) The ecology of <strong>in</strong>vasions by animals and plants.<br />
Chapman and Hall Ltd., London, 181 pp.<br />
Enser<strong>in</strong>k, M. (1999) Biological <strong>in</strong>vaders sweep <strong>in</strong>. Science 285, 1834–<br />
1836.<br />
Essl, F. and W. Rabitsch (2002) Neobiota <strong>in</strong> Österreich. Federal<br />
Environment Agency, 432 pp.<br />
Gherardi F., Baldacc<strong>in</strong>i, G.N., Barbaresi, S., Ercol<strong>in</strong>i, P., De Luise, G.,<br />
Mazzoni D. and M. Mori (1999) Alien crayfish: the situation of Italy.<br />
Crustacean Issues 11, 107–128.<br />
Haas, G., Brunke, M. and B. Streit (2002) Fast turnover <strong>in</strong> dom<strong>in</strong>ance of<br />
exotic <strong>species</strong> <strong>in</strong> the river Rh<strong>in</strong>e determ<strong>in</strong>es biodiversity and ecosystem<br />
function: An affair between amphipods and mussels. In: Leppäkoski, E.,<br />
Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic <strong>species</strong> of Europe.<br />
Distribution, impacts and management. Kluwer Academic Publishers,<br />
Dordrecht, Boston, London, pp. 426–432.<br />
Hendrickson, O. (2002) <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong> <strong>in</strong> Canadian forests. In:<br />
Claudi, R., Nantel, P. and Muckle-Jeffs, E. (eds) Alien <strong>in</strong>vaders <strong>in</strong> Canada’s<br />
waters, wetlands, and forests. Canadian Forest Service, Ottawa, Canada,<br />
320 pp.<br />
Karateyev, A.Y., Burlakova, L.E. and D.K. Padilla (2002) Impacts of zebra<br />
mussels on aquatic communities and their role as ecosystem eng<strong>in</strong>eers.<br />
In: Leppäkoski, E., Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic<br />
<strong>species</strong> of Europe. Distribution, impacts and management. Kluwer<br />
Academic Publishers, Dordrecht, Boston, London, pp. 433–446.<br />
Kl<strong>in</strong>genste<strong>in</strong>, F., Kornacker, P.M., Martens, H. and U. Schippmann (2005)<br />
Gebietsfremde Arten; Positionspapier des Bundesamtes für Naturschutz.<br />
BfN, Bonn, 30 pp.<br />
Kowarik, I. (2003) Biologische Invasionen: Neophyten und Neozoen <strong>in</strong><br />
Mitteleuropa. Verlag Eugen Ulmer, Stuttgart, Germany, 380 pp.<br />
Kowarik, I. and U. Starf<strong>in</strong>ger (eds) Biologische Invasionen:<br />
Herausforderung zum Handeln? Lentz-Druck, Berl<strong>in</strong>, Germany, pp. 5–24.<br />
Leppäkoski, E., Gollasch, S. and S. Olen<strong>in</strong> (eds) <strong>Invasive</strong> aquatic <strong>species</strong> of<br />
Europe. Distribution, impacts and management. Kluwer Academic<br />
Publishers, Dordrecht, pp. 412–425.<br />
Long, J.L. (2003) Introduced mammals of the world. Their history,<br />
distribution and <strong>in</strong>fluence. CABI Publish<strong>in</strong>g, Wall<strong>in</strong>gford, UK, 589 pp.<br />
Low, T. (2001) Feral future; the untold story of Australia’s exotic <strong>in</strong>vaders.<br />
Pengu<strong>in</strong>s Books Australia Ltd., Victoria, Australia, 394 pp.<br />
McCarthy, S.A. and F.M. Khambaty (1994) International dissem<strong>in</strong>ation of<br />
epidemic Vibrio cholerae by cargo ship ballast and other non-potable<br />
waters. Applied Environmental Microbiology 60, 2597–2601.<br />
McNeely J.A. (ed.) (2001) The great reshuffl<strong>in</strong>g: human dimensions of <strong>in</strong>vasive<br />
<strong>alien</strong> <strong>species</strong>. IUCN, Gland, <strong>Switzerland</strong> and Cambridge, UK, 242 pp.<br />
M<strong>in</strong>ch<strong>in</strong>, D. and S. Gollasch (2002) Vectors – How exotics get around. In:<br />
Leppäkoski, E., Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic <strong>species</strong><br />
of Europe. Distribution, impacts and management. Kluwer Academic<br />
Publishers, Dordrecht, Boston, London, pp. 183–192.<br />
Mooney, H.A. and R.J. Hobbs (eds) <strong>Invasive</strong> <strong>species</strong> <strong>in</strong> a chang<strong>in</strong>g world.<br />
Island Press, Wash<strong>in</strong>gton, USA, 457 pp.<br />
Mooney, H.A., Mack, R.N., McNeely, J.A., Neville, L.E., Schei, P.J. and J.K.<br />
Waage (eds) (2005) <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong>: a new synthesis. Island Press,<br />
Wash<strong>in</strong>gton, Covelo & London, 368 pp.<br />
Müller, J.C. and E.M. Griebeler (2002) Genetics on <strong>in</strong>vasive <strong>species</strong>. In:<br />
Leppäkoski, E., Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic <strong>species</strong><br />
of Europe. Distribution, impacts and management. Kluwer Academic<br />
Publishers, Dordrecht, pp. 173–182.<br />
Pimentel, D. (ed.) (2002) Biological <strong>in</strong>vasions; economics and<br />
environmental costs of <strong>alien</strong> plant, animal, and microbe <strong>species</strong>. CRC<br />
Press, Boca Raton, London, New York, Wash<strong>in</strong>gton, D.C., 369 pp.<br />
Pimentel, D., Lach, L., Zuniga, R. and D. Morrison (2000) Environmental<br />
and economic costs of non<strong>in</strong>digenous <strong>species</strong> <strong>in</strong> the United States.<br />
BioScience 50, 53–65.<br />
Sh<strong>in</strong>e, C., Williams, N. and L. Gündl<strong>in</strong>g (2000) A guide to design<strong>in</strong>g legal<br />
and <strong>in</strong>stitutional frameworks on <strong>alien</strong> <strong>in</strong>vasive <strong>species</strong>. IUCN, Gland,<br />
<strong>Switzerland</strong> and Cambridge and Bonn, 138 pp.<br />
S<strong>in</strong>den, J., Jones, R., Hester, S., Odom, D., Kalisch, C., James, R. and O.<br />
Cacho (2004) The economic impact of weeds <strong>in</strong> Australia. CRC for<br />
Australian Weed Management, Technical Series No. 8, 65 pp.<br />
Thiess, A., Schuster, R., Nöckler, K. and H. Mix (2001) Helm<strong>in</strong>thenfunde<br />
beim e<strong>in</strong>heimischen Marderhund (Nyctereutes procyonoides, Gray, 1834).<br />
Berl<strong>in</strong>er und Münchner Tierärztliche Wochenschrift 114, 273–276.<br />
US Congress (1993) Harmful non<strong>in</strong>digenous <strong>species</strong> <strong>in</strong> the United States.<br />
Office of Technology Assessment, OTA-F–565. U.S. Congress Government<br />
Pr<strong>in</strong>t<strong>in</strong>g Office, Wash<strong>in</strong>gton, D.C., USA<br />
Williamson, M. (1996) Biological <strong>in</strong>vasions. Chapman & Hall, London, 244<br />
pp.<br />
Wittenberg, R. and M.J.W. Cock (eds) (2001) <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong>: a<br />
toolkit of best prevention and management practices. CABI Publish<strong>in</strong>g,<br />
Wall<strong>in</strong>gford, UK, 228 pp.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 36<br />
2 > Vertebrates – Vertebrata<br />
Prepared by Rüdiger Wittenberg<br />
All the Fact Sheets for vertebrates have been placed at the end of this chapter. They are<br />
presented <strong>in</strong> alphabetical order to make <strong>in</strong>dividual ones easier to locate, not <strong>in</strong> taxonomical<br />
sequence as <strong>in</strong> the text and lists below.<br />
2.1 Mammals – Mammalia<br />
There are currently about eight mammalian neozoa with established populations <strong>in</strong><br />
<strong>Switzerland</strong> (Table 2.1), another <strong>species</strong> (coypu) is irregularly found, and the grey<br />
squirrel is discussed as a threat to <strong>Switzerland</strong>. Of the established <strong>species</strong>, one is greatly<br />
reduced <strong>in</strong> numbers (European rabbit) due to unfavourable climatic conditions,<br />
hunt<strong>in</strong>g, and <strong>in</strong>troduced diseases such as myxomatosis and rabbit haemorrhagic disease<br />
(RHD) (see below). Three have only small, localized populations (Siberian chipmunk,<br />
sika deer, mouflon), one is currently <strong>in</strong> the process of <strong>in</strong>vasion (raccoon dog), two are<br />
well-established and spread<strong>in</strong>g through <strong>Switzerland</strong> (raccoon, musk rat) and one is<br />
abundant (brown rat).<br />
The European rabbit (Oryctolagus cuniculus L.) was widely distributed throughout<br />
Central and Southern Europe before the last glacial period (Kaetzke et al., 2003).<br />
Dur<strong>in</strong>g this period the distribution retracted dramatically and the European rabbit<br />
survived only on the Iberian Pen<strong>in</strong>sula. Later it was distributed by humans, because it<br />
was prized for its meat, first along the Mediterranean Sea coast and then to Central<br />
Europe. However, it arrived relatively recently <strong>in</strong> <strong>Switzerland</strong>, where it was locally<br />
<strong>in</strong>troduced <strong>in</strong> the 19 th century (Long, 2003). The distribution <strong>in</strong> <strong>Switzerland</strong> was always<br />
very patchy <strong>in</strong> lowlands around Basel, Genève and <strong>in</strong> the Valais and Tic<strong>in</strong>o<br />
(Hausser, 1995) with a decreas<strong>in</strong>g trend <strong>in</strong> its population sizes. While the European<br />
rabbit was able to adapt very well to the hot and arid <strong>in</strong>terior of Australia, it suffers<br />
under cold w<strong>in</strong>ter temperatures and is restricted by the depth of the snow which covers<br />
its food (Flux, 1994). The small population of the European rabbit <strong>in</strong> <strong>Switzerland</strong> and<br />
the very few suitable habitats render this <strong>alien</strong> <strong>species</strong> unimportant. This is an <strong>in</strong>terest<strong>in</strong>g<br />
contrast to many other countries around the globe where the <strong>species</strong> is recognized<br />
for its detrimental effects on agriculture, forestry and the environment, <strong>in</strong>clud<strong>in</strong>g the<br />
more than 800 islands where it has been <strong>in</strong>troduced. It should be noted that releases of<br />
other <strong>alien</strong> members of the family Leporidae should be discouraged. A North American<br />
<strong>species</strong> <strong>in</strong>troduced for hunt<strong>in</strong>g at several locations <strong>in</strong> Europe, <strong>in</strong>clud<strong>in</strong>g <strong>Switzerland</strong>,<br />
France and Italy is Sylvilagus floridanus (J.A. Allan). The ma<strong>in</strong> concern is the possibility<br />
of transmission of diseases to native <strong>species</strong>, e.g. myxomatosis and RHD.
2 > Vertebrates – Vertebrata 37<br />
The only <strong>in</strong>troduced squirrel is the Siberian chipmunk (Tamias sibiricus (Laxmann)).<br />
It was released <strong>in</strong>to parks <strong>in</strong> Genève by pet lovers and established a small but stable<br />
population (Long, 2003). Its native distribution covers a large part of northern Asia and<br />
it is expand<strong>in</strong>g its range westwards, and arrived <strong>in</strong> F<strong>in</strong>land <strong>in</strong> recent years (Grzimek,<br />
1975). This small beautiful squirrel is frequently kept as a pet and has escaped or been<br />
deliberately <strong>in</strong>troduced <strong>in</strong> several European countries. In <strong>Switzerland</strong> only one small<br />
colony is known, but it has also established populations <strong>in</strong> Freiburg im Breisgau <strong>in</strong><br />
Germany and <strong>in</strong> Italy (Personal observation; Andreotti et al., 2001). In Russia it is<br />
known to destroy half the average forest nut production and cause great damage to<br />
gra<strong>in</strong> crops. S<strong>in</strong>ce the chipmunks ma<strong>in</strong>ly feed on the ground, they can severely affect<br />
crops and also damage gardens and orchards (Long, 2003). A Belgian study, which<br />
compared the abundance of birds <strong>in</strong> areas with and without the <strong>alien</strong> squirrel, concluded<br />
that T. sibiricus has no impact on bird populations (Riegel, 2001). In most of<br />
Europe, released populations have not <strong>in</strong>creased and spread significantly, and it seems<br />
that the Siberian chipmunk is dependent on a rich food source, as it is most often found<br />
<strong>in</strong> parks and graveyards with a variety of different plant <strong>species</strong> (Krapp, 1978a, b).<br />
Thus, it should be regarded as a <strong>species</strong> with a low <strong>in</strong>vasion potential. However, the<br />
release of pet animals should be of great concern and should probably be better regulated.<br />
In particular, the danger of diseases carried by <strong>alien</strong> pets to wildlife, other pets<br />
and humans should not be underrated, as demonstrated by the recent arrival of diseases<br />
with <strong>in</strong>troduced pet rats from Africa to the USA.<br />
Another squirrel <strong>species</strong>, although one that has not yet arrived <strong>in</strong> <strong>Switzerland</strong>, is of<br />
great concern, as it is established <strong>in</strong> Italy and spread<strong>in</strong>g. The grey squirrel (Sciurus<br />
carol<strong>in</strong>ensis Gmel<strong>in</strong>) (see Fact Sheet) is one of the rare cases where the impact on a<br />
native congeneric <strong>species</strong>, i.e. the red squirrel (Sciurus vulgaris L.), is well documented.<br />
In Great Brita<strong>in</strong>, except for some mounta<strong>in</strong>ous conifer forests, the red squirrel<br />
has been replaced completely by the grey squirrel. Today, the population <strong>in</strong> northern<br />
Italy is rapidly expand<strong>in</strong>g its range and threatens to <strong>in</strong>vade <strong>Switzerland</strong> <strong>in</strong> the next 20<br />
years or so. Swiss authorities should follow the expansion closely and put prevention<br />
methods <strong>in</strong> place to stop or delay the spread at the border. Whereas the <strong>in</strong>vasion <strong>in</strong>to<br />
the Tic<strong>in</strong>o is more imm<strong>in</strong>ent, the modelled expansion of the grey squirrel to France is<br />
of greater concern, as from here it can expand <strong>in</strong>to <strong>Switzerland</strong> around Genève. Moreover,<br />
if isolated populations outside the <strong>in</strong>vasion front are encountered, they should be<br />
eradicated as long as they are small. Swiss authorities should also <strong>in</strong>itiate discussion<br />
with Italian authorities to implement the approved action plan to eradicate the <strong>species</strong><br />
<strong>in</strong> the Italian part of the Tic<strong>in</strong>o valley.<br />
The re-<strong>in</strong>troduction of beavers <strong>in</strong> some countries has been problematic <strong>in</strong> the past. In<br />
some areas, at least <strong>in</strong> Austria and F<strong>in</strong>land and perhaps Germany (Geiter et al., 2002),<br />
the American congeneric Castor canadensis Kuhl was <strong>in</strong>troduced <strong>in</strong>stead of the European<br />
beaver (C. fiber L.) (Freye, 1978; Englisch, 2002). These two <strong>species</strong> are very<br />
similar and <strong>in</strong>deed were considered to be one <strong>species</strong> until recently, but the karyotypes<br />
differ. No further <strong>in</strong>troductions of the American <strong>species</strong> should be made <strong>in</strong> Europe. The<br />
<strong>in</strong>troductions of C. fiber of different orig<strong>in</strong>s with<strong>in</strong> Europe seem to be of less concern<br />
<strong>in</strong> many parts, s<strong>in</strong>ce the <strong>species</strong> is genetically relatively uniform, with only a few
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 38<br />
obvious sub<strong>species</strong>. In general, specimens used for re-<strong>in</strong>troductions should come from<br />
the same sub<strong>species</strong> or populations, orig<strong>in</strong>al to the specific area, whenever possible.<br />
The North American muskrat (Ondatra zibethicus (L.)) (see Fact Sheet) showed a<br />
tremendous expansion of its range <strong>in</strong> Europe <strong>in</strong> the 100 years after its first <strong>in</strong>troduction<br />
to Prague (Elton, 1958). A discussion of the process of the spread is given by Williamson<br />
(1996). The <strong>in</strong>vasion front is mov<strong>in</strong>g swiftly southwards <strong>in</strong> <strong>Switzerland</strong>. It has the<br />
potential to colonize all aquatic bodies <strong>in</strong> the lowland parts of <strong>Switzerland</strong>. The damage<br />
to waterways and the costs to control this <strong>species</strong> are enormous (Re<strong>in</strong>hardt et al.,<br />
2003). However, given the severity of the damage and the negative impact on native<br />
mussels (Unionidae) through predation, an eradication or control programme should be<br />
considered. Options should be explored together with neighbour<strong>in</strong>g countries, e.g.<br />
Germany. Eradication is probably not achievable given the large population already<br />
present <strong>in</strong> <strong>Switzerland</strong> and the existence of sources for new <strong>in</strong>vasions <strong>in</strong> Germany and<br />
France. On the British Isle the muskrat was eradicated <strong>in</strong> 1939, but the population was<br />
still small and conf<strong>in</strong>ed to several centres of <strong>in</strong>troduction <strong>in</strong> England, Scotland and<br />
Ireland. The British Isles are not connected to cont<strong>in</strong>ental Europe, thus it is easier to<br />
prevent re-<strong>in</strong>festation. It has also been suggested that the UK climate is not ideal for<br />
the <strong>species</strong>, so that eradication was more easily achieved.<br />
Another large rodent, the coypu (Myocastor coypus (Mol<strong>in</strong>a)) has been found <strong>in</strong><br />
<strong>Switzerland</strong> on several occasions and <strong>in</strong> different areas, ma<strong>in</strong>ly <strong>in</strong> the north-west<br />
(Hausser, 1995). Its orig<strong>in</strong> is the southern half of South America and it is conf<strong>in</strong>ed to<br />
water edges. Like the muskrat, it was deliberately <strong>in</strong>troduced <strong>in</strong>to the wild for hunt<strong>in</strong>g<br />
because of its fur; like most aquatic animals, it has very dense fur to help it conserve<br />
heat <strong>in</strong> cold water. However, most coypu colonies are derived from animals that escaped<br />
from fur farms and established short-lived populations <strong>in</strong> Central Europe. They<br />
are not well-adapted to cold w<strong>in</strong>ters, thus severe w<strong>in</strong>ters often cause high mortality and<br />
local ext<strong>in</strong>ction. In the Mediterranean area (e.g. southern France and northern Italy)<br />
coypu populations thrive and they damage dams by burrow<strong>in</strong>g and reduce crop yields<br />
(Southern, 1964). It can be concluded that the coypu is unlikely to establish permanent<br />
populations and cause an environmental impact <strong>in</strong> <strong>Switzerland</strong>, because of the prolonged<br />
frost periods. Most of the sight<strong>in</strong>gs will be of s<strong>in</strong>gle specimens or temporary<br />
populations. However, it needs to be stressed that it is difficult to be confident about<br />
this prediction. The coypu was successfully eradicated from England, and casualties<br />
amongst non-target organisms, which had been considerable dur<strong>in</strong>g the muskrat eradication,<br />
were reduced by us<strong>in</strong>g cage traps and releas<strong>in</strong>g non-target <strong>species</strong> back <strong>in</strong>to the<br />
wild (Williamson, 1996).<br />
The ubiquitous brown rat (Rattus norvegicus (Berkenhout)) (see Fact Sheet) is a<br />
<strong>species</strong> that adapted very well to human environments. The orig<strong>in</strong> of the genus Rattus<br />
is the warm climate of South-east Asia, but R. norvegicus is from the northern part of<br />
the distribution, and is thus better adapted to a cold climate. However, it is surpris<strong>in</strong>g<br />
how well this <strong>species</strong> spread with humans all around the globe. Its omnivorous character,<br />
together with its ability to live <strong>in</strong> close vic<strong>in</strong>ity to humans, led to its success. The<br />
recorded impacts from this rodent are considerable. Its negative impact on the environment<br />
is ma<strong>in</strong>ly evident on islands, where it has even caused bird ext<strong>in</strong>ctions. In
2 > Vertebrates – Vertebrata 39<br />
urban areas, the normal habitat <strong>in</strong> Central Europe, the environmental impact is probably<br />
less pronounced, although it is an additional omnivorous predator. However, its<br />
impact on structures and foodstuffs and its role as a disease vector <strong>in</strong> human settlements<br />
are unambiguous and the brown rat is controlled <strong>in</strong> many cities.<br />
The raccoon (Procyon lotor (L.)) (see Fact Sheet) is the only member of the family<br />
Procyonidae <strong>in</strong>troduced to Europe. It was <strong>in</strong>troduced from North America and released<br />
<strong>in</strong> Germany <strong>in</strong> 1934. From that and other subsequent releases, it spread through Germany<br />
and reached <strong>Switzerland</strong> <strong>in</strong> 1976. The spread cont<strong>in</strong>ues and it will eventually<br />
<strong>in</strong>habit the lowland parts of <strong>Switzerland</strong> to Genève <strong>in</strong> the south and the Valais. The<br />
impact of the <strong>species</strong> is disputed. A def<strong>in</strong>itive conclusion cannot be reached without<br />
studies be<strong>in</strong>g carried out <strong>in</strong> <strong>Switzerland</strong> to assess its impact on the native wildlife and<br />
as a vector of diseases (Hohmann et al., 2002). This addition to the native fauna may<br />
have to be accepted, s<strong>in</strong>ce control measures are not very effective and its impacts have<br />
not (yet) been demontrated. If <strong>in</strong>cidences of diseases carried by the raccoon <strong>in</strong>crease,<br />
hunters will have to focus more on this <strong>species</strong>.<br />
The only <strong>in</strong>troduced dog (Canidae) <strong>in</strong> Central Europe is the east Asian raccoon dog<br />
(Nyctereutes procyonoides (Gray)) (see Fact Sheet), which was for decades and at<br />
many places <strong>in</strong> the former USSR extensively released as a fur animal. The acclimatization<br />
of the <strong>species</strong> and the value of the fur are questionable. However, it spread westwards<br />
and recently reached France. Incorrect identification based on field observations<br />
by laypeople may be frequent, because they are not familiar with this <strong>species</strong> and it can<br />
be confused with other predators, such as the raccoon. Thus, identification of dead<br />
animals is the most reliable way of evaluat<strong>in</strong>g its spread through Europe. Op<strong>in</strong>ions<br />
about its impact vary between it be<strong>in</strong>g a benign <strong>species</strong> and a <strong>species</strong> caus<strong>in</strong>g severe<br />
impact on native wildlife. One argument is that, as a generalist on abundant food<br />
resources, it does no harm to rare organisms. Others argue that predation on bird<br />
<strong>species</strong> and amphibians might have a negative impact (Kauhala, 1996). The status of<br />
many amphibians as endangered <strong>in</strong> Central Europe and the fact that amphibians were<br />
found <strong>in</strong> proportions of up to 45 % <strong>in</strong> the diet of the raccoon dog (Barbu, 1972) <strong>in</strong>dicates<br />
the potential harm. It also plays a role as a vector of diseases to humans and other<br />
predators (Weidema, 2000). It is recommended that the spread of the raccoon dog<br />
should be monitored and its impact <strong>in</strong> <strong>Switzerland</strong> evaluated. However, a control<br />
programme seems unlikely to achieve success, because of the raccoon dog’s secretive<br />
behaviour and its cont<strong>in</strong>u<strong>in</strong>g <strong>in</strong>vasion from the East. The raccoon dog and the raccoon<br />
are two additional predators with which native animals have to cope. Thus, although<br />
the impacts of the <strong>in</strong>troduced predatory <strong>species</strong> are not so different from those of native<br />
orig<strong>in</strong>, if they add to the total predator load, they are likely <strong>in</strong>crease the predatory<br />
pressure, and destabilize exist<strong>in</strong>g populations and population cycles.<br />
The only established <strong>alien</strong> deer (Cervidae) <strong>in</strong> <strong>Switzerland</strong> is the sika deer (Cervus<br />
nippon Temm<strong>in</strong>ck). It is a fairly small deer <strong>species</strong>, <strong>in</strong>termediate <strong>in</strong> size between the<br />
native roe deer (Capreolus capreolus (L.)) and red deer (Cervus elaphus L.). The<br />
summer coat is chestnut brown to buff brown with white spots, while it is uniform<br />
grey-brown dur<strong>in</strong>g w<strong>in</strong>ter. The rump patch is white with a black upper border and the<br />
white tail has a black stripe. The typical antlers have four po<strong>in</strong>ts each. The sika deer is
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 40<br />
mostly nocturnal, although this might be an adaptation to disturbance and hunt<strong>in</strong>g,<br />
s<strong>in</strong>ce most deer <strong>species</strong>, if undisturbed, would probably be diurnal. It was <strong>in</strong>troduced<br />
from its orig<strong>in</strong>al range <strong>in</strong> eastern Asia to several countries <strong>in</strong> Europe, as well as North<br />
America, Africa, Madagascar, Australia and New Zealand (Long, 2003), as a game<br />
animal, for meat, and as a conservation measure. The sika deer <strong>in</strong> its natural distribution<br />
range, like most (Asiatic) deer <strong>species</strong>, is under threat due to high hunt<strong>in</strong>g pressure<br />
and many sub<strong>species</strong> are rare (Kurt, 1988). Many sika deer <strong>in</strong>troductions were successful,<br />
<strong>in</strong>clud<strong>in</strong>g one <strong>in</strong> southern Germany, which was the source of the Swiss population.<br />
The enclosure near Schaffhausen was opened <strong>in</strong> 1941 and a slowly <strong>in</strong>creas<strong>in</strong>g population<br />
established (Hausser, 1995). This population is still restricted to the north of the<br />
river Rh<strong>in</strong>e. The sika deer is a serious forest pest, brows<strong>in</strong>g young trees and also<br />
remov<strong>in</strong>g bark (Welch et al., 2001). Native deer show the same behaviour and their<br />
large populations have created an unresolved problem <strong>in</strong> many places. However, an<br />
additional, <strong>alien</strong>, deer <strong>species</strong> can <strong>in</strong>tensify tree damage (Eisfeld and Fischer, 1996). It<br />
can damage crops, as do other ungulates, and can be a road hazard. A serious threat to<br />
the native red deer could potentially come from hybridization with the <strong>alien</strong> sika deer<br />
(Krapp und Niethammer, 1986; Welch et al., 2001). However, hybridization has not yet<br />
been demonstrated <strong>in</strong> Germany or Austria (Geiter et al., 2002; Welch et al., 2001).<br />
Thus, there is an urgent need to monitor the Swiss population of sika deer and study<br />
whether hybridization with the red deer occurs. The outcome of the study will be<br />
important for a decision on how to manage the sika deer population – eradication,<br />
conta<strong>in</strong>ment or ‘harvest’ hunt<strong>in</strong>g. The other fact to be taken <strong>in</strong>to consideration is the<br />
status of the endangered populations of sika deer <strong>in</strong> its native range. A population <strong>in</strong><br />
<strong>Switzerland</strong> could be valuable for the conservation of the <strong>species</strong>. A problem with<br />
many European populations is the lack of knowledge concern<strong>in</strong>g which sub<strong>species</strong><br />
were <strong>in</strong>troduced, and many are of mixed orig<strong>in</strong>s. Thus, given that sika deer is common<br />
<strong>in</strong> other <strong>in</strong>troduced areas, the <strong>species</strong> should preferably be eradicated <strong>in</strong> <strong>Switzerland</strong> or<br />
at least conta<strong>in</strong>ed <strong>in</strong> the <strong>in</strong>fested area.<br />
The mouflon (Ovis orientalis ssp. musimon Gmel<strong>in</strong> or O. ammon ssp. musimon (Pallas))<br />
has established a small population over the last 20 years or so <strong>in</strong> the Valais, where<br />
they are slowly <strong>in</strong>creas<strong>in</strong>g <strong>in</strong> numbers. The mouflon is a member of a <strong>species</strong>-complex<br />
of wild sheep. The views on <strong>species</strong> status <strong>in</strong> that group vary considerably between<br />
authors. The mouflon was <strong>in</strong>troduced to Central Europe from Corsica and Sard<strong>in</strong>ia.<br />
However, the status of these populations is not clear. While some argue that the mouflon<br />
is native to these islands, Popl<strong>in</strong> (1979) and others provide evidence that the<br />
populations on Corsica and Sard<strong>in</strong>ia are descendants from domestic animals that were<br />
<strong>in</strong>troduced from Asia <strong>in</strong> the Neolithic Age. The animals were still very similar to their<br />
wild ancestors when brought to the Mediterranean islands. In about 1980, the mouflon<br />
entered the Valais from a population <strong>in</strong> France and established populations <strong>in</strong> the lower<br />
Valais. In some localities the mouflon competes with the chamois (Rupicapra rupicapra<br />
(L.)), but <strong>in</strong> general the environmental impact will not be significant (Andreotti<br />
et al., 2001). As long as the population <strong>in</strong> <strong>Switzerland</strong> is small, no significant economic<br />
damage will occur. A decision on the future of the mouflon <strong>in</strong> <strong>Switzerland</strong> needs to be<br />
taken before the populations spread any further. Exclud<strong>in</strong>g the game hunt<strong>in</strong>g aspect, it<br />
is an <strong>alien</strong> <strong>species</strong> with no value, and today the value of native <strong>species</strong> is a predom<strong>in</strong>ant<br />
consideration <strong>in</strong> nature conservation, therefore control of the <strong>species</strong> could be consid-
2 > Vertebrates – Vertebrata 41<br />
ered. It is recommended that the <strong>species</strong> at least be conta<strong>in</strong>ed to the present <strong>in</strong>fested<br />
area, us<strong>in</strong>g hunt<strong>in</strong>g as a measure of control.<br />
A list of <strong>species</strong> present<strong>in</strong>g a potential threat to native biodiversity is not complete<br />
without mention<strong>in</strong>g domestic animals. Cats (Felis catus L.), ferrets (Mustela furo L.),<br />
dogs (Canis familiaris L.) and goats (Capra hircus L.) are examples of domestic<br />
animals for which an environmental impact is documented (Long, 2003). As with most<br />
IAS, the impacts of the above-mentioned domestic animals are greatest on islands,<br />
however there are demonstrated impacts <strong>in</strong> Europe. Domestic cats are very efficient<br />
hunters and will kill wild prey, despite be<strong>in</strong>g fed. Based on a survey, the total number<br />
of mammals, birds, amphibians and reptiles killed by domestic cats <strong>in</strong> England per year<br />
has been estimated to exceed 89 million, <strong>in</strong>clud<strong>in</strong>g 25 million birds (Woods et al.,<br />
2003). Feral cats, which depend entirely on wild food resources, add to this figure.<br />
Domestic populations can occur at high densities, because they are fed and belong to<br />
households. Urban and rural habitats, where the domestic cats ma<strong>in</strong>ly hunt, do not have<br />
many endangered <strong>species</strong>, but are a valuable habitat <strong>in</strong> highly populated Central Europe.<br />
Thus, the effect on native biodiversity is significant. However, a management plan<br />
for domestic cats, such as by-laws to keep cats <strong>in</strong>doors near nature reserves, is a thorny<br />
topic. The impact of hybridization with the wild cat (Felis silvestris Schreber) is not<br />
clear, but it seems to be a conservation issue <strong>in</strong> some areas (Randi et al., 2001).<br />
The feral ferret is a domesticated form of the eastern polecat (Mustela putorius L.) and<br />
escaped animals are widely found. Potentially there could be damage to native small<br />
mammal and ground-nest<strong>in</strong>g bird populations and from <strong>in</strong>trogression of genes <strong>in</strong>to the<br />
wild form, but there is very little <strong>in</strong>formation about feral ferrets <strong>in</strong> the wild (e.g.<br />
whether there are self-susta<strong>in</strong><strong>in</strong>g populations). Feral dogs are more of a problem for<br />
game animals than to biodiversity <strong>in</strong> Central Europe. Problems with feral goats <strong>in</strong><br />
Central Europe, <strong>in</strong> contrast to island ecosystems on a global scale, are rare and local<br />
and limited to hybridization with the native ibex (Capra ibex (L.)).<br />
About half of the <strong>alien</strong> mammal <strong>species</strong> occurr<strong>in</strong>g <strong>in</strong> (or threaten<strong>in</strong>g to <strong>in</strong>vade) <strong>Switzerland</strong><br />
(i.e. muskrat, coypu, grey squirrel, sika deer, raccoon and raccoon dog) are<br />
given as examples of <strong>species</strong> threaten<strong>in</strong>g European wildlife and habitats <strong>in</strong> the Appendix<br />
of Recommendation No. 77 (1999) on eradication of non-native terrestrial vertebrates<br />
of the Bern Convention on the Conservation of European Wildlife and Natural<br />
Habitats (for details on the recommendation for management, see<br />
http://www.nature.coe.<strong>in</strong>t/english/ma<strong>in</strong>/Bern/texts/rec9977.htm). This recommendation<br />
was adopted by the Stand<strong>in</strong>g Committee and specifies the recommended action to be<br />
taken aga<strong>in</strong>st IAS, <strong>in</strong>clud<strong>in</strong>g monitor<strong>in</strong>g, control of sale, and efficacy of eradication<br />
efforts. The recommendation sets another framework for deal<strong>in</strong>g with IAS <strong>in</strong> Europe.<br />
Despite the low number of mammalian neozoa, some patterns can be observed and are<br />
discussed below. The ten <strong>species</strong> listed above are members of n<strong>in</strong>e families. Two of<br />
these families have no native members <strong>in</strong> Europe: Capromyidae occur only <strong>in</strong> the New<br />
World, and Procyonidae <strong>in</strong> either the New World and east Asia or exclusively <strong>in</strong> the<br />
New World, depend<strong>in</strong>g on whether the red panda, Ailurus fulgens (Cuvier), is <strong>in</strong>cluded<br />
or not. The different phylogenetic orig<strong>in</strong> of the ten <strong>species</strong> is reflected <strong>in</strong> the very
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 42<br />
different biologies of the <strong>species</strong>, which <strong>in</strong>clude small rodents, large herbivores and<br />
carnivores.<br />
Five <strong>species</strong> are of Asian orig<strong>in</strong> (four from eastern Asia and one from western Asia),<br />
four from the New World (one of those from South America), and one is Mediterranean.<br />
The pathways of their <strong>in</strong>troductions reveal a clear pattern, which is generally true for<br />
<strong>in</strong>troduced mammals: they were deliberately <strong>in</strong>troduced <strong>in</strong>to Europe. Only one <strong>species</strong><br />
arrived as a stowaway animal <strong>in</strong> cargo, ships and other vehicles – the ubiquitous brown<br />
rat. The majority of the <strong>species</strong> are fur animals which were released and escaped from<br />
farms (four <strong>species</strong>); the two large ungulates were released to enrich the hunt<strong>in</strong>g fauna,<br />
one was used as a food resource and the other to enrich the fauna and for aesthetic<br />
reasons. One other <strong>in</strong>terest<strong>in</strong>g aspect is how the <strong>species</strong> arrived <strong>in</strong> <strong>Switzerland</strong>. The<br />
pathway analysis presented above shows how the <strong>species</strong> were transported <strong>in</strong>to Europe.<br />
Only four of the ten <strong>species</strong> <strong>in</strong> Table 2.1 were actually <strong>in</strong>troduced directly <strong>in</strong>to <strong>Switzerland</strong>.<br />
The most (potentially) damag<strong>in</strong>g <strong>species</strong> have spread <strong>in</strong>to <strong>Switzerland</strong> from an<br />
<strong>in</strong>troduction elsewhere <strong>in</strong> Europe, ma<strong>in</strong>ly Germany and France. This highlights the<br />
importance of <strong>in</strong>ternational dialogue and collaboration on IAS issues.<br />
Some of the most destructive <strong>alien</strong> <strong>species</strong> on a global scale are mammals (Long,<br />
2003). Although the number of <strong>in</strong>troduced <strong>species</strong> is fairly small <strong>in</strong> comparison to<br />
other groups, the impact is considered to be enormous, especially <strong>in</strong> island ecosystems.<br />
The large herbivores cause a major impact on plants because of their size and the<br />
amount they eat, while small carnivores (e.g. Mustelidae) and omnivorous mammals<br />
(e.g. Muridae) are <strong>in</strong>telligent and adaptable <strong>species</strong>, which are able to prey on a varied<br />
diet and can thrive <strong>in</strong> close proximity to humans. The latter trait facilitates their wide<br />
spread by human-mediated transport. The potential impacts of mammalian neozoa <strong>in</strong><br />
<strong>Switzerland</strong> are discussed <strong>in</strong> the <strong>species</strong> descriptions and the Fact Sheets. Fact Sheets<br />
are provided for the five <strong>species</strong> which are likely to cause some impact. There is no<br />
obvious general pattern, besides the fact that the small mammals have had the highest<br />
impact through predation on native <strong>species</strong> and damage to human constructions, <strong>in</strong>clud<strong>in</strong>g<br />
waterways. The date of <strong>in</strong>troduction and perceived impact show no correlation,<br />
s<strong>in</strong>ce the potential for impact rather than the actual impact <strong>in</strong> <strong>Switzerland</strong> is used.<br />
Thus, the grey squirrel, which has not yet been found <strong>in</strong> <strong>Switzerland</strong>, is one of the<br />
<strong>species</strong> of concern. On the other hand, the two <strong>species</strong> <strong>in</strong>troduced <strong>in</strong> the 19 th century<br />
comprise one highly damag<strong>in</strong>g <strong>species</strong> and one which is now almost ext<strong>in</strong>ct.<br />
The small number of <strong>in</strong>troduced <strong>species</strong> means that general patterns observed <strong>in</strong> larger<br />
data pools are unlikely to be replicated, due to chance.<br />
There are about 86 mammal <strong>species</strong> currently reproduc<strong>in</strong>g <strong>in</strong> <strong>Switzerland</strong>, of which<br />
n<strong>in</strong>e are neozoa (coypu <strong>in</strong>cluded), thus about 10 % are <strong>in</strong>troduced. When the list of <strong>species</strong><br />
is compared to those of the neighbour<strong>in</strong>g countries Austria and Germany, there is a<br />
high overlap <strong>in</strong> <strong>species</strong> composition (Englisch, 2002; Geiter et al., 2002). The total<br />
number is a little smaller (Austria: 11; Germany: 11); the most obvious difference is<br />
the absence of fallow deer (Cervus dama (L.)) and m<strong>in</strong>k (Mustela vison Schreber), both
2 > Vertebrates – Vertebrata 43<br />
established <strong>in</strong> the other two countries. Gibb and Flux (1973) list twice as many <strong>in</strong>troduced<br />
mammals to New Zealand, i.e. 25, and the damage they cause is enormous, s<strong>in</strong>ce<br />
the islands had only two native mammal <strong>species</strong> (both bats). The niches occupied elsewhere<br />
by terrestrial mammals were occupied <strong>in</strong> New Zealand by native bird <strong>species</strong>.<br />
The management options for the <strong>species</strong> are discussed <strong>in</strong> the text on <strong>in</strong>dividual <strong>species</strong>,<br />
above. The <strong>in</strong>vasive or potentially <strong>in</strong>vasive <strong>species</strong> are very difficult targets for eradication<br />
and control. The same traits that make them successful <strong>in</strong> the <strong>in</strong>vasion process<br />
renders them difficult to control, e.g. adaptability and a high fecundity.<br />
Tab. 2.1 > Established <strong>alien</strong> mammals <strong>in</strong> <strong>Switzerland</strong>.<br />
Scientific name<br />
Oryctolagus cuniculus<br />
(L., 1758)<br />
Tamias sibiricus<br />
(Laxmann, 1769)<br />
Sciurus carol<strong>in</strong>ensis<br />
Gmel<strong>in</strong>, 1788<br />
Ondatra zibethicus<br />
(L., 1766)<br />
Myocastor coypus<br />
(Mol<strong>in</strong>a, 1782)<br />
Rattus norvegicus<br />
(Berkenhout, 1769)<br />
Procyon lotor<br />
(L. 1758)<br />
Nyctereutes procyonoides<br />
Gray, 1834<br />
Cervus nippon<br />
Temm<strong>in</strong>ck, 1836<br />
Ovis orientalis<br />
musimon Gmel<strong>in</strong>,<br />
1774<br />
Family Orig<strong>in</strong> Year Pathway Impact Note<br />
Leporidae Iberian Pen<strong>in</strong>sula 19th century<br />
Sciuridae Asian 1975? Escaped and released to<br />
establish population<br />
from pet lovers<br />
Sciuridae Eastern North<br />
America<br />
Released for food Alteration of plant succession<br />
Agricultural pest<br />
- Released for aesthetic<br />
reasons<br />
Arvicolidae North America 1935 Escaped from fur farms<br />
and released to provide<br />
wild fur harvest<br />
Capromyidae South America - Escaped from fur farms<br />
and released to provide<br />
wild fur harvest<br />
Muridae Southeast Russia<br />
and northern Ch<strong>in</strong>a<br />
Procyonidae North and Central<br />
America<br />
19 th<br />
century<br />
Transported <strong>in</strong>advertently<br />
<strong>in</strong> ships and other<br />
vehicles<br />
1976 Releases as a fur animal<br />
and to enrich the fauna<br />
Escapees from captivity<br />
Canidae East Asia 2003 Acclimatization as a fur<br />
animal<br />
Cervidae South-eastern<br />
Russia, eastern<br />
Ch<strong>in</strong>a, Japan and<br />
Korea<br />
1941 Released for sport<br />
hunt<strong>in</strong>g and for conservation<br />
of the <strong>species</strong><br />
Bovidae Western Asia 1985? Released as a game<br />
animal<br />
Great damage to gra<strong>in</strong> crops and forest<br />
nut production <strong>in</strong> its native area<br />
Replaces the native red squirrel<br />
Stripp<strong>in</strong>g bark of trees can cause<br />
damage <strong>in</strong> plantations<br />
Dramatic economic costs due to<br />
damage to waterways<br />
Predation on native mussel populations<br />
Damage to crops by feed<strong>in</strong>g and water<br />
banks by burrow<strong>in</strong>g<br />
Transmission of human diseases<br />
High control costs<br />
Damage to crops and structures<br />
Predator of <strong>in</strong>vertebrates and vertebrates,<br />
with a possible impact through<br />
bird nest predation<br />
Nuisance <strong>in</strong> urban areas<br />
Problem <strong>in</strong> orchards?<br />
Vector of diseases<br />
Predator of vertebrates<br />
Vector of diseases<br />
Serious forest pest<br />
Hybridization with red deer (everywhere?)<br />
The <strong>species</strong> does not<br />
thrive <strong>in</strong> <strong>Switzerland</strong><br />
due to the cold w<strong>in</strong>ters<br />
Only small local<br />
population <strong>in</strong> parks <strong>in</strong><br />
Genève<br />
This <strong>species</strong> is not yet<br />
found <strong>in</strong> <strong>Switzerland</strong>,<br />
but is rapidly spread<strong>in</strong>g<br />
<strong>in</strong> Italy<br />
Effective control<br />
options should be<br />
considered<br />
A permanent establishment<br />
seems<br />
unlikely because of<br />
harsh w<strong>in</strong>ters<br />
Generally, urban<br />
populations are<br />
controlled<br />
It will spread through<br />
the entire Mittelland<br />
with<strong>in</strong> the next few<br />
years/decades<br />
It is only currently<br />
spread<strong>in</strong>g <strong>in</strong>to<br />
<strong>Switzerland</strong> from<br />
Germany and France<br />
Only one restricted<br />
population <strong>in</strong> <strong>Switzerland</strong><br />
Local competition with chamois Only a small population<br />
<strong>in</strong> the lower Valais
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 44<br />
2.2 Birds – Aves<br />
About a quarter of the 510 bird <strong>species</strong> recorded for <strong>Switzerland</strong> are <strong>alien</strong>. This is not<br />
surpris<strong>in</strong>g, s<strong>in</strong>ce most birds are good flyers and many migrate regularly. The latter can<br />
easily go astray, especially dur<strong>in</strong>g freak weather events. Hurricane ‘Lothar’, for example,<br />
was responsible for the accidental occurrence of a pelagic bird (generally found far<br />
out at sea), the European storm-petrel (Hydrobates pelagicus (L.)), <strong>in</strong> land-locked<br />
<strong>Switzerland</strong> (Keller and Zb<strong>in</strong>den, 2001). About 200 <strong>species</strong> breed regularly <strong>in</strong> <strong>Switzerland</strong>.<br />
Some bird <strong>species</strong> are spread<strong>in</strong>g naturally through Europe and <strong>in</strong>to <strong>Switzerland</strong>. They<br />
are not <strong>in</strong>cluded <strong>in</strong> the list, although their spread might be triggered by human alterations<br />
to the landscape. The spread of these <strong>species</strong> is <strong>in</strong>directly facilitated by human<br />
action, but they are not <strong>alien</strong> as def<strong>in</strong>ed here. The collared dove (Streptopelia decaocto<br />
(Frivaldszky)) has spread phenomenally from the Balkans north-west through most of<br />
Europe (Glutz von Blotzheim, 1980). Several hypotheses have been put forward to<br />
account for this spread, <strong>in</strong>clud<strong>in</strong>g an <strong>in</strong>creased amount of food <strong>in</strong> rural areas and<br />
genetic changes <strong>in</strong> the bird population, but there is no agreement on the explanation.<br />
The <strong>species</strong> considered <strong>in</strong> this report are exclusively those which were released or<br />
escaped and became established, but not those that expanded their range to <strong>in</strong>clude<br />
<strong>Switzerland</strong>. However, <strong>species</strong> escap<strong>in</strong>g from captivity and observed <strong>in</strong> the wild or<br />
found to breed irregularly are not <strong>in</strong>cluded. Thus, the list provided here takes <strong>in</strong>to<br />
account only birds with established populations <strong>in</strong> <strong>Switzerland</strong> or <strong>in</strong> neighbour<strong>in</strong>g<br />
countries, from which spread <strong>in</strong>to <strong>Switzerland</strong> can be reasonably expected.<br />
Firstly, bird <strong>species</strong> with established populations <strong>in</strong> <strong>Switzerland</strong> (six <strong>species</strong>) are<br />
discussed, and secondly, <strong>alien</strong> <strong>species</strong> established <strong>in</strong> nearby countries which have the<br />
potential to spread to <strong>Switzerland</strong> are considered.<br />
The cormorant (Phalacrocorax carbo L.) is a common guest dur<strong>in</strong>g w<strong>in</strong>ter <strong>in</strong> <strong>Switzerland</strong>,<br />
but the first breed<strong>in</strong>g pairs are now established along the Aare near Bern. The<br />
birds are probably from the population kept under semi-wild conditions <strong>in</strong> the zoo.<br />
Follow<strong>in</strong>g the bann<strong>in</strong>g of hunt<strong>in</strong>g <strong>in</strong> numerous countries, the cormorant is spread<strong>in</strong>g<br />
aga<strong>in</strong> <strong>in</strong> Europe and <strong>in</strong> many places the conflict between conservation and fish<strong>in</strong>g<br />
<strong>in</strong>terests is fervidly debated. Thus, <strong>in</strong> this specific case and <strong>in</strong> general, <strong>species</strong> should<br />
be held <strong>in</strong> secure captivity. Bear<strong>in</strong>g <strong>in</strong> m<strong>in</strong>d recently acquired knowledge about <strong>in</strong>vasive<br />
bird <strong>species</strong> and the ease with which some are able to establish, keep<strong>in</strong>g populations<br />
under semi-wild conditions <strong>in</strong> zoos and similar facilities is no longer always<br />
appropriate. In the recent past, zoos have <strong>in</strong>creased this k<strong>in</strong>d of open conf<strong>in</strong>ement to<br />
simulate a more natural situation for the benefit of visitors. Unfortunately, as it creates<br />
a new pathway for <strong>in</strong>troductions, this approach has to be changed. A similar case is the<br />
cattle egret (Bubulcus ibis (L.)), which is frequently kept as free-fly<strong>in</strong>g populations <strong>in</strong><br />
zoos, and <strong>in</strong>dividuals escape occasionally from captivity. Birds of exotic orig<strong>in</strong> kept<br />
under semi-natural conditions can cause another problem: <strong>in</strong>trogression of genes <strong>in</strong>to<br />
native populations.
2 > Vertebrates – Vertebrata 45<br />
The mute swan (Cygnus olor (Gmel<strong>in</strong>)) (see Fact Sheet) was released on park ponds <strong>in</strong><br />
the 17 th century and has subsequently spread to all suitable habitats. It has a stable<br />
population, so only local problems occur, and it is also a popular <strong>species</strong> with the<br />
human population, so that control should be limited to public education concern<strong>in</strong>g the<br />
impact of feed<strong>in</strong>g waterfowl (Schmid et al., 1998). Stopp<strong>in</strong>g the feed<strong>in</strong>g of the swans<br />
will elim<strong>in</strong>ate the high densities of mute swans. The mute swans would then distribute<br />
themselves more evenly, due to their territoriality based on <strong>in</strong>traspecific aggression.<br />
A second member of the Anatidae, the greylag goose (Anser anser (L.)), the wild form<br />
of the domestic goose, is not native <strong>in</strong> <strong>Switzerland</strong>, but has a grow<strong>in</strong>g breed<strong>in</strong>g population<br />
<strong>in</strong> several places, apparently orig<strong>in</strong>at<strong>in</strong>g from illegally released specimens (Kestenholz<br />
and Heer, 2001), although the presence of wild birds that expanded their range<br />
cannot be ruled out. Greylag geese are native <strong>in</strong> an area extend<strong>in</strong>g from north-western<br />
to south-eastern Europe. They were probably released to enrich the local avifauna <strong>in</strong><br />
<strong>Switzerland</strong>. It can be assumed that the populations will grow, without hunt<strong>in</strong>g pressure.<br />
No environmental impact is expected, s<strong>in</strong>ce the Swiss population is not far from<br />
the southern border of the natural distribution, but there might be damage to crops and<br />
dropp<strong>in</strong>gs on lawns close to lakes could mean the swans are considered a nuisance.<br />
The ruddy shelduck (Tadorna ferrug<strong>in</strong>ea (Pallas)) (see Fact Sheet) is a Central Asian<br />
and northern African <strong>species</strong>, which is a favourite asset of waterfowl collections<br />
throughout Europe for its strik<strong>in</strong>g plumage of an almost orange-brown body and a<br />
whitish head. In many cases the birds are not kept <strong>in</strong> cages and regularly escape. Some<br />
isolated breed<strong>in</strong>g pairs are recorded <strong>in</strong> many countries. However, <strong>Switzerland</strong> has the<br />
only viable population of this <strong>species</strong> <strong>in</strong> Europe, and it is <strong>in</strong> the process of build<strong>in</strong>g up<br />
numbers. Its spread to neighbour<strong>in</strong>g countries should be prevented, s<strong>in</strong>ce if it were to<br />
cause subsequent damage and environmental impact, <strong>Switzerland</strong> would have some<br />
responsibility. Therefore, it is recommended that this unnecessary population should be<br />
elim<strong>in</strong>ated while it is of manageable size.<br />
The mandar<strong>in</strong> duck (Aix galericulata (L.)) (see Fact Sheet) is the only <strong>alien</strong> duck<br />
<strong>species</strong> to have established <strong>in</strong> large numbers <strong>in</strong> Europe. As it is one of the most ornate<br />
waterfowl <strong>species</strong>, it was and is frequently held on park ponds <strong>in</strong> Europe, from where it<br />
escapes <strong>in</strong>to the wild. Its orig<strong>in</strong> is the eastern part of Asia, where populations have been<br />
dramatically reduced by habitat changes (ma<strong>in</strong>ly logg<strong>in</strong>g) and over-hunt<strong>in</strong>g. Consequently,<br />
the European populations might be of some importance for the <strong>species</strong>, especially<br />
if there is no demonstrated impact. Thus, a strategy for this <strong>species</strong> <strong>in</strong> Europe<br />
might recommend accept<strong>in</strong>g it.<br />
The native range of the common pheasant (Phasianus colchicus (L.)) covers a large<br />
part of Asia with about 40 sub<strong>species</strong> described. It has been kept <strong>in</strong> pheasant exhibitions<br />
for at least a thousand years <strong>in</strong> Europe (Geiter et al., 2002). However, it is believed<br />
that it established <strong>in</strong> the wild much later, perhaps only <strong>in</strong> the 18 th century, but its<br />
history cannot be established with certa<strong>in</strong>ty, s<strong>in</strong>ce the numerous releases obscure the<br />
susta<strong>in</strong>ability of wild populations. It is a favourite game bird all over Europe and<br />
millions are released every year. Thus naturalized populations are supported by frequent<br />
releases and are harvested by hunt<strong>in</strong>g, e.g. each year up to 20 million are re-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 46<br />
leased and 12 million are shot <strong>in</strong> the UK (Kestenholz and Heer, 2001). Almost all wild<br />
populations <strong>in</strong> Europe seem to be of hybrid orig<strong>in</strong> from several sub<strong>species</strong>, because of<br />
the many releases of different genetic material. Apparently, many wild populations are<br />
not able to susta<strong>in</strong> themselves without human help, e.g., by releases and w<strong>in</strong>ter feed<strong>in</strong>g.<br />
Schmid et al. (1998) estimate a population of fewer than 1,000 for <strong>Switzerland</strong>, ma<strong>in</strong>ly<br />
<strong>in</strong> the lowland parts. It is recommended that releases of this <strong>alien</strong> <strong>species</strong> are stopped,<br />
so that naturalized populations may dw<strong>in</strong>dle away.<br />
One domestic <strong>species</strong> needs to be mentioned <strong>in</strong> this section, the feral pigeon (Columba<br />
livia f. domestica L.). It is descended from the rock dove (Columba livia L.), which<br />
lives on sea-cliffs and <strong>in</strong> mounta<strong>in</strong>s <strong>in</strong> southern Europe and the UK. The feral pigeon is<br />
abundant <strong>in</strong> most cities of Europe and elsewhere because members of the public supplement<br />
their natural food. Today they are often recognized as a problem <strong>species</strong> due to<br />
their faeces alter<strong>in</strong>g the colour and destroy<strong>in</strong>g the surfaces of old build<strong>in</strong>gs, statues and<br />
other artificial structures, and for their role <strong>in</strong> spread<strong>in</strong>g disease. Most control methods<br />
have not been successful. The ma<strong>in</strong> h<strong>in</strong>drance is feed<strong>in</strong>g, mak<strong>in</strong>g education of the<br />
public a crucial aspect of feral pigeon control <strong>in</strong> cities.<br />
Species <strong>in</strong> other European countries which are expand<strong>in</strong>g their range are discussed<br />
below, as they may enter <strong>Switzerland</strong> <strong>in</strong> the near future.<br />
The most successful avian <strong>in</strong>vader <strong>in</strong> Europe is the Canada goose (Branta canadensis<br />
(L.)), follow<strong>in</strong>g extensive releases and the availability of suitable habitat for this North<br />
American <strong>species</strong> throughout Central and northern Europe. It is rapidly <strong>in</strong>creas<strong>in</strong>g its<br />
population and expand<strong>in</strong>g its range (Delany, 1993). If this spread cont<strong>in</strong>ues, it is likely<br />
to <strong>in</strong>vade <strong>Switzerland</strong> <strong>in</strong> the near future. While s<strong>in</strong>gle <strong>in</strong>dividuals ahead of an <strong>in</strong>vasion<br />
front could be elim<strong>in</strong>ated, the spread of the <strong>species</strong> <strong>in</strong> Europe could only be addressed<br />
through an <strong>in</strong>ternational effort. Competition with native waterfowl has been frequently<br />
observed (Madsen et al., 1999) and hybridization with greylag goose is of concern <strong>in</strong><br />
countries with native populations of the latter <strong>species</strong> (Gebhardt, 1996). The grow<strong>in</strong>g<br />
populations of Canada goose are a cause for concern due to damage to crops and faeces<br />
deposited <strong>in</strong> parks and on golf ranges (Kestenholz and Heer, 2001). In spr<strong>in</strong>g pastures<br />
they cause damage by graz<strong>in</strong>g, trampl<strong>in</strong>g and foul<strong>in</strong>g. Foul<strong>in</strong>g is also responsible for<br />
eutrophication and associated algal blooms on small, still waters (Welch et al., 2001).<br />
Another goose <strong>species</strong>, which <strong>in</strong> recent times rapidly spread from a nucleus population<br />
<strong>in</strong> the Netherlands, is the Egyptian goose (Alopochen aegyptiacus (L.)). Its native<br />
range is Sub-Saharan Africa, where it is a ubiquitous <strong>species</strong>. The Egyptian goose was<br />
already present <strong>in</strong> captivity <strong>in</strong> the UK <strong>in</strong> the 17 th century, and probably around 1967<br />
some birds escaped conf<strong>in</strong>ement <strong>in</strong> the Netherlands (Bezzel, 1996). If the affected<br />
countries employ no counter-measures it seems likely that the Egyptian goose will<br />
eventually spread <strong>in</strong>to <strong>Switzerland</strong>, but that will take some time, unless further escapes<br />
occur <strong>in</strong> or close to <strong>Switzerland</strong>. Generally, as for other waterfowl, this <strong>species</strong> should<br />
be kept <strong>in</strong> closed cages to m<strong>in</strong>imize escapes. Dur<strong>in</strong>g the breed<strong>in</strong>g season, Egyptian<br />
geese are very aggressive, and this could have some impact on native waterfowl.
2 > Vertebrates – Vertebrata 47<br />
The <strong>in</strong>troduction of North American ruddy duck (Oxyura jamaicensis (Gmel<strong>in</strong>)) (see<br />
Fact Sheet) <strong>in</strong>to Europe led to one of the best-known cases of concern about an <strong>alien</strong><br />
<strong>species</strong> <strong>in</strong> relation to conservation of a globally threatened native <strong>species</strong>, i.e. the<br />
white-headed duck (Oxyura leucocephala Scopoli)), classified as Vulnerable by IUCN<br />
(the World Conservation Union) (Hughes et al., 1999). The two <strong>species</strong> readily hybridize<br />
and it is likely that unless counter-measures are taken the white-headed duck population<br />
will be completely absorbed. A European action plan for eradication of the<br />
destructive <strong>in</strong>vader has been set up to safeguard the small populations of white-headed<br />
duck <strong>in</strong> the western and eastern Mediterranean. The plan does not cover Asia, but it is<br />
to be hoped that the ruddy duck will be eradicated <strong>in</strong> cont<strong>in</strong>ental Europe and considerably<br />
reduced <strong>in</strong> the UK, so that its spread to Asia becomes unlikely. The spread of<br />
the ruddy duck with<strong>in</strong> the UK and then through Europe was facilitated by the fact that<br />
it is a migratory <strong>species</strong>. While the ‘Action Plan for the White-headed Duck’ (Green<br />
and Hughes, 1996) names eight threats and limit<strong>in</strong>g factors, the <strong>in</strong>troduction of the<br />
ruddy duck is the only one that is considered critical for its implementation. Tragically<br />
the white-headed duck population was just recover<strong>in</strong>g <strong>in</strong> Spa<strong>in</strong>, from a m<strong>in</strong>imum of 22<br />
birds <strong>in</strong> 1977 to about 2,700 today due to a conservation programme, but it is now<br />
be<strong>in</strong>g hit by hybridization with the ruddy duck. Over 4,200 ruddy ducks have now been<br />
shot <strong>in</strong> the UK, where shoot<strong>in</strong>g at large w<strong>in</strong>ter<strong>in</strong>g sites is crucial to the success of the<br />
action plan. <strong>Switzerland</strong> has endorsed the eradication plans and agreed to take action<br />
aga<strong>in</strong>st the ruddy duck with<strong>in</strong> its borders. Shoot<strong>in</strong>g of the ruddy duck has to be conducted<br />
<strong>in</strong> a co-ord<strong>in</strong>ated way by hunt<strong>in</strong>g and conservation authorities. Firstly, the<br />
correct identity has to be confirmed, s<strong>in</strong>ce the two Oxyura <strong>species</strong> can be very similar.<br />
It is clear from the current situation that ruddy ducks thrive <strong>in</strong> Europe, build up large<br />
viable populations, and spread rapidly. Therefore, keep<strong>in</strong>g ruddy ducks <strong>in</strong> captivity<br />
needs to be regulated. While the ultimate goal for such a serious problem <strong>species</strong> is the<br />
prohibition of keep<strong>in</strong>g them <strong>in</strong> captivity, an <strong>in</strong>terim measure should be the monitor<strong>in</strong>g<br />
of all specimens kept and acceptance of collections only <strong>in</strong> secure facilities.<br />
In Europe several parrot <strong>species</strong> frequently escape from captivity and some of those are<br />
able to establish <strong>in</strong> the wild under favourable conditions. Most of these colonies are <strong>in</strong><br />
parks <strong>in</strong> cities, where a rich food source of fruit-bear<strong>in</strong>g trees and bird feeders dur<strong>in</strong>g<br />
the harsh w<strong>in</strong>ter time together with temperatures on average 2° C above the surround<strong>in</strong>g<br />
landscape allow their survival. The climate seems an important factor for these<br />
<strong>species</strong> that naturally <strong>in</strong>habit lower latidudes. The most common parrot <strong>species</strong> are the<br />
rose-r<strong>in</strong>ged parakeet (Psittacula krameri (Scopoli)) and the monk parakeet (Myiopsitta<br />
monachus (Boddaert)), which have established large colonies <strong>in</strong> several European<br />
cities, <strong>in</strong>clud<strong>in</strong>g some <strong>in</strong> neighbour<strong>in</strong>g countries: France, Germany, Austria and Italy.<br />
There are s<strong>in</strong>gle reports of breed<strong>in</strong>g attempts of both <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>. They are<br />
apparently limited to city environments and it is not known whether there is any genetic<br />
exchange between the colonies <strong>in</strong> Europe. The colonies are probably isolated and<br />
will not spread that far. The rose-r<strong>in</strong>ged parakeet nests <strong>in</strong> tree cavities, so that it will<br />
compete with native wildlife. The monk parakeet is unusual, build<strong>in</strong>g large communal<br />
nests on trees, build<strong>in</strong>gs and power pylons. In their home ranges both <strong>species</strong> are<br />
recognized as agricultural pests. Other parrot <strong>species</strong> are found irregularly throughout<br />
Europe. Any parrots encountered <strong>in</strong> the wild <strong>in</strong> <strong>Switzerland</strong> should be captured.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 48<br />
This section, deal<strong>in</strong>g with <strong>alien</strong> bird <strong>species</strong> established <strong>in</strong> <strong>Switzerland</strong> and those<br />
<strong>species</strong> that are established <strong>in</strong> neighbour<strong>in</strong>g countries and spread<strong>in</strong>g, illustrates the<br />
varied importance of bird families <strong>in</strong>volved. There are four Anatidae, one Phasianidae<br />
and one Phalacrocoracidae established. Moreover, three further Anatidae are spread<strong>in</strong>g<br />
<strong>in</strong> neighbour<strong>in</strong>g countries and are expected to arrive <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> the near future.<br />
In addition, two Psittacidae are well established <strong>in</strong> cities of neighbour<strong>in</strong>g countries.<br />
Thus, Anatidae dom<strong>in</strong>ate the <strong>species</strong> list of <strong>alien</strong> birds currently establish<strong>in</strong>g colonies<br />
<strong>in</strong> the wild <strong>in</strong> Europe. Estimates of the number of established <strong>alien</strong> bird <strong>species</strong> <strong>in</strong><br />
Europe vary depend<strong>in</strong>g on the author, but for example Kestenholz and Heer (2001) list<br />
22 <strong>species</strong>, of which eight belong to the Anatidae and eight to the Phasianidae. In<br />
conclusion, the Swiss composition of <strong>alien</strong> birds mirrors the European situation, with<br />
the exception of hav<strong>in</strong>g fewer <strong>in</strong>troduced Phasianidae. The latter group has been<br />
predom<strong>in</strong>antly released <strong>in</strong> the UK and France as game birds. The six established <strong>species</strong><br />
<strong>in</strong> <strong>Switzerland</strong> represent about 3 % of all breed<strong>in</strong>g <strong>species</strong> (approximately 205). As<br />
birds are a vagile and migratory group, a comparison of breed<strong>in</strong>g <strong>species</strong> is obviously<br />
the best approach to compare native versus <strong>alien</strong> <strong>species</strong>.<br />
The total of six <strong>alien</strong> bird <strong>species</strong> established <strong>in</strong> <strong>Switzerland</strong> is comparable with neighbour<strong>in</strong>g<br />
countries (Table 2.2), especially given that Germany is much bigger. Schuster<br />
(2002) lists five <strong>species</strong> (three Anatidae, one Phasianidae, one Psittacidae) for Austria.<br />
Ten <strong>alien</strong> bird <strong>species</strong> (six Anatidae, three Phasianidae, one Psittacidae) are found <strong>in</strong><br />
Germany (Geiter et al., 2002). Italy (Andreotti et al., 2001) has about eight <strong>species</strong>, but<br />
the taxonomic composition is different with one Anatidae, one Odontophoridae, two<br />
Phasianidae, two Psittacidae, one Paradoxornithidae and one Estrildidae.
2 > Vertebrates – Vertebrata 49<br />
Tab. 2.2 > Alien birds established () <strong>in</strong> selected countries of Europe.<br />
Family Species Country<br />
Germany Austria <strong>Switzerland</strong> Italy<br />
Anatidae Aix galericulata (L.) <br />
Cygnus olor (Gmel<strong>in</strong>) native <br />
Branta canadensis (L.) <br />
Alopochen aegyptiacus (L.) <br />
Anser cygnoides (L.) <br />
Anser <strong>in</strong>dicus (Latham) <br />
Cygnus atratus (Latham) <br />
Tadorna ferrug<strong>in</strong>ea (Pallas) <br />
Anser anser (L.) native native <br />
Phasianidae Phasianus colchicus (L.) <br />
Meleagris gallopavo L. <br />
Syrmaticus reevesi (Gray) <br />
Alectoris chukar (Gray) <br />
Odontophoridae Col<strong>in</strong>us virg<strong>in</strong>ianus (L.) <br />
Phalacrocoracidae Phalacrocorax carbo L. native native native<br />
Psittacidae Psittacula krameri (Scopoli) <br />
Myiopsitta monachus (Boddaert) <br />
Paradoxornithidae Paradoxornis alphonsianus (Verreaux) <br />
Estrildidae Amandava amandava (L.) <br />
After Geiter et al., 2002 (Germany); Schuster, 2002 (Austria); this report (<strong>Switzerland</strong>); Andreotti et al., 2001 (Italy).<br />
The composition of established <strong>alien</strong> birds <strong>in</strong> <strong>Switzerland</strong> seems to reflect human<br />
activities, i.e. the frequency of release of the <strong>species</strong>, rather than their ecological traits.<br />
Therefore, an evaluation of what makes a bird <strong>species</strong> <strong>in</strong>vasive is less <strong>in</strong>formative than<br />
a discussion of the attractiveness of <strong>species</strong> to humans.<br />
Due to the small number of six established <strong>species</strong> (or 11, if we consider the spread<strong>in</strong>g<br />
<strong>species</strong> of neighbour<strong>in</strong>g countries), there is no obvious pattern to their orig<strong>in</strong>. Three of<br />
the established <strong>species</strong> are from with<strong>in</strong> Europe, two from Asia, and one from Africa/Asia.<br />
The year of <strong>in</strong>troduction (i.e. when it was first found established <strong>in</strong> the wild) varies<br />
greatly. However, the very recent significant <strong>in</strong>crease of establishment of <strong>species</strong> of<br />
Anatidae and Psittacidae, not only <strong>in</strong> <strong>Switzerland</strong> but also other European countries,<br />
gives rise to concern and po<strong>in</strong>ts to the need to prevent further <strong>in</strong>troductions.<br />
The pathways of <strong>in</strong>troduction differ greatly between the three ma<strong>in</strong> groups, but are<br />
characteristic for each group. The Anatidae are either released for aesthetic reasons<br />
(e.g. mute swan) or have escaped from the numerous ornamental waterfowl collections.<br />
The members of the Phasianidae are released as game birds for hunt<strong>in</strong>g. The Psittacidae<br />
are, of course, escapees from captivity. The success of establishment <strong>in</strong> all three
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 50<br />
groups is based on the support they receive from the human population, s<strong>in</strong>ce some are<br />
released and most are fed <strong>in</strong> the wild (or receive some other human support).<br />
The environmental and economic impact of established <strong>alien</strong> birds <strong>in</strong> <strong>Switzerland</strong> is<br />
probably fairly negligible and limited to some local effects. The exception on a European<br />
scale is the ruddy duck, which is the critical issue for a native globally endangered<br />
<strong>species</strong>. The possible and certa<strong>in</strong> impacts are summarized <strong>in</strong> Table 2.3.<br />
Birds seem to cause less concern and actual impact on biodiversity, as outl<strong>in</strong>ed above,<br />
than mammals, but their impressive capability for spread underl<strong>in</strong>es the concern about<br />
both potential future spread and the unsatisfactory level of knowledge for predict<strong>in</strong>g<br />
spread and impact. The rapid <strong>in</strong>crease <strong>in</strong> the number of newly <strong>in</strong>troduced bird <strong>species</strong><br />
recorded <strong>in</strong> recent times and the spread of some older <strong>in</strong>troductions underl<strong>in</strong>e the<br />
urgency of implement<strong>in</strong>g effective strategies to address the issue.<br />
The pathway analysis <strong>in</strong>dicates where prevention of future <strong>in</strong>troductions will be most<br />
efficient. The three major pathways identified are listed below, together with measures<br />
to help close them.<br />
Escaped birds from captivity (Psittacidae and Anatidae). Measures to prevent escapes<br />
of <strong>alien</strong> bird <strong>species</strong> from captive collections can <strong>in</strong>clude strict standards of security for<br />
aviaries, a register and documented bird monitor<strong>in</strong>g, and penal and adm<strong>in</strong>istrative<br />
sanctions <strong>in</strong> the event of violation.<br />
Released birds for aesthetic reasons and to enrich the native fauna (Anatidae). Legislation<br />
to prevent deliberate <strong>in</strong>troductions should be established, or improved and implemented.<br />
As can be easily seen from the cases of <strong>species</strong> spread<strong>in</strong>g through Europe<br />
towards <strong>Switzerland</strong>, this issue needs to be tackled on a European scale. The <strong>in</strong>ternational<br />
conventions are <strong>in</strong> place and need to be implemented, e.g. Convention on Biological<br />
Diversity (CBD), Bern Convention, Bonn Convention, Ramsar Convention. It<br />
should be noted here that the domestic duck can also pose a problem for the wild<br />
mallard (Anas platyrhynchos (L.)) because of hybridization. These noticeable hybrids<br />
are most often observed <strong>in</strong> urban areas, but they should be elim<strong>in</strong>ated <strong>in</strong> the wild.<br />
Species released as game birds (Phasianidae). An environmental risk assessment needs<br />
to be undertaken for all <strong>species</strong> considered for release <strong>in</strong> Europe. In the past chukar<br />
(Alectoris chukar (Gray)) were released <strong>in</strong> the Alps, <strong>in</strong>clud<strong>in</strong>g <strong>Switzerland</strong>. They can<br />
hybridize with the native rock partridge (Alectoris graeca (Meisner)). The chukar is the<br />
eastern equivalent of the rock partridge. These releases are unnecessary and pose a<br />
potential danger to the <strong>in</strong>digenous rock partridge, which is one of the n<strong>in</strong>e European<br />
endemic bird <strong>species</strong>.<br />
Management options for the <strong>species</strong> which can cause problems are detailed <strong>in</strong> the text<br />
relat<strong>in</strong>g to the <strong>species</strong> and <strong>in</strong> the Fact Sheets, e.g. ruddy duck and ruddy shelduck<br />
should be shot.
2 > Vertebrates – Vertebrata 51<br />
Four ma<strong>in</strong> recommendations are drawn from this compilation:<br />
> Based on <strong>in</strong>ternational obligations to safeguard the globally endangered whiteheaded<br />
duck, the ruddy duck has to be elim<strong>in</strong>ated (shot) whenever it is found <strong>in</strong><br />
<strong>Switzerland</strong>. Furthermore, the <strong>species</strong> should not be kept <strong>in</strong> captivity.<br />
> <strong>Switzerland</strong> has a responsibility <strong>in</strong> relation to the potential spread of the ruddy<br />
shelduck. It should prevent the spread of the Swiss population and consider eradicat<strong>in</strong>g<br />
it.<br />
> Birds held <strong>in</strong> captivity should be monitored closely and escape prevented. Conta<strong>in</strong>ment<br />
of birds under semi-wild conditions should be restricted or prohibited.<br />
> All releases of birds <strong>in</strong>to the wild should be subject to authorization, whereby releases<br />
of <strong>alien</strong> <strong>species</strong> should be avoided and releases of native birds should be made<br />
us<strong>in</strong>g genetic material typical of the region.<br />
Tab. 2.3 > Established birds (neozoa) <strong>in</strong> <strong>Switzerland</strong> and <strong>species</strong> to watch for (last three <strong>species</strong>).<br />
Scientific name Family Orig<strong>in</strong> Year Pathway Impact Note<br />
Phalacrocorax Phalacro- Europe 2000 Escaped from Breed<strong>in</strong>g colonies destroy trees and First breed<strong>in</strong>g attempts recorded<br />
carbo L. coracidae captivity other vegetation below trees.<br />
Impact on fishery heatedly debated.<br />
Fish farms<br />
Cygnus olor Anatidae North-eastern 1690 Releases for Decl<strong>in</strong>e of submerged aquatic A much-liked <strong>species</strong><br />
(Gmel<strong>in</strong>) Europe aesthetic reasons vegetation<br />
Anser anser (L.) Anatidae Parts of 1983 Released as an Probably no environmental impact Wild form of the domestic goose<br />
Europe enrichment of the<br />
avifauna<br />
Damage of crop possible<br />
Tadorna ferrugi- Anatidae Central Asia 1997 Escaped from Aggressive behaviour towards other CH has responsibility for the only<br />
nea (Pallas) and northern captivity waterfowl viable population <strong>in</strong> the <strong>in</strong>troduced<br />
Africa European range<br />
Aix galericulata Anatidae East-Asia 1958 Escaped from Competition for tree holes with other Population is small <strong>in</strong> CH, but other<br />
(L.) captivity cavity breed<strong>in</strong>g <strong>species</strong>? European populations expand<strong>in</strong>g<br />
Phasianus Phasianidae Asia 18th Released as Competition with native game birds? Pheasant densities are ma<strong>in</strong>ly detercolchicus<br />
(L.) cen- game Indirect effects by predator control to m<strong>in</strong>ed by the extent of releases<br />
tury? relieve pheasants from predation<br />
Branta canadensis Anatidae North America - Released as an Competition with native waterfowl Has not yet reached <strong>Switzerland</strong><br />
(L.) enrichment of the Hybridization with greylag goose<br />
avifauna Damage to crops<br />
Dropp<strong>in</strong>gs can be a nuisance and<br />
cause eutrophication <strong>in</strong> water<br />
Alopochen Anatidae Sub-Saharan - Escaped from Aggressive behaviour towards other Has not yet reached <strong>Switzerland</strong><br />
aegyptiacus (L.) Africa captivity waterfowl<br />
Oxyura jamaicen- Anatidae North America - Escaped from Hybridization with the globally There is no established population,<br />
sis (Gmel<strong>in</strong>) captivity endangered white-headed duck only records of s<strong>in</strong>gle birds. These<br />
threatens the ext<strong>in</strong>ction of the latter need to be shot under an EU-wide<br />
<strong>species</strong> effort.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 52<br />
2.3 Reptiles – Reptilia<br />
Three snake <strong>species</strong> have been relocated with<strong>in</strong> <strong>Switzerland</strong>, from southern locations to<br />
colder, northern localities. Two of these relocations were very local. The re<strong>in</strong>troduction<br />
of the European pond terrap<strong>in</strong> used <strong>alien</strong> genetic material. The only <strong>alien</strong><br />
reptile <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> as a country are aquatic turtles released from aquariums,<br />
but they seem not to have established populations yet (Table 2.4).<br />
The case of the Italian wall lizard (Podarcis sicula (Raf<strong>in</strong>esque-Schmaltz)) <strong>in</strong> <strong>Switzerland</strong><br />
is rather mysterious (Hofer et al., 2001). It is not known whether the <strong>species</strong> is<br />
native or <strong>alien</strong> nor is it proven that an established population exists at all. S<strong>in</strong>ce specimens<br />
were found along the railway tracks <strong>in</strong> the Tic<strong>in</strong>o, an accidental <strong>in</strong>troduction<br />
from Italy seems likely.<br />
The dice snake (Natrix tessellata (Laurenti)) is one of the most endangered snakes <strong>in</strong><br />
<strong>Switzerland</strong>, because it is at the north-western limit of its range (thus it is naturally<br />
rare). It is native to the Tic<strong>in</strong>o, but was released north of the Alps at several lakes<br />
(Gruschwitz et al., 1999). Although it is rare <strong>in</strong> <strong>Switzerland</strong>, it should not be relocated<br />
to the north of the Alps. At the Lac de Genève, where it now occurs together with<br />
another rare snake <strong>species</strong>, the viper<strong>in</strong>e snake (Natrix maura (L.)), it is probably compet<strong>in</strong>g<br />
with the native <strong>species</strong> for food and habitat, s<strong>in</strong>ce the two <strong>species</strong> have a rather<br />
similar biology, one occurr<strong>in</strong>g <strong>in</strong> south-western Europe and the other <strong>in</strong> south-eastern<br />
Europe (Hofer et al., 2001).<br />
Small populations of the western whip snake (Coluber viridiflavus (Lacépède)) might<br />
have established at the Neuenburgersee and <strong>in</strong> the Valais, from specimens collected <strong>in</strong><br />
the Tic<strong>in</strong>o and released <strong>in</strong> these areas. The impacts of these populations are probably<br />
negligible and the populations themselves might not persist. Populations of the Aesculapian<br />
snake (Elaphe longissima (Laurenti)) at the Neuenburgersee and Bieler See<br />
may have orig<strong>in</strong>ated <strong>in</strong> the same way and aga<strong>in</strong> have little impact. The latter <strong>species</strong><br />
does seem to be established (Hofer et al., 2001).<br />
The status of the European pond terrap<strong>in</strong> (Emys orbicularis (L.)) <strong>in</strong> <strong>Switzerland</strong> is<br />
not yet resolved (Hofer et al., 2001). Some populations might still be native, but there<br />
were many re-<strong>in</strong>troductions of the <strong>species</strong>. In most <strong>in</strong>stances the orig<strong>in</strong> of the specimens<br />
released is not known, but releases of non-native material could lead to genetic<br />
<strong>in</strong>trogression <strong>in</strong>to possibly surviv<strong>in</strong>g native populations. However, this concern rema<strong>in</strong>s<br />
hypothetical because of the lack of knowledge of the status of extant populations.<br />
The North American red-eared slider (Trachemys scripta (Seidel)) (see Fact Sheet) is<br />
representative here of a guild of potential IAS – aquatic turtles from several genera. In<br />
past years, several countries restricted the importation of this <strong>species</strong>, because of the<br />
potential threat. However, this merely leads to a shift to other <strong>species</strong> by the aquarium<br />
trade. The red-eared slider is still the <strong>species</strong> most often found <strong>in</strong> the wild <strong>in</strong> high<br />
numbers and many places, although, because of the direct releases <strong>in</strong> parks etc., the
2 > Vertebrates – Vertebrata 53<br />
<strong>species</strong> is predom<strong>in</strong>antly recorded around agglomerations (Geiger and Waitzmann,<br />
1996). The red-eared slider is probably not yet established <strong>in</strong> the wild <strong>in</strong> <strong>Switzerland</strong>,<br />
but this might change through adaptation and future releases of specimens from a more<br />
northern natural distribution <strong>in</strong> North America. Reproduction has been repeatedly<br />
observed <strong>in</strong> areas of Europe with a Mediterranean climate. However, even without<br />
reproduction, the populations <strong>in</strong> certa<strong>in</strong> areas of <strong>Switzerland</strong> are extremely high ow<strong>in</strong>g<br />
to frequent releases and the longevity of the <strong>species</strong>. Therefore there is the potential for<br />
native biodiversity to be affected, even if reproduction fails. A recent study by Cadi<br />
and Joly (2004) found both weight loss and high mortality <strong>in</strong> the native turtle Emys<br />
orbicularis (L.) <strong>in</strong> mixed groups and argues for apply<strong>in</strong>g a precautionary pr<strong>in</strong>ciple. The<br />
<strong>in</strong>troduced <strong>species</strong> also outcompetes the native turtle for preferred bask<strong>in</strong>g places (Cadi<br />
and Joly, 2003). The red-eared slider is one of the <strong>species</strong> mentioned <strong>in</strong> the Recommendations<br />
of the Bern Convention, as described above.<br />
S<strong>in</strong>ce no <strong>alien</strong> reptile <strong>species</strong> has established populations <strong>in</strong> <strong>Switzerland</strong>, no general<br />
pattern can be discussed here, except to note that the situation <strong>in</strong> neighbour<strong>in</strong>g countries<br />
is rather similar.<br />
Reptiles depend to a large extent on climate, especially temperature, because they can<br />
regulate their body temperature only to a certa<strong>in</strong> degree. Thus, tropical and subtropical<br />
<strong>species</strong>, which are most frequently kept <strong>in</strong> terrariums, are not of concern, apart from<br />
escapes of the occasional crocodilian or dangerous snake. However, this is more a<br />
matter of <strong>in</strong>terest for the media than a true threat. A real concern for native biodiversity<br />
are <strong>species</strong> from North America and east Asia (Ch<strong>in</strong>a), ma<strong>in</strong>ly turtles, which could<br />
become established <strong>in</strong> the Swiss climate. Two measures should be implemented to<br />
restrict this threat:<br />
1. Rais<strong>in</strong>g awareness <strong>in</strong> the human population of the potential problem, and especially<br />
<strong>in</strong> herpetological clubs and organizations.<br />
2. Implement<strong>in</strong>g laws to m<strong>in</strong>imize releases and escapes of <strong>species</strong> which could potentially<br />
cause problems, e.g. article 25 of the Eidgenössisches Tierschutzgesetz (Swiss<br />
Animal Protection Act) stipulates the penalties for releases of this k<strong>in</strong>d.<br />
Tab. 2.4 > Established <strong>alien</strong> reptiles <strong>in</strong> <strong>Switzerland</strong>.<br />
Scientific name Family Orig<strong>in</strong> Year Pathway Impact Note<br />
Natrix tessellata Colubridae Tic<strong>in</strong>o – 1920s Released Competition with Natrix maura Relocation with<strong>in</strong> <strong>Switzerland</strong><br />
(Laurenti) <strong>Switzerland</strong><br />
Emys orbicularis (L.) Emydidae ? 1800s? Released Genetic <strong>in</strong>trogression with native populations? Re-<strong>in</strong>troduction<br />
Trachemys scripta Emydidae North America Fairly recent Released Competition with native turtles <strong>in</strong> Europe Perhaps not established, but<br />
(Seidel) Escaped? Predator<br />
Destroy<strong>in</strong>g and disturb<strong>in</strong>g float<strong>in</strong>g bird nests<br />
long-lived
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 54<br />
2.4 Amphibians – Amphibia<br />
Currently 20 amphibian <strong>species</strong> are found <strong>in</strong> <strong>Switzerland</strong>. One <strong>species</strong> (Rana ridibunda<br />
Pallas), i.e. 5 % of the total Swiss <strong>species</strong> number, is <strong>in</strong>troduced and established and<br />
will be discussed below and <strong>in</strong> a Fact Sheet. A second <strong>species</strong> has been relocated<br />
with<strong>in</strong> <strong>Switzerland</strong> (Triturus carnifex (Laurenti)) and a third <strong>species</strong> is found occasionally<br />
but has not yet been able to establish a population, i.e. the North American bullfrog<br />
Rana catesbeiana Shaw (Table 2.5).<br />
The marsh frog (Rana ridibunda) (see Fact Sheet) is the only <strong>in</strong>troduced <strong>alien</strong> amphibian<br />
<strong>species</strong> at the national level. It was <strong>in</strong>troduced between 1920 and 1950, probably<br />
from Hungary, as was the case with specimens released <strong>in</strong> the UK (Zeisset and<br />
Beebee, 2003). The most probable explanation for the <strong>in</strong>troductions is that they were<br />
imports for human consumption. The marsh frog has now established with a wide<br />
distribution <strong>in</strong> <strong>Switzerland</strong>, ma<strong>in</strong>ly <strong>in</strong> the west and the Valais and <strong>in</strong> the north-eastern<br />
region. The <strong>species</strong> should not be supported, as native amphibian <strong>species</strong> are, by local<br />
conservation groups, and releases on conservation grounds or as a food resource need<br />
to be stopped. The role of the genetic peculiarities <strong>in</strong> the green frog complex are not<br />
fully understood, but <strong>in</strong> most places where the marsh frog is <strong>in</strong>creas<strong>in</strong>g the other two<br />
related <strong>species</strong> are decreas<strong>in</strong>g. Thus, a displacement is evident.<br />
Another <strong>species</strong> <strong>in</strong>troduced <strong>in</strong>to a new range is the Italian crested newt (Triturus<br />
carnifex) (see Fact Sheet), which is native to the south of <strong>Switzerland</strong> (Tic<strong>in</strong>o) but was<br />
<strong>in</strong>troduced to the canton of Genève. Genetic studies showed that the Genève population<br />
is closest genetically to a population <strong>in</strong> Tuscany, Italy (Arntzen, 2001). Most probably<br />
the <strong>species</strong> was imported for zoological experiments and released <strong>in</strong>to a pond at the<br />
University of Genève. It completely replaced the closely related great crested newt<br />
(Triturus cristatus (Laurenti)) <strong>in</strong> that area. However, it is a complex situation and there<br />
is hybridization between the two <strong>species</strong>. The distribution and spread of the Italian<br />
crested newt should be monitored to provide <strong>in</strong>formation for future decisions about this<br />
<strong>species</strong> and its threat.<br />
The American bullfrog (Rana catesbeiana) (see Fact Sheet) has not (yet) colonized<br />
<strong>Switzerland</strong>, but there are some recent unconfirmed records of the <strong>species</strong>. However, it<br />
is established <strong>in</strong> the neighbour<strong>in</strong>g countries of Germany, France, and especially Italy <strong>in</strong><br />
the Po valley with some rapidly expand<strong>in</strong>g populations. Research has demonstrated its<br />
high potential to cause a decl<strong>in</strong>e <strong>in</strong> native amphibians and reptiles. Thus, bullfrogs<br />
should not be tolerated <strong>in</strong> the wild. Monitor<strong>in</strong>g of suspect water bodies and rais<strong>in</strong>g<br />
awareness <strong>in</strong> the human population for this potential menace to native biodiversity is of<br />
high priority.<br />
In <strong>Switzerland</strong> only one frog is <strong>in</strong>troduced and established, this is 5 % of the 20 amphibian<br />
<strong>species</strong> currently found <strong>in</strong> <strong>Switzerland</strong>. In Austria (Schuster and Rabitsch,<br />
2002) and Germany (Geiter et al., 2002) no established amphibians are listed, although<br />
it is possible that the American bullfrog has established unnoticed <strong>in</strong> the recent past –
2 > Vertebrates – Vertebrata 55<br />
there is an <strong>in</strong>creas<strong>in</strong>g number of records from Germany. The <strong>in</strong>troduced <strong>species</strong> <strong>in</strong><br />
<strong>Switzerland</strong>, the marsh frog, is native <strong>in</strong> Austria and Germany.<br />
Species of edible frogs may be illegally released as a food resource and game animal.<br />
In certa<strong>in</strong> areas, notably western <strong>Switzerland</strong>, R. ridibunda is still imported <strong>in</strong> large<br />
numbers for human consumption and it is likely that some escape. This has been<br />
particularly true <strong>in</strong> the past with less rigid control of transport and conta<strong>in</strong>ment. Thus,<br />
the current populations have most probably developed from these escapees. It seems<br />
likely also that some amphibians are released or escape from captivity as pets. Thus the<br />
pet trade should have a responsibility to educate the public about the potential threat of<br />
<strong>alien</strong> <strong>species</strong> to native biodiversity. Trade <strong>in</strong> certa<strong>in</strong> <strong>species</strong>, such as the American<br />
bullfrog, should be prohibited.<br />
All three <strong>species</strong> discussed here are known to cause severe damage to native amphibians<br />
by prey<strong>in</strong>g on the smaller <strong>species</strong>, through competition as tadpoles, as vector of<br />
diseases, or through hybridization. No negative economic impact has been reported.<br />
The major recommendations to reduce the risks caused by <strong>alien</strong> amphibian <strong>species</strong> are:<br />
> A stricter regulation of the (pet) trade and implementation of laws and conventions.<br />
> Besides the legislative options, the public needs to be made aware of the potential<br />
threats to native biodiversity. Some of them might voluntarily act <strong>in</strong> a more responsible<br />
way.<br />
> Any established populations of American bullfrogs should be eradicated.<br />
Tab. 2.5 > Established <strong>alien</strong> amphibians <strong>in</strong> <strong>Switzerland</strong>.<br />
Scientific name Family Orig<strong>in</strong> Year Pathway Impact Note<br />
Rana ridibunda Ranidae East Europe, 1920s? Released (as food resource?) Competition with native amphibians Spread<strong>in</strong>g<br />
Pallas probably Hungary Escaped from imports for Genetic changes <strong>in</strong> native green frog<br />
human consumption complex<br />
Triturus carnifex Salaman- South Europe, Few Released or escaped from Replaced the great crested newt Native to <strong>Switzerland</strong><br />
(Laurenti) dridae <strong>in</strong>clud<strong>in</strong>g the Tic<strong>in</strong>o decades conta<strong>in</strong>ment for scientific<br />
ago studies<br />
Rana catesbeiana Ranidae Central and eastern Recent Released or escaped Feed<strong>in</strong>g on native amphibians and Probably no breed<strong>in</strong>g<br />
Shaw North America reptiles population <strong>in</strong> CH, as<br />
Competition with native amphibians<br />
Disease vector<br />
yet<br />
2.5 Fish – Pisces<br />
The situation with regard to <strong>in</strong>troduced fish <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> is <strong>in</strong>terest<strong>in</strong>g because,<br />
with the exception of asp (Aspius aspius (L.)), all other <strong>species</strong> are <strong>in</strong>dividually<br />
mentioned <strong>in</strong> Appendixes 2 or 3 of the Fisheries Act of 1993 (Ord<strong>in</strong>ance relevant to the<br />
federal fisheries law). In this Act, Appendix 2 describes situations (garden ponds,
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 56<br />
aquaculture plants, etc.) <strong>in</strong> which certa<strong>in</strong> <strong>alien</strong> fish taxa can be released without authorization,<br />
while Appendix 3 names fish taxa which are unwanted <strong>in</strong> <strong>Switzerland</strong>. In<br />
this section, <strong>species</strong> mentioned <strong>in</strong> these two Appendixes are referred to as Appendix 2<br />
and Appendix 3 <strong>species</strong>, respectively. A total of 15 <strong>species</strong> will be discussed below<br />
(Table 2.7), although <strong>in</strong> some cases natural breed<strong>in</strong>g has not yet been reported from<br />
<strong>Switzerland</strong>. In such cases, populations are based on frequent releases. However, the<br />
<strong>species</strong> lists <strong>in</strong> Appendixes 2 and 3 suggest all 15 <strong>species</strong> should be dealt with here,<br />
s<strong>in</strong>ce they are regulated <strong>in</strong> <strong>Switzerland</strong>. The common carp (Cypr<strong>in</strong>us carpio L.) is not<br />
<strong>in</strong>cluded <strong>in</strong> the <strong>species</strong> discussed <strong>in</strong> this section, because it is assumed to be an archaeozoa,<br />
arriv<strong>in</strong>g with the Romans <strong>in</strong> Central Europe. It was also found <strong>in</strong> most of<br />
Europe before the glacial era. Organisms which occurred before glacial times <strong>in</strong> Central<br />
Europe often stimulate discussions about def<strong>in</strong>itions of native and <strong>alien</strong> <strong>species</strong>.<br />
The bighead carp (Aristichthys nobilis Richardson) (see Fact Sheet) has not yet been<br />
reported to reproduce <strong>in</strong> <strong>Switzerland</strong>. However, it is assumed that the <strong>species</strong> could<br />
breed, and that is the reason for add<strong>in</strong>g the <strong>species</strong> to Appendix 3. It is reported to<br />
breed <strong>in</strong> the Danube (Donau).<br />
The asp (Aspius aspius) is a very recent <strong>in</strong>vader <strong>in</strong> <strong>Switzerland</strong>, first recorded <strong>in</strong> 1994<br />
<strong>in</strong> the Rh<strong>in</strong>e (Rhe<strong>in</strong>) at Basel (Zaugg et al., 2003). It is probably expand<strong>in</strong>g its range<br />
follow<strong>in</strong>g its <strong>in</strong>troduction <strong>in</strong>to the Rh<strong>in</strong>e <strong>in</strong> Germany, downstream of Basel. Its natural<br />
range seems to be Central and eastern Europe from Germany eastwards, <strong>in</strong>clud<strong>in</strong>g the<br />
Danube system. The Rh<strong>in</strong>e population was either <strong>in</strong>troduced for fish<strong>in</strong>g purposes<br />
(Ladiges and Vogt, 1979) or spread from the Danube after the completion of the Rh<strong>in</strong>e–Danube<br />
Canal. This is the only <strong>alien</strong> fish <strong>species</strong> which was not <strong>in</strong>troduced directly<br />
<strong>in</strong>to <strong>Switzerland</strong> but <strong>in</strong>to a neighbour<strong>in</strong>g country, <strong>in</strong> this case Germany, and is naturally<br />
spread<strong>in</strong>g up the river Rh<strong>in</strong>e. The asp is a large fish (up to one metre long) and prefers<br />
large rivers. It is one of the few cypr<strong>in</strong>ids that is a piscivore. The adults eat ma<strong>in</strong>ly fish,<br />
but also mammals and birds, while the more gregarious liv<strong>in</strong>g young eat smaller animals<br />
such as <strong>in</strong>vertebrates. It is assumed that the <strong>species</strong> will colonize more parts of<br />
<strong>Switzerland</strong>. S<strong>in</strong>ce the natural range is close to <strong>Switzerland</strong>, the <strong>species</strong> might be<br />
acceptable, as a natural expansion rather than an <strong>in</strong>vasion.<br />
The goldfish (Carassius auratus (L.)) is a favourite <strong>species</strong> for aquariums and garden<br />
and park ponds. Today, it has acquired an almost pan-global distribution through its<br />
ornamental use. It either escapes or is released <strong>in</strong>to the wild. Its native range is Central<br />
and eastern Asia. The Carassius <strong>species</strong> (two more are discussed below) are difficult to<br />
identify and some records might be misidentifications (Arnold, 1990). Goldfish feed on<br />
a wide range of food <strong>in</strong>clud<strong>in</strong>g plants and small animals. In some places they are<br />
regarded as a nuisance due to the production of stunted populations. They produce<br />
large numbers of <strong>in</strong>dividuals, which mature at a much reduced size, thereby dim<strong>in</strong>ish<strong>in</strong>g<br />
the usefulness of the population for sport or commercial fish<strong>in</strong>g (Lehtonen, 2002).<br />
There are <strong>in</strong>dications that they compete with native fish <strong>species</strong> and <strong>in</strong>crease turbidity<br />
through their bottom dwell<strong>in</strong>g behaviour, as does the common carp, thereby alter<strong>in</strong>g<br />
the aquatic community. Introductions limited to closed systems are possible without<br />
authorization (Appendix 2 <strong>species</strong>).
2 > Vertebrates – Vertebrata 57<br />
The crucian carp (Carassius carassius (L.)) is probably native to Central and eastern<br />
Europe, but was widely distributed by humans throughout western Europe <strong>in</strong> the<br />
Middle Ages for fish<strong>in</strong>g. It was <strong>in</strong> the recent past, if not still, be<strong>in</strong>g sold as a bait fish.<br />
The crucian carp is a hardy <strong>species</strong> that can survive adverse conditions, such as low<br />
oxygen levels and frost. It is also an Appendix 2 <strong>species</strong>, and therefore its use as a bait<br />
fish is illegal. The crucian carp is rarely found <strong>in</strong> <strong>Switzerland</strong>.<br />
The third Carassius <strong>species</strong> is the Prussian carp (Carassius gibelio (Bloch)). The<br />
taxonomy of the three Carassius <strong>species</strong> is complicated and this <strong>species</strong> is sometimes<br />
considered conspecific with C. auratus. The <strong>species</strong> are very similar <strong>in</strong> many ways,<br />
<strong>in</strong>clud<strong>in</strong>g appearance, biology, food and potential impact. The actual distribution and<br />
the history of <strong>in</strong>troductions and spread are not well known, s<strong>in</strong>ce the <strong>species</strong> were<br />
probably often misidentified (Arnold, 1990). The Prussian carp is listed <strong>in</strong> Appendix 2,<br />
thus its release outside conta<strong>in</strong>ed captivity must be authorized.<br />
The grass carp (Ctenopharyngodon idella (Cuvier & Valenciensis) (see Fact Sheet) is<br />
of Ch<strong>in</strong>ese orig<strong>in</strong>, but as it is one of the most important aquacultural fish, it has acquired<br />
a wide distribution on five cont<strong>in</strong>ents. Although natural reproduction has not<br />
been reported <strong>in</strong> the wild <strong>in</strong> <strong>Switzerland</strong>, it is a <strong>species</strong> of concern because of its potential<br />
to cause massive impacts on ecosystems by remov<strong>in</strong>g higher aquatic plants and<br />
thereby caus<strong>in</strong>g shifts <strong>in</strong> the producer guilds to other plant <strong>species</strong>. This impact at the<br />
bottom of the food cha<strong>in</strong> can cause major alterations to ecosystems. The grass carp<br />
showed some potential to control water weeds <strong>in</strong> <strong>Switzerland</strong>, but Müller (1995)<br />
concludes that it only controls the symptoms of eutrophication <strong>in</strong>stead of ameliorat<strong>in</strong>g<br />
the causes of the deteriorat<strong>in</strong>g water quality. The grass carp is listed <strong>in</strong> Appendix 3 and<br />
therefore all releases are prohibited.<br />
The silver carp (Hypophthalmichthys molitrix (Valenciensis)) (see Fact Sheet) is a<br />
highly specialized phytoplankton feeder. In many cases it was released to reduce<br />
phytoplankton densities or stop algal blooms. Its orig<strong>in</strong> is Ch<strong>in</strong>a, but today it is found<br />
<strong>in</strong> many countries around the world. Spawn<strong>in</strong>g requires very specific conditions, but<br />
has been recorded <strong>in</strong> the Danube. Its reproduction <strong>in</strong> <strong>Switzerland</strong> cannot be ruled out<br />
with certa<strong>in</strong>ty, so the <strong>species</strong> was <strong>in</strong>cluded <strong>in</strong> Appendix 3.<br />
The stone moroko (Pseudorasbora parva (Temm<strong>in</strong>ck & Schlegel)) (see Fact Sheet) is<br />
a very small cypr<strong>in</strong>id. It is the only established fish <strong>species</strong> accidentally <strong>in</strong>troduced to<br />
Europe and <strong>Switzerland</strong>, with shipments of grass carp from Ch<strong>in</strong>a. Whereas the deliberately<br />
<strong>in</strong>troduced fish <strong>species</strong> are of value to the commercial or sports sectors, the<br />
stone moroko has no value to humans. Thus, the potential for conflicts over management<br />
and legislative measures for the <strong>species</strong> is reduced and it is an Appendix 3 <strong>species</strong>.<br />
The American catfish (Ameiurus melas and A. nebulosus (Le Sueur)) (see Fact Sheet)<br />
belong to the family Ictaluridae, which is restricted to the subtropical and temperate<br />
zones of North America, and were <strong>in</strong>troduced <strong>in</strong>to Europe to <strong>in</strong>vestigate their potential<br />
as a fish for human consumption. However, they are of little value. The two <strong>species</strong> are
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 58<br />
very similar not only <strong>in</strong> appearance but also <strong>in</strong> their biology. They are listed <strong>in</strong> Appendix<br />
3, because of their potential negative impacts on native biodiversity.<br />
The brightly coloured pumpk<strong>in</strong>seed (Lepomis gibbosus (L.)) (see Fact Sheet) is an<br />
example of an ornamental fish <strong>in</strong>troduced <strong>in</strong>to Europe. It has an <strong>in</strong>terest<strong>in</strong>g breed<strong>in</strong>g<br />
behaviour and is very showy. However, <strong>in</strong> some cases the <strong>species</strong> flourished and has<br />
reached high densities. In these circumstances, this predatory fish will almost certa<strong>in</strong>ly<br />
have an impact on the food web through selective feed<strong>in</strong>g. It is an Appendix 3 <strong>species</strong>.<br />
One of the most popular sport fish <strong>in</strong> North America, the largemouth bass (Micropterus<br />
salmoides (Lacépède) (see Fact Sheet), was widely distributed <strong>in</strong> North America<br />
outside its natural range, and imported to Europe <strong>in</strong> the 1880s. The adult is a specialized<br />
fish predator and a decl<strong>in</strong>e <strong>in</strong> native fish <strong>species</strong> was observed <strong>in</strong> Italy after its<br />
<strong>in</strong>troduction (Welcomme, 1988). It was also one factor, amongst others, <strong>in</strong> the ext<strong>in</strong>ction<br />
of the Atitlán grebe (Podilymbus gigas Griscom) which was endemic to Guatemala<br />
(BirdLife International, 2000). The bird’s population dropped drastically to 80 as<br />
a result of competition and predation by the <strong>in</strong>troduced largemouth bass, but recovered<br />
to a high of 232 <strong>in</strong> 1975 when the numbers of the bass plummeted (LaBastille, 1984).<br />
Later the grebe became ext<strong>in</strong>ct through other factors.<br />
The only established <strong>alien</strong> Percidae is the pike-perch (Sander lucioperca (L.)), a<br />
<strong>species</strong> from Central and eastern Europe. It is one of the most popular sport fish<strong>in</strong>g<br />
<strong>species</strong> and a highly priced commercial fish. Hence it has been widely released and has<br />
built up self-susta<strong>in</strong><strong>in</strong>g populations. It prefers large rivers and lakes, where it is a<br />
ferocious solitary pelagic predator. In the UK negative impacts on native fish populations<br />
have been confirmed; populations of Esox lucius L. and Perca fluviatilis L.<br />
decl<strong>in</strong>ed after the <strong>in</strong>troduction of pike-perch (Welcomme, 1988). In <strong>Switzerland</strong>,<br />
releases of the pike-perch without authorization are allowed <strong>in</strong> conta<strong>in</strong>ed water bodies<br />
and where the pike-perch already occurs without negative effects on the fauna and flora<br />
(Appendix 2 <strong>species</strong>). However, the latter is difficult to prove or disprove, so that this<br />
predatory <strong>species</strong> can be released <strong>in</strong> many open waters. It was found <strong>in</strong> 137 localities<br />
dur<strong>in</strong>g the survey for the fish atlas of <strong>Switzerland</strong> (Zaugg et al., 2003). More rigid<br />
legislation would be desirable for this <strong>species</strong>.<br />
The ra<strong>in</strong>bow trout (Oncorhynchus mykiss Walbaum) is probably the most widely<br />
distributed freshwater fish <strong>species</strong> and may be regarded as a <strong>species</strong> of global distribution<br />
today. It is a highly prized game fish as well as be<strong>in</strong>g widely valued for commercial<br />
use. In <strong>Switzerland</strong> its natural reproduction is suspected but has been proved only<br />
<strong>in</strong> the ‘Alp<strong>in</strong>e Rh<strong>in</strong>esystem’. The observed populations are probably based on extensive<br />
stock<strong>in</strong>g of the <strong>species</strong>. Zaugg et al. (2003) found it <strong>in</strong> 39 % of the Swiss lakes and<br />
regarded it as a common <strong>species</strong>. A self-susta<strong>in</strong><strong>in</strong>g population is undesirable, because it<br />
would be difficult to control its spread or prevent competition for breed<strong>in</strong>g grounds<br />
with the native brown trout (Salmo trutta ssp. fario L.). Where ra<strong>in</strong>bow trout is extensively<br />
released, it will have a negative impact on native salmonids. Mahan (2002)<br />
showed that the <strong>in</strong>troduction of the ra<strong>in</strong>bow trout <strong>in</strong>to a North American lake caused a<br />
decl<strong>in</strong>e of an endemic congeneric <strong>species</strong> (O. negratis) with local ext<strong>in</strong>ctions. Drake<br />
and Naiman (2000) expla<strong>in</strong> the impact on the habitat. The exception of authorization
2 > Vertebrates – Vertebrata 59<br />
(Appendix 2 <strong>species</strong>) for stock<strong>in</strong>g <strong>in</strong> alp<strong>in</strong>e lakes should be re-addressed, because of<br />
the potential impact on amphibians. Amphibian populations <strong>in</strong> naturally fishless ponds<br />
and lakes will suffer from <strong>in</strong>troductions of <strong>alien</strong> predatory fish <strong>species</strong>.<br />
A favourite <strong>species</strong> for fly-fish<strong>in</strong>g, the brook trout (Salvel<strong>in</strong>us font<strong>in</strong>alis (Mitchill)), is<br />
today a common <strong>species</strong> <strong>in</strong> alp<strong>in</strong>e and subalp<strong>in</strong>e lakes of <strong>Switzerland</strong> (Zaugg et al.,<br />
2003). This <strong>species</strong> probably competes with the native brown trout. Impacts on lake<br />
communities have not been studied <strong>in</strong> <strong>Switzerland</strong>. In a North American study Bechara<br />
et al. (1992) <strong>in</strong>vestigated the impact of brook trout on native communities. Overall,<br />
their results suggest that size-selective predation by brook trout can cause profound<br />
changes <strong>in</strong> the structure of epibenthic communities at primary as well as secondary<br />
trophic levels. Releases of this <strong>species</strong> are restricted, s<strong>in</strong>ce it is also an Appendix 2<br />
<strong>species</strong>.<br />
The lake trout (Salvel<strong>in</strong>us namaycush (Walbaum)) was chosen to stock many high<br />
altitude lakes based on its cold-tolerant, northern distribution <strong>in</strong> North America. It is a<br />
large predatory fish, which can probably cause negative impacts on native fish <strong>species</strong>.<br />
The lake trout is a favourite <strong>in</strong> recreational fish<strong>in</strong>g. Stock<strong>in</strong>g alp<strong>in</strong>e lakes with this<br />
Appendix 2 <strong>species</strong> is permitted. This will have an effect on amphibians shar<strong>in</strong>g the<br />
habitat.<br />
The frequent stock<strong>in</strong>g of water bodies with <strong>alien</strong> fish <strong>species</strong>, and also native fish<br />
<strong>species</strong> from abroad, for sport and commercial fish<strong>in</strong>g <strong>in</strong>creases the possibility of<br />
<strong>in</strong>troduc<strong>in</strong>g diseases (see for example the nematode Anguillicola crassus <strong>in</strong> the Nemathelm<strong>in</strong>thes<br />
section <strong>in</strong> the chapter on ‘other selected <strong>in</strong>vertbrate groups’), <strong>in</strong> addition<br />
to the issue of the potential establishment of the <strong>alien</strong> fish <strong>species</strong> themselves. Environmental<br />
risk studies are recommended before any fish <strong>in</strong>troductions to <strong>in</strong>vestigate<br />
potential threats. Once the decision to <strong>in</strong>troduce a <strong>species</strong> is taken, the material may be<br />
imported as eggs and some type of quarant<strong>in</strong>e measures should be adopted for imported<br />
material prior to its release <strong>in</strong>to natural waters.<br />
The 15 <strong>in</strong>troduced <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> belong to five families (Fig. 1.1) (as expla<strong>in</strong>ed<br />
above, not all of the <strong>species</strong> def<strong>in</strong>itely have established populations, but they<br />
are <strong>in</strong>cluded here, s<strong>in</strong>ce they are listed <strong>in</strong> Appendixes 2 and 3). Two families are<br />
naturally restricted to North America, thus they are new to <strong>Switzerland</strong>. The Cypr<strong>in</strong>idae<br />
are the most <strong>species</strong>-rich family <strong>in</strong> the world with about 2,000 <strong>species</strong>, so it is not<br />
surpris<strong>in</strong>g that the highest number of native as well as <strong>in</strong>troduced <strong>species</strong> belong to this<br />
family. It is also <strong>in</strong>terest<strong>in</strong>g to note that eleven families with only one <strong>species</strong> occur <strong>in</strong><br />
<strong>Switzerland</strong>, although several had more members before some became ext<strong>in</strong>ct. Today<br />
50 % of the salmonids <strong>in</strong> <strong>Switzerland</strong> are <strong>in</strong>troduced (three <strong>species</strong>).<br />
The high number of <strong>in</strong>troduced salmonids reflects their popularity as game fish and for<br />
aquaculture. On the other hand, as Table 2.6 shows by compar<strong>in</strong>g the total number of<br />
<strong>species</strong> per family and the number of <strong>species</strong> <strong>in</strong>troduced to <strong>Switzerland</strong>, the number of<br />
<strong>species</strong> is a stochastic phenomenon – the smaller families produced a higher percentage<br />
of <strong>in</strong>vaders.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 60<br />
Fig. 2.1 > Numbers of native and <strong>in</strong>troduced fish <strong>species</strong> <strong>in</strong> different families <strong>in</strong> <strong>Switzerland</strong>.<br />
Salmonidae<br />
Thymallidae<br />
Coregonidae<br />
Esocidae<br />
Cottidae<br />
Percidae<br />
Gobiidae<br />
Centrarchidae<br />
Blenniidae<br />
Gasterosteidae<br />
Gadidae<br />
Siluridae<br />
Ictaluridae<br />
Cypr<strong>in</strong>idae<br />
Cobitidae<br />
Balitoridae<br />
Clupeidae<br />
Anguillidae<br />
0 5 10 15 20 25 30 35<br />
No. <strong>species</strong><br />
Tab. 2.6 > Total number of members of five fish families worldwide and the number of <strong>species</strong><br />
<strong>in</strong> these families <strong>in</strong>troduced <strong>in</strong>to <strong>Switzerland</strong>.<br />
Total no. <strong>species</strong> per<br />
family<br />
No. <strong>species</strong> <strong>in</strong>troduced to<br />
<strong>Switzerland</strong><br />
Native<br />
Introduced<br />
Percentage of <strong>in</strong>troduced to total<br />
number [%]<br />
Cypr<strong>in</strong>idae 2000 8 0.4<br />
Ictaluridae 35 1 2.9<br />
Centrarchidae 30 2 6.7<br />
Percidae 159 1 0.6<br />
Salmonidae 66 3 4.5<br />
The lack of certa<strong>in</strong>ty about which <strong>species</strong> are actually established, and which of these<br />
are <strong>in</strong>vasive, comb<strong>in</strong>ed with the different def<strong>in</strong>itions used <strong>in</strong> national reports, which<br />
leads to anomalies regard<strong>in</strong>g which <strong>species</strong> are listed and how they are categorized,<br />
makes a comparison with other European countries such as Austria and Germany<br />
difficult. However, the situation appears to be very similar, with the exception of<br />
<strong>species</strong> from Central Europe which are native to Germany. As discussed above, many<br />
<strong>species</strong> are of North American and Asian orig<strong>in</strong> and were widely <strong>in</strong>troduced to Europe,<br />
so that they occur also <strong>in</strong> neighbour<strong>in</strong>g countries.<br />
The 15 <strong>in</strong>troduced <strong>species</strong> represent about 25 % of the current fish fauna. This value is<br />
fairly high compared with the figures for the other groups of vertebrates, <strong>in</strong> which<br />
<strong>in</strong>troduced <strong>species</strong> form less than 10 % of the Swiss fauna. This reflects the economic<br />
importance of fish <strong>species</strong>, but also <strong>in</strong>dicates the potential threat to native biodiversity.
2 > Vertebrates – Vertebrata 61<br />
The orig<strong>in</strong> of the 15 <strong>species</strong> is obviously correlated with the climate, s<strong>in</strong>ce all <strong>species</strong><br />
are from temperate climates <strong>in</strong> the Northern Hemisphere, with six each from North<br />
America and Asia, and three from other parts of Europe. Mikschi (2002) lists two<br />
cichlid <strong>species</strong> for thermal waters <strong>in</strong> Austria, and these tropical <strong>species</strong> can only survive<br />
<strong>in</strong> these warm waters. Three other <strong>species</strong> from the Neotropics (Poeciliidae)<br />
became ext<strong>in</strong>ct after their release <strong>in</strong> the same waters.<br />
The exact pathways for fish <strong>in</strong>troductions are often not known and <strong>in</strong> many cases<br />
<strong>in</strong>troductions have been carried out with more than one motive. Tak<strong>in</strong>g the most likely<br />
pathways for each <strong>species</strong>, those released for commercial and sport fish<strong>in</strong>g and aquaculture<br />
will amount to 11 of the 15 <strong>species</strong>. Two <strong>species</strong> are ornamental releases and<br />
escapes, and one was <strong>in</strong>troduced as a control measure for unwanted vegetation. Interest<strong>in</strong>gly,<br />
only one <strong>species</strong> arrived accidentally. However, its arrival is also connected<br />
with aquacultural practices, s<strong>in</strong>ce it was a contam<strong>in</strong>ant <strong>in</strong> grass carp shipments. In<br />
conclusion, the potential threats to native biodiversity are primarily due to fish<strong>in</strong>g<br />
activities. Fish releases need to be considered carefully, their threats evaluated, and<br />
specimens quarant<strong>in</strong>ed to prevent spread of diseases. The number of <strong>in</strong>troductions on a<br />
global scale has dramatically decreased s<strong>in</strong>ce the 1960s, when they peaked, partially<br />
because grow<strong>in</strong>g awareness of possible negative consequences has led to legislation,<br />
but also because of a saturation effect as some <strong>species</strong> had been <strong>in</strong>troduced to all<br />
suitable recipient areas (Welcomme, 1988).<br />
The demonstrated impacts of the 15 <strong>species</strong> are detailed <strong>in</strong> Table 2.7 and <strong>in</strong> the accounts<br />
for <strong>in</strong>dividual <strong>species</strong>, above. They encompass the entire range of effects measured,<br />
i.e. predatory and graz<strong>in</strong>g pressure, competition with native <strong>species</strong>, changes <strong>in</strong><br />
water quality, community and food-web changes, disease vector<strong>in</strong>g and hybridization.<br />
All potentially harmful <strong>species</strong> are regulated by Swiss law and are listed <strong>in</strong> Appendix 2<br />
or 3. This is a very good basis for management of <strong>alien</strong> fish <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>,<br />
although some of the Appendix 2 <strong>species</strong> which harm native biodiversity can be stocked<br />
<strong>in</strong> alp<strong>in</strong>e lakes without authorization. This is especially worrisome <strong>in</strong> the case of<br />
predatory fish released <strong>in</strong>to previously fishless lakes, where they can damage the<br />
amphibian populations.<br />
Therefore it is recommended that the <strong>species</strong> of Appendix 2 are re-addressed and<br />
stricter regulations are provided for those <strong>species</strong>.<br />
Another po<strong>in</strong>t of concern are <strong>species</strong> native to <strong>Switzerland</strong> released outside their native<br />
range, as noted <strong>in</strong> the sections of this chapter on reptiles and amphibians. The roach<br />
(Rutilus rutilus (L.)), for example, is native to the northern side of the Alps but was<br />
released <strong>in</strong> the Tic<strong>in</strong>o, where it competes with the native fish fauna.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 62<br />
Tab. 2.7 > Alien fish <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>.<br />
Scientific name Family Orig<strong>in</strong> Year Pathway Impact Note<br />
Aristichthys nobilis Richardson Cypr<strong>in</strong>idae Ch<strong>in</strong>a ? Released for fish<strong>in</strong>g Changes of community and habitat? Appendix 3<br />
<strong>species</strong><br />
Aspius aspius (L.) Cypr<strong>in</strong>idae Central and 1994 Released for fish<strong>in</strong>g <strong>in</strong> Harmless Not <strong>in</strong> the<br />
eastern Europe Germany Appendixes<br />
Migration through new<br />
canal systems<br />
Carassius auratus (L.) Cypr<strong>in</strong>idae Central and ? Released and escaped; Competition with native fish Appendix 2<br />
eastern Asia imported as ornamental <strong>species</strong>? <strong>species</strong><br />
fish Community changes by <strong>in</strong>creas<strong>in</strong>g<br />
water turbidity?<br />
Carassius carassius (L.) Cypr<strong>in</strong>idae Europe ? Released for fish<strong>in</strong>g Competition with native fish Appendix 2<br />
Bait fish <strong>species</strong>? <strong>species</strong><br />
Community changes by <strong>in</strong>creas<strong>in</strong>g<br />
water turbidity?<br />
Carassius gibelio (Bloch) Cypr<strong>in</strong>idae Probably Asia ? Released for fish<strong>in</strong>g Competition with native fish Appendix 2<br />
<strong>species</strong>? <strong>species</strong><br />
Community changes by <strong>in</strong>creas<strong>in</strong>g<br />
water turbidity?<br />
Ctenopharyngodon idella (Cuvier Cypr<strong>in</strong>idae Ch<strong>in</strong>a ? Aquaculture Can change the ecosystem by Appendix 3<br />
& Valenciensis) Released to control remov<strong>in</strong>g aquatic plants <strong>species</strong><br />
aquatic vegetation<br />
Hypophthalmichthys molitrix Cypr<strong>in</strong>idae Ch<strong>in</strong>a 1970 Released for control of Community and food-web changes Appendix 3<br />
(Valenciensis) phytoplankton by feed<strong>in</strong>g on phytoplankton <strong>species</strong><br />
Pseudorasbora parva (Temm<strong>in</strong>ck Cypr<strong>in</strong>idae East Asia 1990 Accidental <strong>in</strong>troductions Community and food-web changes Appendix 3<br />
& Schlegel) with other cypr<strong>in</strong>id imports by selective feed<strong>in</strong>g on zooplankton <strong>species</strong><br />
Chances <strong>in</strong> water chemistry<br />
No commercial value<br />
Ameiurus melas and A. nebulosus Ictaluridae Central and ? Aquaculture Predator Appendix 3<br />
(Le Sueur) eastern North Aquarium releases Competition with native fish <strong>species</strong> <strong>species</strong><br />
America Little commercial value<br />
Lepomis gibbosus (L.) Centrarchidae Eastern North ? Ornamental reasons Predator of small <strong>in</strong>vertebrates and Appendix 3<br />
America Released for fish<strong>in</strong>g vertebrates <strong>species</strong><br />
Micropterus salmoides (Lacé- Centrarchidae Central and ? Released for fish<strong>in</strong>g Decl<strong>in</strong>e of native fish <strong>species</strong> Appendix 3<br />
pède) eastern North <strong>species</strong><br />
America<br />
Sander lucioperca (L.) Percidae Central and ? Released for fish<strong>in</strong>g Ferocious predator Appendix 2<br />
eastern Europe <strong>species</strong><br />
Oncorhynchus mykiss Walbaum Salmonidae North America 1887 Released for fish<strong>in</strong>g Predator of native fish and am- Appendix 2<br />
Stock from Aquaculture phibians <strong>species</strong><br />
Germany Competition with native salmonids<br />
Salvel<strong>in</strong>us font<strong>in</strong>alis (Mitchill) Salmonidae Eastern parts of 1883 Released for fish<strong>in</strong>g Competition with native salmonids Appendix 2<br />
North America Aquaculture <strong>species</strong><br />
Stock from<br />
Germany<br />
Salvel<strong>in</strong>us namaycush (Walbaum) Salmonidae North America 1888 Released for fish<strong>in</strong>g Predator of native fish Appendix 2<br />
Competition with native salmonids <strong>species</strong>
2 > Vertebrates – Vertebrata 63<br />
References<br />
Andreotti, A., Baccetti, N., Perfetti, A., Besa, M., Genovesi, P. and V.<br />
Guberti (2001) Mammiferi e ucelli esotici <strong>in</strong> Italia: analisi del fenomeno,<br />
impatto sulla biodiversità e l<strong>in</strong>ee guida gestionali. Quaderni di<br />
Conservazione della Natura No 2, M<strong>in</strong>. Ambiente – Ist. Naz. Fauna<br />
Selvatica, 189 pp.<br />
Arnold, A. (1990) E<strong>in</strong>gebürgerte Fischarten. Die Neue Brehm-Bücherei. A.<br />
Ziemsen Verlag, Wittenberg Lutherstadt, 144 pp.<br />
Arntzen, J.W. (2001) Genetic variation <strong>in</strong> the Italian crested newt, Triturus<br />
carnifex, and the orig<strong>in</strong> of a non-native population north of the Alps.<br />
Biodiversity and Conservation 10, 971–987.<br />
Arntzen, J.W. and R.S. Thorpe (1999) Italian crested newts (Triturus<br />
carnifex) <strong>in</strong> the Bas<strong>in</strong> of Geneva: distribution and genetic <strong>in</strong>teractions with<br />
autochthonous <strong>species</strong>. Herpetologica 55 (4), 423–433.<br />
Barbu, P. (1972) Beiträge zum Studium des Marderhundes, Nyctereutes<br />
procyonoides ussuriensis Matschie, 1907, aus dem Donaudelta.<br />
Säugetierk. Mitt. 20, 375–405.<br />
Bechara, J. A., Moreau, G. and D. Planas (1992) Top-down effects of<br />
brook trout Salvel<strong>in</strong>us font<strong>in</strong>alis <strong>in</strong> a boreal forest stream. Canadian<br />
Journal of Fisheries and Aquatic Sciences 49, 2093–2103.<br />
Bezzel, E. (1996) Neubürger <strong>in</strong> der Vogelwelt Europas: Zoogeographischökologische<br />
Situationsanalyse – Konsequenzen für den Naturschutz. In:<br />
Gebhardt, H., K<strong>in</strong>zelbach, R. and S. Schmidt-Fischer (eds) Gebietsfremde<br />
Tierarten. Auswirkungen auf e<strong>in</strong>heimische Arten, Lebensgeme<strong>in</strong>schaften<br />
und Biotope; Situationsanalyse. Ecomed Verlagsgesellschaft, Landsberg,<br />
pp. 287–296.<br />
Birdlife International (2000) Threatened birds of the world. Lynx Edicions<br />
and BirdLife International, Barcelona, Spa<strong>in</strong> and Cambridge, UK, 864 pp.<br />
Cady, A. and P. Joly (2003) Competition for bask<strong>in</strong>g places between the<br />
endangered European pond turtle (Emys orbicularis galloitalica) and the<br />
red-eared slider (Trachemys scripta elegans). Canadian Journal of<br />
Zoology 81 (8), 1392–1398.<br />
Cady, A. and P. Joly (2004) Impact of the <strong>in</strong>troduction of the red-eared<br />
slider (Trachemys scripta elegans) on survival rates of the European pond<br />
turtle (Emys orbicularis). Biodiversity and Conservation 13 (13), 2511–<br />
2518.<br />
Delany, S.N. (1993) Introduced and escaped geese <strong>in</strong> Brita<strong>in</strong> <strong>in</strong> summer<br />
1991. Brit. Birds 86, 591–599.<br />
Drake C.D. and J.R. Naiman (2000) An evaluation of restoration efforts <strong>in</strong><br />
fishless lakes stocked with exotic trout. Conservation Biology 14 (6),<br />
1807–1820.<br />
Elton, C.S. (1958) The ecology of <strong>in</strong>vasions by animals and plants.<br />
Methuen, London, UK, 181 pp.<br />
Eisfeld, D. and U. Fischer (1996) Das Management e<strong>in</strong>gebürgerter<br />
Huftierarten – widerstreitende Interessen, fehlende Konzepte. In:<br />
Gebhardt, H., K<strong>in</strong>zelbach, R. and S. Schmidt-Fischer (eds) Gebietsfremde<br />
Tierarten. Auswirkungen auf e<strong>in</strong>heimische Arten, Lebensgeme<strong>in</strong>schaften<br />
und Biotope; Situationsanalyse. Ecomed Verlagsgesellschaft, Landsberg,<br />
pp. 287–296.<br />
Englisch, H. (2002) Säugetiere (Mammalia). In: Essl, F. and W. Rabitsch<br />
(eds) Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, pp. 214–221.<br />
Flux, J.E.C. (1994) World distribution. In: Thompson, H.V. and C.M. K<strong>in</strong>g<br />
(eds) The European rabbit – the history of a successful colonizer, pp. 8–<br />
21.<br />
Freye, H.-A. (1978) Castor fiber L<strong>in</strong>naeus, 1758 – Europäischer Biber. In:<br />
Niethammer, J. and F. Krapp (eds) Handbuch der Säugetiere Europas.<br />
Band 1. Aula Verlag, Wiebelsheim, pp. 184–200.<br />
Gebhardt, H. (1996) Ecological and economical consequences of<br />
<strong>in</strong>troductions of exotic wildlife (birds and mammals) <strong>in</strong> Germany. Wildlife<br />
Biology 2, 205–211.<br />
Geiger, A. and M. Waitzmann (1996) Überlebensfähigkeit allochthoner<br />
Amphibien und Reptilien <strong>in</strong> Deutschland – Konsequenzen für den<br />
Artenschutz. In: Gebhardt, H., K<strong>in</strong>zelbach, R. and S. Schmidt-Fischer (eds)<br />
Gebietsfremde Tierarten. Auswirkungen auf e<strong>in</strong>heimische Arten,<br />
Lebensgeme<strong>in</strong>schaften und Biotope; Situationsanalyse. Ecomed<br />
Verlagsgesellschaft, Landsberg, pp. 227–240.<br />
Geiter, O., Homma, S. and R. K<strong>in</strong>zelbach (2002) Bestandsaufnahme und<br />
Bewertung von Neozoen <strong>in</strong> Deutschland. Umweltbundesamt, 308 pp.<br />
Gibb, J.A. and J.E.C. Flux (1973) The mammals. In: Williams, G.R. (ed)<br />
The natural history of New Zealand. A.H. and A.W. Reed, Welll<strong>in</strong>gton, pp.<br />
334–371.<br />
Glutz von Blotzheim, U.N. (ed) (1994) Handbuch der Vögel Mitteleuropas,<br />
Band 9, Columbiformes – Piciformes. Aula-Verlag, Wiebelsheim,<br />
Germany, 1148 pp.<br />
Green, A.J. and B. Hughes (1996) Action plan for the white-headed duck<br />
oxyura leucocephala. In: Heredia, B., Rose, L. and M. Pa<strong>in</strong>ter (eds)<br />
Globally threatened birds <strong>in</strong> Europe. Council of Europe Publish<strong>in</strong>g,<br />
Strasbourg, pp. 119–146.<br />
Gruschwitz, M., Sigrid, L., Mebert, K. and V. Lanka (1999) Natrix<br />
tessellata (Laurenti, 1768) – Würfelnatter. In: Böhme, W. (ed) Handbuch<br />
der Reptilien und Amphibien Europas, Band 3 II A: Schlangen II,<br />
Serpentes II: Colubridae 2 (Boig<strong>in</strong>ae, Natric<strong>in</strong>ae). Aula-Verlag,<br />
Wiebelsheim, pp. 482–815.<br />
Grzimek, B. (1975) Grzimek’s Tierleben, Band 10–13.<br />
Hausser, J. (1995) Säugetiere der Schweiz. Birkhäuser Verlag, Basel,<br />
Boston, Berl<strong>in</strong>, 502 pp.<br />
Hofer, U., Monney, J.-C. and G. Dušej (2001) Die Reptilien der Schweiz.<br />
Verbreitung, Lebensräume, Schutz. Birkenhäuser, Basel, Boston, Berl<strong>in</strong>,<br />
202 pp.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 64<br />
Hohmann, U., Voigt, S. and U. Andreas (2002) Racoons take the offensive.<br />
A current assessment. In: Kowarik, I. and U. Starf<strong>in</strong>ger (eds) Biologische<br />
Invasionen. Herausforderung zum Handeln? Neobiota 1, 191–192.<br />
Hughes, B., Criado, J., Delany, S., Gallo-Ursi, U., Green, A.J., Grussu, M.,<br />
Perrenou, C. and J.A. Torres (1999) The status of the North American<br />
ruddy duck Oxyura jamaicensis <strong>in</strong> the Western Palearctic: towards an<br />
action plan for eradication. Report by the Wildlife & Wetlands Trust to the<br />
Council of Europe.<br />
Kaetzke, P., Niedermeier, J. and M. Masseti (2003) Oryctolagus cuniculus<br />
(L<strong>in</strong>naeus, 1758) – Europäisches Wildkan<strong>in</strong>chen. In: Niethammer, J. and<br />
F. Krapp (eds) Handbuch der Säugetiere Europas. Band 3/II. Aula Verlag,<br />
Wiebelsheim, pp. 187–289.<br />
Kauhala, K. (1996) Introduced carnivores <strong>in</strong> Europe with special reference<br />
to central and northern Europe. Wildlife Biology 2, 197–204.<br />
Keller, V. and N. Zb<strong>in</strong>den (2001) Die <strong>Schweizer</strong> Vogelwelt an der<br />
Jahrhundertwende. Avifauna Report Sempach 1, 64 pp.<br />
Kestenholz, M. and L. Heer (2001) Absichtlich und unabsichtlich<br />
ausgesetzte Vogelarten <strong>in</strong> der Schweiz: Situationsanalyse und<br />
Massnahmenplan. <strong>Schweizer</strong>ische Vogelwarte Sempach und <strong>Schweizer</strong><br />
Vogelschutz SVS – BirdLife Schweiz, Sempach und Zürich, 44 pp.<br />
Krapp, F. (1978a) Tamias striatus (L<strong>in</strong>naeus, 1758) – Chipmunk. In:<br />
Niethammer, J. and F. Krapp (eds) Handbuch der Säugetiere Europas.<br />
Band 1. Aula Verlag, Wiebelsheim, p. 115.<br />
Krapp, F. (1978b) Tamias sibiricus (L<strong>in</strong>naeus, 1758) – Burunduk. In:<br />
Niethammer, J. and F. Krapp (eds) Handbuch der Säugetiere Europas.<br />
Band 1. Aula Verlag, Wiebelsheim, pp. 116–121.<br />
Krapp, F. and J. Niethammer (1986) Cervus nippon Temm<strong>in</strong>ck, 1836 –<br />
Sikahirsch. In: Niethammer, J. and F. Krapp (eds) Handbuch der<br />
Säugetiere Europas. Band 2/II. Aula Verlag, Wiesbaden, 159–172.<br />
Kurt, F. (1988) Sikahirsche (Untergattung Sika). In: Grzimeks<br />
Enzyklopädie Säugetiere. K<strong>in</strong>dler Verlag GmbH, München, pp. 174–175.<br />
LaBastille, A. (1984) Drastic decl<strong>in</strong>e <strong>in</strong> Guatemala’s giant pied-billed<br />
grebe population. Environmental Conservation 11, 346–348.<br />
Ladiges, W. and D. Vogt (1979) Die Süsswasserfische Europas. Paul<br />
Parey, Hamburg und Berl<strong>in</strong>, 299 pp.<br />
Lehtonen, H. (2002) Alien freshwater fishes of Europe. In: Leppaekoski,<br />
E., Gollasch, S. and S. Olen<strong>in</strong> (eds) <strong>Invasive</strong> aquatic <strong>species</strong> <strong>in</strong> Europe,<br />
Kluwer, pp. 153–161.<br />
Long, J.L. (2003) Introduced mammals of the world. Their history,<br />
distribution and <strong>in</strong>fluence. CABI, Wall<strong>in</strong>gford, UK, 589 pp.<br />
Madsen, J., Cracknell, G. and T. Fox (1999) Goose populations of the<br />
Western Palearctic. Wetlands International Publication, No. 48, 343 pp.<br />
Mahan, K. (2002) The effect of an exotic <strong>species</strong> (Oncorhynchus mykiss),<br />
on an endemic population <strong>in</strong> an isolated caldera lake. Journal of<br />
Atawapaskat Research 1, 8–15.<br />
Mikschi, E. (2002) Fische (Pisces). In: Essl, F. and W. Rabitsch (eds)<br />
Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, 197–204.<br />
Müller, R. (1995) Besatzversuche mit ostasiatischen pflanzenfressenden<br />
Fischarten <strong>in</strong> der Schweiz. BUWAL, Schriftenreihe Umwelt Nr. 236, 75 pp.<br />
Popl<strong>in</strong>, F. (1979) Orig<strong>in</strong>e du mouflon de Corse dans uns nouvelle<br />
perspective paléontologique: par marronage. Ann. Génét. Sél. Anim. 11,<br />
133–143.<br />
Randi, E., Pierpaoli, M., Beaumont, M., Ragni, B. and A. Sforzi (2001)<br />
Genetic identification of wild and domestic cats (Felis silvestris) and their<br />
hybrids us<strong>in</strong>g Bayesian cluster<strong>in</strong>g methods. Molecular Biology and<br />
Evolution 18 (9) 1679–1693.<br />
Re<strong>in</strong>hardt, F., Herle, M. Bastiansen, F. and B. Streit (2003) Economic<br />
impact of the spread of <strong>alien</strong> <strong>species</strong> <strong>in</strong> Germany. Federal Environmental<br />
Agency of Germany, 190 pp.<br />
Riegel , J., Lafonta<strong>in</strong>e, R.-M., Pasteels, J. and P. Devillers (2001) Potential<br />
<strong>in</strong>fluence of the Siberian chipmunk Tamias sibiricus (Laxmann) on the<br />
regression of the bird fauna of the Forêt de Soignes, Brussels. Cahiers<br />
d'Éthologie.<br />
Schmid, H., Luder, R., Naef-Daenzer, B., Graf, R. and N. Zb<strong>in</strong>den (1998)<br />
<strong>Schweizer</strong> Brutvogelatlas. Verbreitung der Brutvögel <strong>in</strong> der Schweiz und<br />
im Fürstentum Liechtenste<strong>in</strong> 1993–1996. <strong>Schweizer</strong>ische Vogelwarte,<br />
Sempach.<br />
Schuster, A. (2002) Vögel (Aves). In: Essl, F. and W. Rabitsch (eds)<br />
Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, pp. 209–214.<br />
Schuster, A. and W. Rabitsch (2002) Lurche und Kriechtiere (Amphibia &<br />
Reptilia). In: Essl, F. and W. Rabitsch (eds) Neobiota <strong>in</strong> Österreich.<br />
Umweltbundesamt, Wien, pp. 205–209.<br />
Southern, H.N. (1964) The handbook of British mammals. Oxford, 465 pp.<br />
Weidema, I.R. (ed) (2000) Introduced <strong>species</strong> <strong>in</strong> the Nordic countries.<br />
Nordic Council of M<strong>in</strong>isters, Copenhagen, 242 pp.<br />
Welch, D., Carss, D.N., Gornall, J., Manchester, S.J., Marquiss, M.,<br />
Preston, C.D., Telfer, M.G., Arnold, H. and J. Holbrook (2001) An audit of<br />
<strong>alien</strong> <strong>species</strong> <strong>in</strong> Scotland. Scottish Natural Heritage Review No 139, 225<br />
pp.<br />
Welcomme, R.L. (1988) International <strong>in</strong>troductions of <strong>in</strong>land aquatic<br />
<strong>species</strong>. FAO Fisheries Technical Paper 294, 318 pp.<br />
Williamson, M. (1996) Biological <strong>in</strong>vasions. Chapman & Hall, London,<br />
We<strong>in</strong>haim, New York, Tokyo, Melbourne, Madras, 244 pp.<br />
Woods, M., McDonald, R. and S. Harris (2001) Predation of wildlife by<br />
domestic cats Felis catus <strong>in</strong> Great Brita<strong>in</strong>. Mammal Review 33, 174–188.<br />
Zaugg, B., Stucki, P., Pedroli, J.-C. and A. Kirchhofer (2003) Pisces, Atlas.<br />
Fauna Helvetica 7. Centre Suisse de Cartographie de la Faune, 233 pp.<br />
Zeisset, I. and T.J.C. Beebee (2003) Population genetics of a successful<br />
<strong>in</strong>vader: the marsh frog Rana ridibunda <strong>in</strong> Brita<strong>in</strong>. Molecular Ecology 12,<br />
639–646.
3 > Crustaceans – Crustacea 65<br />
3 > Crustaceans – Crustacea<br />
Prepared by Rüdiger Wittenberg<br />
It is not possible to draw up a comprehensive list of established <strong>alien</strong> crustaceans <strong>in</strong><br />
<strong>Switzerland</strong> ow<strong>in</strong>g to gaps <strong>in</strong> knowledge for some groups and some regions. Another<br />
complicat<strong>in</strong>g factor is the rapid changes <strong>in</strong> <strong>species</strong> composition <strong>in</strong> the large rivers.<br />
Some new <strong>in</strong>vaders are show<strong>in</strong>g an explosive expansion <strong>in</strong> their ranges and densities.<br />
Despite these shortcom<strong>in</strong>gs, a prelim<strong>in</strong>ary list has been compiled us<strong>in</strong>g available<br />
<strong>in</strong>formation. As with all lists of <strong>alien</strong> and <strong>in</strong>vasive <strong>alien</strong> <strong>species</strong>, it is crucial to update<br />
and add new <strong>in</strong>formation about distribution, impacts and management options as it<br />
becomes available. However, this list is believed to be a good basis for future expansion,<br />
s<strong>in</strong>ce a comparison with neighbour<strong>in</strong>g Austria shows similarities <strong>in</strong> <strong>species</strong> and<br />
<strong>species</strong> numbers.<br />
Table 3.1 summarizes available <strong>in</strong>formation on 17 established <strong>alien</strong> crustacean <strong>species</strong><br />
<strong>in</strong> <strong>Switzerland</strong>. Six of them are considered to be harmful to the environment. More<br />
<strong>in</strong>formation on these <strong>species</strong> is presented <strong>in</strong> the Fact Sheets at the end of this chapter.<br />
There is little <strong>in</strong>formation available on Daphnia parvula Fordyce (Cladocera) and<br />
Atyaephyra desmaresti (Millet) (Atyidae) and their <strong>in</strong>vasions might be of little significance<br />
for native biodiversity.<br />
The copepod Cyclops vic<strong>in</strong>us Uljan<strong>in</strong> is thought to have impacted native <strong>species</strong> by its<br />
predatory behaviour.<br />
The amphipod Corophium curvisp<strong>in</strong>um (Sars) (Corophiidae) (see Fact Sheet) is a<br />
considerable threat to native ecosystems. This <strong>species</strong> is an ecosystem eng<strong>in</strong>eer and<br />
occurs at fairly high densities. In the Rh<strong>in</strong>e between Basel and Bodensee it was the<br />
<strong>species</strong> with the third highest number of specimens per square metre, i.e. about 9,200<br />
<strong>in</strong>dividuals (Rey and Ortlepp, 2002).<br />
Six members of the family Gammaridae are established <strong>alien</strong> <strong>species</strong> <strong>in</strong> Swiss waters.<br />
Four of these are of Ponto–Caspian orig<strong>in</strong>, while one <strong>in</strong>vaded Central Europe from<br />
south-western Europe and one <strong>species</strong> came from North America. Of the three <strong>species</strong><br />
Ech<strong>in</strong>ogammarus ischnus (Behn<strong>in</strong>g), E. trichiatus (Martynov) and E. berilloni Catta<br />
only the first-named is known to have a negative impact on the environment (see Table<br />
3.1), while <strong>in</strong>sufficient is currently known about the other <strong>species</strong> to judge. The American<br />
<strong>species</strong> Gammarus tigr<strong>in</strong>us Sexton may change the food web after <strong>in</strong>vasion. Two<br />
Dikerogammarus <strong>species</strong> have <strong>in</strong>vaded <strong>Switzerland</strong>, D. haemobaphes (Eichwald) and
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 66<br />
D. villosus (Soy<strong>in</strong>ski). The latter <strong>species</strong> is the largerst of the <strong>in</strong>vaders and has considerable<br />
impact on the ecosystem (see Fact Sheet).<br />
Introduced freshwater crayfish <strong>species</strong>, compris<strong>in</strong>g four <strong>species</strong> <strong>in</strong> two families,<br />
Astacidae and Cambaridae, represent one of the greatest threats to Swiss biodiversity.<br />
Astacus leptodactylus Eschscholtz (see Fact Sheet) <strong>in</strong>troduced from parts of southeastern<br />
Europe and south-western Asia is of least concern, although it can potentially<br />
compete with native crayfish. The other three <strong>species</strong> (Pacifastacus leniusculus (Dana),<br />
Orconectes limosus (Raf<strong>in</strong>esque) and Procambarus clarkii Girard) (see Fact Sheets)<br />
are of North American orig<strong>in</strong>. They are not susceptible to the crayfish plague (see the<br />
chapter on Fungi), but they are vectors of the disease, carry<strong>in</strong>g it <strong>in</strong>to European crayfish<br />
(Astacus astacus (L.)) populations. The severity of the disease <strong>in</strong> these populations<br />
raises grave concerns about the survival of the native crayfish <strong>species</strong>.<br />
The three Isopoda <strong>species</strong> (see Table 3.1) may not pose a threat to Swiss freshwater<br />
ecosystems, despite the enormous densities of the t<strong>in</strong>y Ponto–Caspian <strong>in</strong>vader Jaera<br />
istri Veuille.<br />
With respect to the taxonomic composition of the established <strong>species</strong>, two groups are<br />
dom<strong>in</strong>ant, i.e. the amphipods with seven <strong>species</strong> and the crayfish with four <strong>species</strong>. The<br />
amphipods are a successful group of about 6,000 <strong>species</strong> (Pöckl, 2002). They often<br />
dom<strong>in</strong>ate ecosystems <strong>in</strong> numbers of <strong>in</strong>dividuals as well as <strong>in</strong> biomass ow<strong>in</strong>g to their<br />
high fecundity under optimal conditions. Their omnivorous behaviour renders them<br />
adaptive to chang<strong>in</strong>g <strong>species</strong> compositions. The rapid <strong>in</strong>vasions and build up of tremendous<br />
densities of <strong>alien</strong> amphipods are legendary. The <strong>in</strong>teractions of native and<br />
rapidly <strong>in</strong>vad<strong>in</strong>g <strong>alien</strong> <strong>species</strong> are very complex and difficult to understand. The fluctuations<br />
<strong>in</strong> densities are considerable and sometimes new <strong>in</strong>vaders replace earlier<br />
<strong>in</strong>vaders. Haas et al. (2002) give an overview of changes <strong>in</strong> abundance of <strong>alien</strong> <strong>species</strong><br />
<strong>in</strong> the Rh<strong>in</strong>e. The crayfish, as the second dom<strong>in</strong>ant group, belongs to the Decapoda,<br />
whose major centre of distribution is <strong>in</strong> North America and there are only a few <strong>species</strong><br />
<strong>in</strong> Europe. The economic importance of crayfish is, of course, the major <strong>in</strong>centive for<br />
<strong>in</strong>troduc<strong>in</strong>g <strong>alien</strong> crayfish <strong>in</strong>to Europe.<br />
A comparison of <strong>alien</strong> crustaceans established <strong>in</strong> <strong>Switzerland</strong> and <strong>in</strong> neighbour<strong>in</strong>g<br />
countries is not without flaws, because of gaps <strong>in</strong> knowledge. However, the 17 <strong>species</strong><br />
listed here compare quite well with the 19 <strong>species</strong> recorded for Austria (Essl and<br />
Rabitsch, 2002). Geiter et al. (2002) list 26 <strong>species</strong> for Germany, but this <strong>in</strong>cludes<br />
<strong>species</strong> liv<strong>in</strong>g <strong>in</strong> mar<strong>in</strong>e and brackish environments.<br />
The orig<strong>in</strong> of the <strong>alien</strong> crustaceans established <strong>in</strong> <strong>Switzerland</strong> is equally divided between<br />
North America (<strong>in</strong>clud<strong>in</strong>g three crayfish <strong>species</strong>), the Ponto–Caspian region<br />
(five of the seven amphipods) and Mediterranean parts of Europe. The Ponto–Caspian<br />
region is essentially the area of the Black and Caspian Seas and their adjacent rivers.<br />
Many <strong>species</strong> endemic to the Ponto–Caspian region have become established <strong>in</strong> Europe,<br />
the Baltic Sea and, more recently, the Great Lakes <strong>in</strong> North America. This extraord<strong>in</strong>ary<br />
spread has been facilitated by the construction of numerous canals allow<strong>in</strong>g<br />
<strong>species</strong> to disperse by active migration and, to a greater extent, with ship traffic (<strong>in</strong>
3 > Crustaceans – Crustacea 67<br />
ballast tanks and through hull-foul<strong>in</strong>g). Many Ponto–Caspian crustaceans were also<br />
transplanted between water bodies as food for native fish to stimulate fish production<br />
with<strong>in</strong> the former USSR.<br />
Although the exact time and place of the first <strong>in</strong>troductions of many <strong>species</strong> are not<br />
documented, the data available suggest that many of them have <strong>in</strong>vaded <strong>Switzerland</strong><br />
fairly recently. Thus, further spread and <strong>in</strong>crease <strong>in</strong> <strong>species</strong> densities <strong>in</strong> the short or<br />
long term are very likely. Moreover, other <strong>species</strong> are expected to arrive via river<br />
systems.<br />
With the exception of the crayfish <strong>species</strong>, almost all the crustacean <strong>in</strong>vasions of<br />
<strong>Switzerland</strong> have been facilitated by the construction of canals and ship traffic. In most<br />
cases it is difficult to prove the exact pathways for the various <strong>species</strong>. Natural migration<br />
of some mobile <strong>species</strong> along rivers and canals is likely to be important over short<br />
distances. However, the rapid expansion of most <strong>species</strong> and some isolated records<br />
<strong>in</strong>dicate spread by ship traffic, either <strong>in</strong> ballast tanks or on hulls. The crayfish <strong>species</strong>,<br />
however, were deliberately <strong>in</strong>troduced as a food resource for human consumption.<br />
Additionally, escapees from aquaculture facilities have founded several populations. A<br />
third pathway is the importation of live crayfish for consumption. Crayfish are a delicacy<br />
and have to be added to boil<strong>in</strong>g water alive. Thus, they are imported alive and<br />
some may have escaped. Moreover, some specimens escape from aquariums and<br />
garden ponds, or are released by their owners, who no longer want to keep them.<br />
The impacts of the <strong>species</strong>, especially the six high-risk <strong>species</strong>, are detailed <strong>in</strong> the Fact<br />
Sheets and <strong>in</strong> Table 3.1. The three North American crayfish <strong>species</strong> are of greatest<br />
concern with regard to the survival of the native crayfish populations, because they act<br />
as a vector of the devastat<strong>in</strong>g crayfish plague. The two amphipod <strong>species</strong> D. villosus<br />
and C. curvisp<strong>in</strong>um alter the <strong>in</strong>vaded habitats by predation, competition and caus<strong>in</strong>g<br />
changes to the substrate. Demonstrat<strong>in</strong>g impacts of <strong>alien</strong> <strong>species</strong> on native biodiversity<br />
and ecosystems is always difficult ow<strong>in</strong>g to the complex <strong>in</strong>teractions. In conclusion,<br />
<strong>species</strong> thought to be harmless may turn out to have some, as yet undetected, negative<br />
impacts. This is especially true for freshwater ecosystems. Many of the <strong>alien</strong> <strong>species</strong><br />
occur <strong>in</strong> enormous abundance. This must have some impact on the ecosystem, s<strong>in</strong>ce<br />
each <strong>in</strong>dividual uses some resources and is a resource for others. Rey and Ortlepp<br />
(2002) found that the biomass <strong>in</strong> the Rh<strong>in</strong>e near Basel was dom<strong>in</strong>ated by <strong>alien</strong> <strong>species</strong><br />
– 97 % of the animal biomass and more than 90 % of the <strong>in</strong>dividuals were of <strong>alien</strong><br />
orig<strong>in</strong>. Thus, the orig<strong>in</strong>al character of the Rh<strong>in</strong>e has vanished; <strong>in</strong> this area it is not the<br />
typical Rh<strong>in</strong>e anymore. Most of the <strong>alien</strong> <strong>species</strong> have <strong>in</strong>vaded this stretch of the Rh<strong>in</strong>e<br />
with<strong>in</strong> the past five years. This suggests that their distribution will further expand and<br />
their dom<strong>in</strong>ance <strong>in</strong> other areas will <strong>in</strong>crease. The dom<strong>in</strong>ance of <strong>alien</strong> <strong>species</strong> <strong>in</strong> European<br />
(and other cont<strong>in</strong>ental) <strong>in</strong>land waters is dramatic.<br />
In most cases, eradication of established problematic <strong>species</strong> is not feasible. Control<br />
has limited success <strong>in</strong> open freshwater systems, such as rivers. Some <strong>species</strong>, <strong>in</strong> particular<br />
the crayfish, have economic value and <strong>in</strong>tensive fish<strong>in</strong>g could reduce their<br />
numbers. However, a reduction <strong>in</strong> the crayfish population might not be a sufficient<br />
goal, given the great threat they pose to native crayfish. One crayfish plague-<strong>in</strong>fested
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 68<br />
<strong>alien</strong> crayfish could be enough to wipe out a healthy population of their European<br />
relatives. Migrat<strong>in</strong>g specimens of <strong>alien</strong> crayfish were observed even <strong>in</strong> low-density<br />
situations, so a reduction might not prevent their spread. Thus, prevention of further<br />
<strong>in</strong>troductions of <strong>species</strong> <strong>in</strong>to and with<strong>in</strong> <strong>Switzerland</strong> should have highest priority to<br />
safeguard native biodiversity and ecosystems. North American crayfish <strong>species</strong> are of<br />
greatest concern. With respect to releases of problematic freshwater <strong>species</strong> (e.g. for<br />
fish<strong>in</strong>g and from aquarium dump<strong>in</strong>g) an awareness-rais<strong>in</strong>g campaign would be crucial<br />
to sensitize the public to the potential problems caused by <strong>alien</strong> <strong>species</strong>. Accidental<br />
arrival of new <strong>species</strong> with ship traffic can only be m<strong>in</strong>imized by effective treatment of<br />
ballast water and ships’ hulls. Crayfish <strong>species</strong> other than those mentioned above must<br />
be regulated too. An Australian <strong>species</strong>, Cherax destructor Clark for example, has been<br />
found <strong>in</strong> <strong>Switzerland</strong> (Stucki and Jean-Richard, 2000) and is be<strong>in</strong>g kept as a pet <strong>in</strong><br />
aquariums and garden ponds ow<strong>in</strong>g to its beautiful appearance. The release of pets is<br />
illegal, but the unfortunate fact is that it is used as a way of gett<strong>in</strong>g rid of unwanted<br />
<strong>in</strong>dividuals, and they are also capable of escap<strong>in</strong>g from ponds.
3 > Crustaceans – Crustacea 69<br />
Tab. 3.1 > Established <strong>alien</strong> crustaceans <strong>in</strong> <strong>Switzerland</strong>.<br />
Scientific name Taxonomic Orig<strong>in</strong> Year Pathway Impact Note<br />
group<br />
Daphnia parvula Fordyce Cladocera America 1972 Ship traffic Change <strong>in</strong> food web? First European record <strong>in</strong> the<br />
Bodensee (Lake Constance)<br />
Cyclops vic<strong>in</strong>us Uljan<strong>in</strong> Copepoda Europe 1954 Fish releases? Predator of native crusta- Found <strong>in</strong> the Bodensee.<br />
ceans<br />
Atyaephyra desmaresti Atyidae Mediterranean ? Expand<strong>in</strong>g along No impact shown Likely to expand further throughout<br />
(Millet) canals, ship Europe<br />
traffic?<br />
Corophium curvisp<strong>in</strong>um Corophiidae Ponto– 1980s Ballast water Change of ecosystem by Rapid colonization of the Rh<strong>in</strong>e<br />
(Sars) Caspian Also migration transferr<strong>in</strong>g hard substrate to<br />
muddy areas.<br />
Reduces available habitat for<br />
hard substrate <strong>species</strong><br />
Ech<strong>in</strong>ogammarus ischnus Gammaridae Ponto– 1990s Ship traffic Modifies substrate sediment. Invader of a large portion of Europe<br />
(Behn<strong>in</strong>g) Caspian Migration Alters energy flow between<br />
pelagic and benthic organisms.<br />
Additional prey for fish.<br />
Excludes compet<strong>in</strong>g <strong>species</strong><br />
Ech<strong>in</strong>ogammarus Gammaridae Ponto– - Ship traffic Not known Likely to reach <strong>Switzerland</strong> <strong>in</strong> the<br />
trichiatus (Martynov) Caspian near future<br />
Ech<strong>in</strong>ogammarus berilloni Gammaridae South-western 20 th century Migration through Not known Probably still <strong>in</strong>vad<strong>in</strong>g Europe<br />
Catta Europe canals<br />
Dikerogammarus Gammaridae Ponto– 1990s Ship traffic Not known Decreas<strong>in</strong>g <strong>in</strong> Europe, because of<br />
haemobaphes (Eichwald) Caspian Migration the <strong>in</strong>vasion of D. villosus<br />
Dikerogammarus villosus Gammaridae Ponto– Late 1990s Ship traffic Predator of <strong>alien</strong> and native Replac<strong>in</strong>g earlier <strong>in</strong>vaders<br />
(Sov<strong>in</strong>ski) Caspian gammarids and other prey<br />
Gammarus tigr<strong>in</strong>us Gammaridae North America 1990s Ship traffic Changes <strong>in</strong> food web? Introduced from North America <strong>in</strong><br />
Sexton Migration ballast water<br />
Astacus leptodactylus Astacidae South-eastern 1980s Release for Competition with native Important competitor for native<br />
Eschscholtz Europe fish<strong>in</strong>g crayfish <strong>species</strong> crayfish <strong>species</strong>, but also vulnerable<br />
to the crayfish plague<br />
Pacifastacus leniusculus Astacidae North America ? Release for Competition with native Very destructive <strong>in</strong>vader, displac<strong>in</strong>g<br />
(Dana) fish<strong>in</strong>g crayfish <strong>species</strong> native crayfish <strong>species</strong><br />
Vector of the crayfish plague<br />
Orconectes limosus Cambaridae North America Before 1976 Release for Competition with native Very destructive <strong>in</strong>vader, displac<strong>in</strong>g<br />
(Raf<strong>in</strong>esque) fish<strong>in</strong>g crayfish <strong>species</strong>. native crayfish <strong>species</strong><br />
Vector of the crayfish plague<br />
Procambarus clarkii Cambaridae Southern ? Release for Competition with native Very destructive <strong>in</strong>vader, displac<strong>in</strong>g<br />
Girard North America fish<strong>in</strong>g crayfish <strong>species</strong>. native crayfish <strong>species</strong><br />
Vector of the crayfish plague<br />
Proasellus coxalis Asellidae South-western Second half Ship traffic Not known Rh<strong>in</strong>e, but not common<br />
(Dollfuss) Europe of 20 th<br />
century<br />
Proasellus meridianus Asellidae South-western Second half Ship traffic Not known Rh<strong>in</strong>e, near Basel, but not common<br />
(Racovitza) Europe of 20 th<br />
century<br />
Jaera istri Veuille Jaeridae Ponto– End of Ship traffic Not known, but very abun- One of the <strong>species</strong> with the highest<br />
Caspian 1990s dant abundance – likely to <strong>in</strong>crease<br />
further
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 70<br />
References<br />
Bernauer, D., Kappus, B. and W. Jansen (1996) Neozoen <strong>in</strong><br />
Kraftwerksproben und Begleituntersuchungen am nördlichen Oberrhe<strong>in</strong>.<br />
In: Gebhardt, H., K<strong>in</strong>zelbach, R. and S. Schmidt-Fischer (eds)<br />
Gebietsfremde Tierarten. Auswirkungen auf e<strong>in</strong>heimische Arten,<br />
Lebensgeme<strong>in</strong>schaften und Biotope; Situationsanalyse. Ecomed<br />
Verlagsgesellschaft, Landsberg, pp. 87–95.<br />
Eder, E. (2002) Krebstiere: Flusskrebse (Crustacea: Decapoda: Astacidae<br />
und Cambaridae). In: Essl, F. and W. Rabitsch (eds) Neobiota <strong>in</strong><br />
Österreich. Umweltbundesamt, Wien, pp. 287–291.<br />
Essl, F. and W. Rabitsch (eds) (2002) Neobiota <strong>in</strong> Österreich. Federal<br />
Environment Agency, 432 pp.<br />
Frutiger, A. and R. Müller (2002) Der Rote Sumpfkrebs im Schübelweiher<br />
(Geme<strong>in</strong>de Küsnacht ZH). Auswertung der Massnahmen 1998–2001 und<br />
Erkenntnisse. EAWAG.<br />
Gaviria, S. (2002) Krebstiere: Wasserflöhe & Ruderfusskrebse (Crustacea:<br />
Cladocera und Copepoda). In: Essl, F. and W. Rabitsch (eds) Neobiota <strong>in</strong><br />
Österreich. Umweltbundesamt, Wien, pp.263–268.<br />
Geiter, O., Homma, S. and R. K<strong>in</strong>zelbach (2002) Bestandsaufnahme und<br />
Bewertung von Neozoen <strong>in</strong> Deutschland. Umweltbundesamt, 308 pp.<br />
Haas, G., Brunke, M. and B. Streit (2002) Fast turnover <strong>in</strong> dom<strong>in</strong>ance of<br />
exotic <strong>species</strong> <strong>in</strong> the Rh<strong>in</strong>e river determ<strong>in</strong>es biodiversity and ecosystem<br />
function: an affair between amphipods and mussles. In: Leppäkoski, E.,<br />
Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic <strong>species</strong> of Europe.<br />
Distribution, impacts and management. Kluwer Academic Publishers,<br />
Dordrecht, pp. 426–432.<br />
Löffler, H. (1983) Aspects of the history and evolution of Alp<strong>in</strong>e lakes <strong>in</strong><br />
Austria. Hydrobiologia 100, 143–152.<br />
Ojaveer, H., Leppäkoski, E., Olen<strong>in</strong>, S. and A. Ricciardi (2002) Ecological<br />
impact of ponto-caspian <strong>in</strong>vaders <strong>in</strong> the Baltic Sea, European <strong>in</strong>land<br />
waters and the Great Lakes: an <strong>in</strong>ter-ecosystem comparison. In:<br />
Leppäkoski, E., Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic <strong>species</strong><br />
of Europe. Distribution, impacts and management. Kluwer Academic<br />
Publishers, Dordrecht, pp. 412–425.<br />
Pöckl, M. (2002) Flohkrebse (Crustacea: Amphipoda: Corophiidae und<br />
Gammaridae). In: Essl, F. and W. Rabitsch (eds) Neobiota <strong>in</strong> Österreich.<br />
Umweltbundesamt, Wien, pp. 273–284.<br />
Pöckl, M. and W. Rabitsch (2002) Ausgewählte aquatische Neozoen<br />
(Cnidaria, Plathelm<strong>in</strong>thes, Kamptozoa, Annelida, Isopoda). In: Essl, F. and<br />
W. Rabitsch (eds) Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, pp.<br />
228–239.<br />
Re<strong>in</strong>hardt, F., Herle, M. Bastiansen, F. and B. Streit (2003) Economic<br />
impact of the spread of <strong>alien</strong> <strong>species</strong> <strong>in</strong> Germany. Federal Environmental<br />
Agency of Germany, 190 pp.<br />
Rey, P. and J. Ortlepp (2002) Koord<strong>in</strong>ierte biologische Untersuchungen<br />
am Hochrhe<strong>in</strong> 2000; Makro<strong>in</strong>vertebraten. BUWAL Schriftenreihe Umwelt<br />
Nr. 345, Gewässerschutz, 98 pp.<br />
Stucki, T. and P. Jean-Richard (2000) Verbreitung der Flusskrebse <strong>in</strong> der<br />
Schweiz. Mitteilungen zur Fischerei Nr. 65, 52 pp.<br />
Tittizer, T. (1996) Vorkommen und Ausbreitung aquatischer Neozoen<br />
(Makrozoobenthos) <strong>in</strong> den Bundeswasserstrassen. In: Gebhardt, H.,<br />
K<strong>in</strong>zelbach, R. and S. Schmidt-Fischer (eds) Gebietsfremde Tierarten.<br />
Auswirkungen auf e<strong>in</strong>heimische Arten, Lebensgeme<strong>in</strong>schaften und<br />
Biotope; Situationsanalyse. Ecomed Verlagsgesellschaft, Landsberg, pp.<br />
49–86.<br />
Tittizer, T., Schöll, F., Bann<strong>in</strong>g, M., Haybach, A. and M. Schleuter (2000)<br />
Aquatische Neozoen im Makrozoobenthos der B<strong>in</strong>nenwasserstrassen<br />
Deutschlands. Lauterbornia 39, 1–72.<br />
Vorburger, C. and G. Ribi (1999) Aggression and competition for shelter<br />
between a native and an <strong>in</strong>troduced crayfish <strong>in</strong> Europe. Freshwater<br />
Biology 42, 111–119.<br />
Welch, D., Carss, D.N., Gornall, J., Manchester, S.J., Marquiss, M.,<br />
Preston, C.D., Telfer, M.G., Arnold, H. and J. Holbrook (2001) An audit of<br />
<strong>alien</strong> <strong>species</strong> <strong>in</strong> Scotland. Scottish Natural Heritage Review No 139, 225<br />
pp.<br />
Westman, K. (2002) Alien crayfish <strong>in</strong> Europe: negative and positive<br />
impacts and <strong>in</strong>teractions with native crayfish. In: Leppäkoski, E.,<br />
Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic <strong>species</strong> of Europe.<br />
Distribution, impacts and management. Kluwer Academic Publishers,<br />
Dordrecht, pp. 76–95.<br />
Wittmann, K.J. (2002) Krebstiere: “Schwebgarnelen” und<br />
Süsswassergarnelen (Crustacea: Mysidacea: Mysidae, Decapoda:<br />
Atyidae). In: Essl, F. and W. Rabitsch (eds) Neobiota <strong>in</strong> Österreich.<br />
Umweltbundesamt, Wien, 269–272.
4 > Insects – Insecta 71<br />
4 > Insects – Insecta<br />
Prepared by Marc Kenis<br />
4.1 Introduction<br />
Insects probably represent the most numerous exotic organisms <strong>in</strong> <strong>Switzerland</strong>. A list<br />
of <strong>in</strong>sects that are considered to be of exotic orig<strong>in</strong> has been drawn up, with the help of<br />
specialists (see Tables 4.1–4.6). It <strong>in</strong>cludes 311 <strong>species</strong> of exotic orig<strong>in</strong> that have<br />
certa<strong>in</strong>ly or probably been <strong>in</strong>troduced <strong>in</strong>to <strong>Switzerland</strong> or neighbour<strong>in</strong>g countries by<br />
human activities. It does not <strong>in</strong>clude <strong>species</strong> that are spread<strong>in</strong>g naturally <strong>in</strong>to <strong>Switzerland</strong>,<br />
unless they were first <strong>in</strong>troduced <strong>in</strong>to neighbour<strong>in</strong>g countries. This list is by no<br />
means complete, but it will be ma<strong>in</strong>ta<strong>in</strong>ed and new records will be added when data<br />
become available. In collaboration with taxonomists for the various <strong>in</strong>sect groups, we<br />
also <strong>in</strong>tend to publish this list <strong>in</strong> peer-reviewed journals, with more details on the<br />
occurrence, distribution and biology of the exotic <strong>species</strong>. The two published lists of<br />
<strong>alien</strong> organisms <strong>in</strong> Austria and Germany (Essl and Rabitsch, 2002; Geiter et al., 2002)<br />
have been of great help <strong>in</strong> the construction of the Swiss list.<br />
Many problems were encountered <strong>in</strong> draw<strong>in</strong>g up the list, the ma<strong>in</strong> ones be<strong>in</strong>g detailed<br />
below.<br />
> There is an obvious lack of <strong>in</strong>formation sources <strong>in</strong> <strong>Switzerland</strong>. Up-to-date, published<br />
checklists are available for a few <strong>in</strong>sect groups only (e.g. Diptera, Orthoptera,<br />
Odonata). The ‘Centre Suisse de Cartographie de la Faune’ (CSCF) is ma<strong>in</strong>ta<strong>in</strong><strong>in</strong>g<br />
<strong>in</strong>complete lists with the <strong>in</strong>put of Swiss taxonomists, and several lists of other important<br />
<strong>in</strong>sect groups are <strong>in</strong> the process of be<strong>in</strong>g completed (e.g. Coleoptera, Lepidoptera,<br />
Aphidoidea, etc.). The CSCF and <strong>in</strong>dividual taxonomists were very cooperative<br />
<strong>in</strong> help<strong>in</strong>g us check and complete the list of exotic <strong>in</strong>sects; nevertheless, it<br />
has been a rather difficult task. For some groups, there is presently little expertise <strong>in</strong><br />
<strong>Switzerland</strong>, and no recent checklist is available (e.g. most primitive orders and<br />
Mallophaga, which are therefore not <strong>in</strong>cluded <strong>in</strong> the list, and also parts of the Hymenoptera,<br />
Heteroptera, etc.). Additionally, it must be noted that <strong>in</strong> the few checklists<br />
published, the orig<strong>in</strong> of the <strong>species</strong> is often not clearly <strong>in</strong>dicated. For future<br />
checklists, we suggest <strong>in</strong>clud<strong>in</strong>g the ‘exotic character’ and the area of orig<strong>in</strong> <strong>in</strong> the<br />
data set.<br />
> Many <strong>alien</strong> <strong>in</strong>sects <strong>in</strong> <strong>Switzerland</strong> orig<strong>in</strong>ate from the Mediterranean region. Some of<br />
these <strong>species</strong>, particularly Hemiptera, were evidently <strong>in</strong>troduced with their host<br />
plants, which are often planted as ornamentals or crops <strong>in</strong> <strong>Switzerland</strong>. However, for<br />
other <strong>species</strong>, it is not clear whether they were <strong>in</strong>troduced by human activities or<br />
arrived naturally (e.g. the Heteroptera Arocatus longiceps and Deraeocoris fla-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 72<br />
vil<strong>in</strong>ea). Some <strong>species</strong> undoubtedly migrated unaided from southern Europe to<br />
<strong>Switzerland</strong>, possibly because of global warm<strong>in</strong>g. This is, for example, the case for<br />
the dragonflies Sympetrum meridionale (Selys) and Crocothemis erythraea (Brullé).<br />
Likewise, many Lepidoptera <strong>species</strong> (e.g. Helicoverpa armigera (Hübner) and<br />
Mythimna unipuncta (Haworth), which used to be observed only occasionally <strong>in</strong><br />
summer <strong>in</strong> southern <strong>Switzerland</strong>, are now overw<strong>in</strong>ter<strong>in</strong>g <strong>in</strong> the country more frequently.<br />
Eastern European <strong>in</strong>sect <strong>species</strong> are also <strong>in</strong>creas<strong>in</strong>gly observed <strong>in</strong> <strong>Switzerland</strong>.<br />
For most of them, it is not clear whether their <strong>in</strong>troduction <strong>in</strong>to <strong>Switzerland</strong><br />
was natural or by human-mediated transport, and whether they are firmly established<br />
or not. In general, <strong>species</strong> whose establishment <strong>in</strong> <strong>Switzerland</strong> seems to be the result<br />
of a natural process were not <strong>in</strong>cluded <strong>in</strong> our list. However, this phenomenon would<br />
merit further studies.<br />
> High numbers of tropical or subtropical <strong>in</strong>sects are found <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> houses,<br />
greenhouses and other conf<strong>in</strong>ed environments. Many of them are important pests of<br />
greenhouse crops (e.g. several thrips and whiteflies) or stored products (e.g. various<br />
beetle and moth <strong>species</strong>). However, it is not always easy to determ<strong>in</strong>e whether these<br />
<strong>in</strong>sects are firmly established, or regularly <strong>in</strong>troduced from abroad with new plant<br />
material. Furthermore, some <strong>species</strong> that were thought to be conf<strong>in</strong>ed to greenhouses<br />
are now found outdoors <strong>in</strong> natural environments. Examples <strong>in</strong>clude, among others,<br />
the scale <strong>in</strong>sects Icerya purchasi, Coccus hesperidum and Dynaspidiotus britannicus<br />
(Kozar et al., 1994).<br />
> Today, many <strong>in</strong>sects, particularly those feed<strong>in</strong>g on stored products and crops and<br />
occurr<strong>in</strong>g as ectoparasites of vertebrates are considered to be of cosmopolitan distribution<br />
and, therefore, it is often difficult to assess their exotic status and some will<br />
f<strong>in</strong>ally be concluded to be cryptogenic <strong>species</strong>. Only <strong>in</strong>sects that are suspected to be<br />
exotic, for any reason (e.g. because they feed only on an exotic plant, or because the<br />
genus is considered exotic) are <strong>in</strong>cluded <strong>in</strong> the list.<br />
<strong>Invasive</strong> <strong>alien</strong> <strong>in</strong>sects are serious threats to agriculture, economies, the environment<br />
and human and animal health worldwide. In some regions, such as North America,<br />
South Africa and many oceanic islands, exotic <strong>in</strong>sect pests are considered to be as<br />
important as native pests, if not more so. Traditionally, problems have been less severe<br />
<strong>in</strong> Central Europe. Only a few exotic <strong>in</strong>sects are known to cause serious damage <strong>in</strong><br />
<strong>Switzerland</strong>, usually as stored product pests or on agricultural crops. However, <strong>in</strong><br />
recent years several pests of economic importance have <strong>in</strong>vaded Europe, stimulat<strong>in</strong>g<br />
more <strong>in</strong>terest <strong>in</strong> the issue of exotic <strong>in</strong>sects. For example, the western corn rootworm,<br />
Diabrotica virgifera ssp. virgifera is seriously threaten<strong>in</strong>g European maize production,<br />
and the horse chestnut leaf m<strong>in</strong>er, Cameraria ohridella, is caus<strong>in</strong>g much public concern<br />
because of its spectacular damage to urban trees <strong>in</strong> Central Europe.<br />
Worldwide, environmental impacts have been studied less for <strong>alien</strong> <strong>in</strong>sects than for<br />
other groups of <strong>in</strong>vasive organisms such as plants or mammals. However, many <strong>in</strong>sects<br />
are known to cause serious environmental damage <strong>in</strong> <strong>in</strong>vaded habitats. Here aga<strong>in</strong>,<br />
most of these cases are from outside Europe. For example, the hemlock woolly adelgid,<br />
Adelges tsugae Annand, is threaten<strong>in</strong>g unique forest ecosystems <strong>in</strong> North America<br />
(Jenk<strong>in</strong>s et al., 1999); the scale <strong>in</strong>sect Orthezia <strong>in</strong>signis Browne was <strong>in</strong> the process of<br />
push<strong>in</strong>g the endemic gumwood tree Commidendrum robustum DC. <strong>in</strong> Sa<strong>in</strong>t Helena to
4 > Insects – Insecta 73<br />
the br<strong>in</strong>k ext<strong>in</strong>ction when a successful biological control programme was implemented<br />
(Fowler, 2004). In Europe, there have been few similar examples. Recently, however,<br />
the Argent<strong>in</strong>e ant L<strong>in</strong>epithema humile has been caus<strong>in</strong>g changes <strong>in</strong> <strong>in</strong>vertebrate and<br />
plant communities <strong>in</strong> the Mediterranean region by predation and displacement of native<br />
<strong>species</strong> (Gómez and Oliveras, 2003). The horse chestnut leaf m<strong>in</strong>er is an urban pest <strong>in</strong><br />
Central Europe but, <strong>in</strong> the Balkans, it is threaten<strong>in</strong>g the few rema<strong>in</strong><strong>in</strong>g <strong>in</strong>digenous<br />
horse chestnut populations. In general, most <strong>alien</strong> <strong>in</strong>sects have not been the targets of<br />
serious environmental impact studies. Their <strong>in</strong>teraction with the native fauna and flora<br />
has been rarely <strong>in</strong>vestigated, particularly if their habitat is of little economic concern.<br />
Environmental impacts may be direct (on a plant for a herbivore, or on a prey for a<br />
predator), but the <strong>in</strong>vader may also have an <strong>in</strong>direct impact on co-occurr<strong>in</strong>g organisms,<br />
e.g. through shar<strong>in</strong>g the same food webs. Among the over 300 recorded exotic <strong>in</strong>sects<br />
<strong>in</strong> <strong>Switzerland</strong>, surely more than one has an important environmental impact with<strong>in</strong> the<br />
newly <strong>in</strong>vaded ecosystem? Investigations on the impact of <strong>in</strong>vasive <strong>species</strong> should not<br />
be restricted to <strong>species</strong> of economic importance, but should also focus on abundant<br />
<strong>species</strong> <strong>in</strong> poorly studied ecosystems, <strong>in</strong> which important environmental impacts may<br />
have been overlooked.<br />
The gaps <strong>in</strong> knowledge of the Swiss <strong>in</strong>sect fauna, as expla<strong>in</strong>ed at the beg<strong>in</strong>n<strong>in</strong>g of this<br />
chapter, precludes any conclusive evaluation of its status, i.e. what percentage of the<br />
Swiss <strong>in</strong>sect fauna is <strong>alien</strong>, the orig<strong>in</strong>s of the <strong>alien</strong> <strong>species</strong>, and the pathways by which<br />
they reached <strong>Switzerland</strong>. However, the fact that many <strong>species</strong> were apparently <strong>in</strong>troduced<br />
with their host plants and food <strong>in</strong>dicates that trade <strong>in</strong> commodities is a major<br />
pathway of <strong>in</strong>sect <strong>in</strong>troduction <strong>in</strong>to <strong>Switzerland</strong>. Furthermore, it is certa<strong>in</strong> that the<br />
majority of the <strong>alien</strong> <strong>in</strong>sect fauna was accidentally <strong>in</strong>troduced. Prevention of entry of<br />
pest <strong>species</strong> is addressed by quarant<strong>in</strong>e regulations and <strong>in</strong>terception; however, <strong>species</strong><br />
of environmental concern are mostly not covered by these measures. Control of <strong>alien</strong><br />
<strong>in</strong>sect populations is also focussed on <strong>species</strong> of economic importance.<br />
4.2 Coleoptera<br />
There is no recent checklist of Coleoptera of <strong>Switzerland</strong>. However, Lucht (1987), <strong>in</strong><br />
his catalogue of Coleoptera of Central Europe, <strong>in</strong>cludes distribution data for northern<br />
<strong>Switzerland</strong>. Furthermore, Claude Besuchet, Muséum d’Histoire Naturelle, Genève, is<br />
currently compil<strong>in</strong>g a checklist for <strong>Switzerland</strong>, and he k<strong>in</strong>dly helped us complete our<br />
list with his unpublished data.<br />
Over 120 beetle <strong>species</strong> are known, or suspected, to be of exotic orig<strong>in</strong> <strong>in</strong> <strong>Switzerland</strong>,<br />
which accounts for about 40 % of the list of <strong>alien</strong> <strong>in</strong>sects. Many of these exotic <strong>species</strong><br />
are domestic pests, often feed<strong>in</strong>g on stored products, construction material, etc. The<br />
ma<strong>in</strong> pests of stored products are found <strong>in</strong> the families Anobiidae, Bostrichidae, Bruchidae,<br />
Cucujidae, Curculionidae, Dermestidae, Mycetophagidae, Nitidulidae, Pt<strong>in</strong>idae,<br />
Silvanidae and Tenebrionidae. Most of them have been transported with their food and<br />
are now found worldwide. Their artificial cosmopolitan distribution renders the identification<br />
of their orig<strong>in</strong> difficult. Some <strong>species</strong>, <strong>in</strong> particular many Dermestidae, may<br />
Advisor:<br />
Claude Besuchet, Muséum<br />
d’Histoire Naturelle, Genève
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 74<br />
actually be native to Central Europe. Thus the <strong>in</strong>clusion of some <strong>species</strong> <strong>in</strong> the list is<br />
questionable.<br />
Some of the stored product pests are of tropical orig<strong>in</strong> and are strictly domestic <strong>in</strong><br />
<strong>Switzerland</strong>, whereas others can also survive and reproduce under outdoor conditions.<br />
Many more <strong>species</strong> than those listed <strong>in</strong> this study are found associated with stored<br />
products <strong>in</strong> <strong>Switzerland</strong>. However, it is not always clear whether these <strong>species</strong> are<br />
established, or regularly <strong>in</strong>troduced with imported goods. Our list <strong>in</strong>cludes only the<br />
<strong>species</strong> thought to be established, although, <strong>in</strong> several cases, the decision on whether to<br />
<strong>in</strong>clude them or not was rather subjective. For example, we did not <strong>in</strong>clude the coffee<br />
bean beetle, Araecerus fasciculatus De Geer, a <strong>species</strong> often found <strong>in</strong> imported coffee<br />
and cocoa beans <strong>in</strong> Swiss food factories, but not clearly established <strong>in</strong> the country.<br />
Hoppe (1981) surveyed stored commodities <strong>in</strong> <strong>Switzerland</strong> and found the most destructive<br />
beetles to be the curculionids Sitophilus granarius and S. orizae, the silvanid<br />
Oryzaephilus sur<strong>in</strong>amensis and the tenebrionids Tribolium castaneum and T. confusum,<br />
all considered to be of exotic orig<strong>in</strong>.<br />
Apart from stored product pests, few exotic beetles are recorded as agricultural pests <strong>in</strong><br />
<strong>Switzerland</strong>. However, two important chrysomelid beetles are worth mention<strong>in</strong>g. The<br />
<strong>in</strong>famous Colorado potato beetle, Lept<strong>in</strong>otarsa deceml<strong>in</strong>eata <strong>in</strong>vaded Europe from<br />
North America <strong>in</strong> the 1920s and reached <strong>Switzerland</strong> <strong>in</strong> 1937. It is a major pest of<br />
potato <strong>in</strong> Europe and North America (see Fact Sheet). More recently, the western corn<br />
rootworm, Diabrotica virgifera ssp. virgifera, a Nearctic <strong>species</strong>, took the same road<br />
and arrived <strong>in</strong> Belgrade <strong>in</strong> 1992. Twelve years on, it has spread to more than 12 European<br />
countries, <strong>in</strong>clud<strong>in</strong>g <strong>Switzerland</strong> (see Fact Sheet). D. virgifera virgifera is considered<br />
the major pest of maize <strong>in</strong> North America and substantial economic damage has<br />
already been observed <strong>in</strong> Central and eastern Europe.<br />
Some exotic beetles are damag<strong>in</strong>g to forest and ornamental trees. Xylophagous <strong>in</strong>sects<br />
are notorious for be<strong>in</strong>g easily <strong>in</strong>troduced <strong>in</strong>to new regions through the importation of<br />
timber or solid wood pack<strong>in</strong>g material. At least six Scolytidae <strong>in</strong> <strong>Switzerland</strong> are of<br />
<strong>alien</strong> orig<strong>in</strong>. In particular, the Asian Xylosandrus germanus and the North American<br />
Gnathotrichus materiarius damage freshly cut logs and reduce timber quality. They<br />
both arrived <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> the 1980s (see Fact Sheet for X. germanus; Hirschheydt<br />
(1992) for G. materiarius). Other wood bor<strong>in</strong>g scolytids arrived <strong>in</strong> <strong>Switzerland</strong> very<br />
recently: Tripodendron laeve from East Asia or Scand<strong>in</strong>avia, Xylebor<strong>in</strong>us alni from<br />
East Asia, and Xyleborus punctulatus from Siberia (C. Besuchet, pers. comm.). Similarly,<br />
the cerambycids Neoclytus acum<strong>in</strong>atus and Xylotrechus stebb<strong>in</strong>gi, xylophagous<br />
beetles from North America and the Himalayas, respectively, have recently been found<br />
<strong>in</strong> the Tic<strong>in</strong>o (C. Besuchet, pers. comm.). Although these <strong>species</strong> have not yet caused<br />
economic or environmental damage <strong>in</strong> <strong>Switzerland</strong>, their <strong>in</strong>troductions show that<br />
<strong>in</strong>ternational movement of timber and timber products is an important <strong>in</strong>troduction<br />
pathway for bark and wood bor<strong>in</strong>g beetles that may eventually lead to the establishment<br />
of serious forest pests. Among these potential <strong>in</strong>troductions are the two Asian<br />
cerambycids Anoplophora glabripennis (Motschulsky) and A. ch<strong>in</strong>ensis (Forster). Both<br />
<strong>species</strong> are important pests <strong>in</strong> Asia, attack<strong>in</strong>g and kill<strong>in</strong>g a whole range of tree <strong>species</strong>
4 > Insects – Insecta 75<br />
(see Fact Sheets for both <strong>species</strong>). They were both <strong>in</strong>troduced <strong>in</strong>to North America <strong>in</strong><br />
the 1990s, where they are now the targets of large-scale eradication programmes.<br />
These <strong>species</strong> have been often <strong>in</strong>tercepted at ports of entry <strong>in</strong>to Europe and, <strong>in</strong> the last<br />
four years, field populations have been found at various sites <strong>in</strong> Europe (<strong>in</strong> Austria,<br />
Germany and France for A. glabripennis; <strong>in</strong> Italy and France for A. ch<strong>in</strong>ensis). Eradication<br />
programmes have been established but the population of A. ch<strong>in</strong>ensis <strong>in</strong> Italy, at<br />
least, is considered to be established. The potential damage these two <strong>species</strong> could<br />
cause <strong>in</strong> Europe is unclear, but it could be spectacular. A. glabripennis is usually<br />
<strong>in</strong>troduced as eggs, larvae or pupae <strong>in</strong> solid wood pack<strong>in</strong>g material, i.e. crat<strong>in</strong>g, pallets<br />
or pack<strong>in</strong>g blocks from Ch<strong>in</strong>a. A. ch<strong>in</strong>ensis has been transported to North America,<br />
France and Italy <strong>in</strong> bonsai trees from Asia.<br />
Not all <strong>in</strong>troduced beetles <strong>in</strong> <strong>Switzerland</strong> are of economic importance. Many exotic<br />
<strong>species</strong> are found <strong>in</strong> decay<strong>in</strong>g plant material, compost, litter, etc., <strong>in</strong> particular several<br />
fungus beetles (Latridiidae) and rove beetles (Staphyl<strong>in</strong>idae), but also <strong>species</strong> <strong>in</strong> the<br />
families Hydrophilidae, Languridae, Merophysiidae, Orthoperidae, Ptiliidae, etc.<br />
Consider<strong>in</strong>g how little attention has been paid to these <strong>in</strong>sects and their ecosystems, it<br />
is likely that many more exotic <strong>species</strong> rema<strong>in</strong> unrecorded <strong>in</strong> <strong>Switzerland</strong>, and <strong>in</strong><br />
Europe <strong>in</strong> general. Some of the recorded <strong>species</strong> are particularly abundant <strong>in</strong> their<br />
habitat, suggest<strong>in</strong>g that direct or <strong>in</strong>direct <strong>in</strong>teractions with the native fauna occur.<br />
F<strong>in</strong>ally, another <strong>in</strong>vasive beetle worth mention<strong>in</strong>g <strong>in</strong> the multicoloured Asian ladybeetle,<br />
Harmonia axyridis. This cocc<strong>in</strong>ellid is a biological control agent widely used <strong>in</strong><br />
greenhouses aga<strong>in</strong>st aphids. It recently became established outdoors <strong>in</strong> several European<br />
countries. The first specimen was found <strong>in</strong> Basel <strong>in</strong> 2004 (Klausnitzer, 2004). It is<br />
also established s<strong>in</strong>ce the late 1980s <strong>in</strong> North America, where it has become a human<br />
nuisance because of its habit of <strong>in</strong>vad<strong>in</strong>g houses and build<strong>in</strong>gs <strong>in</strong> large numbers. Furthermore,<br />
it seems to decrease the diversity of native Cocc<strong>in</strong>ellidae, and it has also<br />
become a pest of fruit production, particularly <strong>in</strong> v<strong>in</strong>eyards (see Fact Sheet).<br />
4.3 Lepidoptera<br />
The Lepidoptera of Central Europe has been quite well studied, and the distribution of<br />
both native and exotic <strong>species</strong> is fairly well known compared with most other <strong>in</strong>sect<br />
orders. In this group, the ma<strong>in</strong> difficulty <strong>in</strong> establish<strong>in</strong>g a list of exotic <strong>species</strong> came<br />
from the high numbers of Mediterranean <strong>species</strong> that are occasionally observed <strong>in</strong><br />
<strong>Switzerland</strong>. The exotic or <strong>in</strong>vasive status of most of these <strong>species</strong> is unclear. For<br />
example, Rezbanyai-Reser (2000) lists Mediterranean Geometridae and Noctuidae<br />
which occasionally overw<strong>in</strong>ter <strong>in</strong> the Tic<strong>in</strong>o. Some <strong>species</strong> apparently became established<br />
only recently, such as the noctuids Mythimna unipuncta and Acantholeucania<br />
loreyi (Duponchel), perhaps as a result of climate change. Others, such as the well<br />
known noctuid pest Spodoptera exigua (Hübner), are occasional visitors that can<br />
overw<strong>in</strong>ter <strong>in</strong> the warmest areas of the country. Many other southern or eastern European<br />
<strong>species</strong>, <strong>in</strong>clud<strong>in</strong>g migrant <strong>species</strong>, are occasionally recorded <strong>in</strong> <strong>Switzerland</strong>. In<br />
general, these Lepidoptera are not <strong>in</strong>cluded <strong>in</strong> our list, because their <strong>in</strong>troduction and<br />
Advisors:<br />
Ladislaus Reser, Natur-Museum<br />
Luzern; Rudolf Bryner, Twann
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 76<br />
establishment <strong>in</strong> <strong>Switzerland</strong> are natural phenomena and not mediated by human<br />
activity. A few <strong>species</strong> whose <strong>in</strong>vasive status is <strong>in</strong> doubt are <strong>in</strong>cluded <strong>in</strong> the list, such as<br />
the geometrid Eupithecia s<strong>in</strong>uosaria, a European <strong>species</strong> for which the spread through<br />
Central Europe has been particularly well studied (Rezbanyai-Reser et al., 1998).<br />
Similarly, Caradr<strong>in</strong>a <strong>in</strong>grata is a noctuid of eastern Mediterranean orig<strong>in</strong> and is <strong>in</strong>creas<strong>in</strong>gly<br />
observed <strong>in</strong> urban areas, where it could have become established <strong>in</strong> warmer<br />
microclimates (Rezbanyai-Reser et al., 1997; Whitebread, 1997).<br />
Many Lepidoptera of exotic orig<strong>in</strong> are primary or secondary pests. Among the bestknown<br />
exotic <strong>species</strong> are several leaf m<strong>in</strong>ers <strong>in</strong> the family Gracillariidae, which all<br />
<strong>in</strong>vaded Europe <strong>in</strong> the last 30 years. Leaf m<strong>in</strong>ers are easily <strong>in</strong>troduced <strong>in</strong>to new areas<br />
because the m<strong>in</strong>es on fresh or dead leaves are often <strong>in</strong>conspicuous and readily carried<br />
over long distances. In addition, several <strong>species</strong> pupate <strong>in</strong> or near the leaf m<strong>in</strong>e, and<br />
these stages are less vulnerable to adverse conditions. The horse chestnut leaf m<strong>in</strong>er,<br />
Cameraria ohridella, is a moth of unknown orig<strong>in</strong> that was first found <strong>in</strong> Macedonia <strong>in</strong><br />
1984 and <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> 1998 (Kenis and Forster, 1998; see also Fact Sheet). In less<br />
than 20 years, it has <strong>in</strong>vaded most of Europe. The rapid spread of the moth <strong>in</strong> Europe is<br />
expla<strong>in</strong>ed by the transport of adults and dead leaves conta<strong>in</strong><strong>in</strong>g pupae <strong>in</strong> or on vehicles.<br />
It causes severe defoliation to horse chestnut (Aesculus hippocastanum L.) trees <strong>in</strong><br />
urban areas <strong>in</strong> most European regions as well as to <strong>in</strong>digenous horse chestnut stands <strong>in</strong><br />
the Balkans, where it represents a threat to the survival of the tree <strong>species</strong> <strong>in</strong> the wild.<br />
The North American <strong>species</strong> Phyllonorycter rob<strong>in</strong>iella and Parectopa rob<strong>in</strong>iella m<strong>in</strong>e<br />
leaves of their orig<strong>in</strong>al host, the North American black locust Rob<strong>in</strong>ia pseudoacacia L.<br />
(see Fact Sheet <strong>in</strong> the plants chapter). Phyllonorycter platani, a moth orig<strong>in</strong>at<strong>in</strong>g from<br />
the Balkans and Asia M<strong>in</strong>or, is commonly found on Platanus trees. Phyllonorycter<br />
leucographella, also of eastern Mediterranean orig<strong>in</strong>, has spread all over Europe and<br />
feeds on Pyracantha and Crataegus. Another gracillariid, the eastern Asian Caloptilia<br />
azaleella, m<strong>in</strong>es on Rhododendron <strong>in</strong> greenhouses. Ornamental Cupressaceae <strong>in</strong> Europe<br />
are attacked by another North American leaf m<strong>in</strong>er, the yponomeutid Argyresthia<br />
thuiella.<br />
The arctiid moth Hyphantria cunea is another <strong>in</strong>terest<strong>in</strong>g case of an <strong>in</strong>vasive Lepidoptera.<br />
Orig<strong>in</strong>at<strong>in</strong>g from North America, H. cunea arrived <strong>in</strong> the 1940s <strong>in</strong> Hungary, from<br />
where it spread to most of Europe. It was first found <strong>in</strong> the Tic<strong>in</strong>o <strong>in</strong> 1991 (Jerm<strong>in</strong>i et<br />
al., 1995; see also Fact Sheet). This polyphagous defoliator is considered a serious pest<br />
of forest and ornamental trees and shrubs <strong>in</strong> some eastern European countries and<br />
eastern Asia, where it was also <strong>in</strong>troduced. Cydia molesta is an Asian <strong>species</strong> present<br />
<strong>in</strong> orchards <strong>in</strong> <strong>Switzerland</strong> where it feeds on various fruit trees. The noctuid Helicoverpa<br />
armigera is a cosmopolitan, polyphagous pest, probably of African orig<strong>in</strong>,<br />
which occurs <strong>in</strong> <strong>Switzerland</strong> ma<strong>in</strong>ly <strong>in</strong> greenhouses. It is now also regularly found<br />
overw<strong>in</strong>ter<strong>in</strong>g <strong>in</strong> the Tess<strong>in</strong>, where it probably arrived by itself. Another greenhouse<br />
pest of African orig<strong>in</strong> is the banana moth, Opogona sacchari, which feeds on various<br />
woody and perennial ornamentals.<br />
Several exotic moths are pests of stored products <strong>in</strong> <strong>Switzerland</strong>. Most of them have<br />
been <strong>in</strong>troduced with commodities <strong>in</strong>to most parts of the world. The pyralid Plodia<br />
<strong>in</strong>terpunctella is a major pest of stored products <strong>in</strong> warehouses, gra<strong>in</strong> elevators and
4 > Insects – Insecta 77<br />
food factories, as well as <strong>in</strong> private households. It feeds on gra<strong>in</strong>s, nuts and various<br />
other dried products. In <strong>Switzerland</strong>, it is particularly prevalent <strong>in</strong> chocolate factories,<br />
together with other pyralids, Ephestia elutella and Cadra cautella (Hoppe, 1981). The<br />
pyralids E. kuehniella and Sitotroga cerealella are also cosmopolitan <strong>species</strong>, found<br />
ma<strong>in</strong>ly <strong>in</strong> stored gra<strong>in</strong>s. These pests of stored products are usually well controlled <strong>in</strong><br />
<strong>Switzerland</strong> and other developed countries, but their management makes extensive<br />
resource demands on the <strong>in</strong>dustry.<br />
4.4 Hymenoptera<br />
Only two Symphyta <strong>in</strong> the Swiss fauna are known to be of exotic orig<strong>in</strong>. The wood<br />
wasp Sirex cyaneus is a secondary forest pest. Larvae live <strong>in</strong> dead or dy<strong>in</strong>g trunks of<br />
Abies, ma<strong>in</strong>ly, but also Larix and Pseudotsuga (Schwenke, 1982). The <strong>in</strong>sect orig<strong>in</strong>ates<br />
from North America but <strong>in</strong>vaded Europe a long time ago, without caus<strong>in</strong>g damage. It is<br />
<strong>in</strong>terest<strong>in</strong>g to note that a European congeneric <strong>species</strong>, S. noctilio F., <strong>in</strong> Europe nearly<br />
as harmless as S. cyaneus, has become a major <strong>in</strong>troduced pest of p<strong>in</strong>e plantations <strong>in</strong><br />
Australia, New Zealand, South Africa and South America. The second exotic sawfly <strong>in</strong><br />
<strong>Switzerland</strong> is the nemat<strong>in</strong>e tenthred<strong>in</strong>id Nematus tibialis, an American <strong>species</strong> established<br />
<strong>in</strong> Europe where it feeds on its orig<strong>in</strong>al host, Rob<strong>in</strong>ia pseudoacacia.<br />
Most Hymenoptera belong to the Apocrita, and of these the vast majority are parasitic<br />
<strong>in</strong>sects. These are among the least known <strong>in</strong>sects. There is no checklist of parasitic<br />
Hymenoptera for <strong>Switzerland</strong>, and new, undescribed <strong>species</strong> are found every year <strong>in</strong><br />
Central Europe. Many European <strong>species</strong> are known to occur on other cont<strong>in</strong>ents as well<br />
but, for most of them, it is impossible to know whether their wide distribution is the<br />
result of an <strong>in</strong>troduction of exotic <strong>species</strong> <strong>in</strong>to Europe, or vice versa. Only the few<br />
<strong>species</strong> deliberately <strong>in</strong>troduced <strong>in</strong>to Europe as part of biological control programmes<br />
are def<strong>in</strong>itely exotic. For example, at least three <strong>alien</strong> parasitoid <strong>species</strong> have been<br />
<strong>in</strong>troduced <strong>in</strong>to <strong>Switzerland</strong> and have become permanently established. The aphel<strong>in</strong>id<br />
Aphel<strong>in</strong>us mali was released aga<strong>in</strong>st the woolly aphid Eriosoma lanigerum as early as<br />
1922 (Greathead, 1976). Another aphel<strong>in</strong>id wasp, Encarsia perniciosi, was <strong>in</strong>troduced<br />
more recently aga<strong>in</strong>st the San José scale Quadraspidiotus perniciosus, and the dry<strong>in</strong>id<br />
Neodry<strong>in</strong>us typhlocybae was released <strong>in</strong> the Tic<strong>in</strong>o <strong>in</strong> 1998–99 to control the flatid<br />
planthopper Metcalfa pru<strong>in</strong>osa (Mani and Baroffio, 1997; Jerm<strong>in</strong>i et al., 2000). Other<br />
parasitoids were released <strong>in</strong> neighbour<strong>in</strong>g countries and were subsequently recovered<br />
<strong>in</strong> <strong>Switzerland</strong>, such as Aphytis proclia, Encarsia berlesei and E. lounhuryi, all <strong>in</strong>troduced<br />
<strong>in</strong> Italy to control scale <strong>in</strong>sects (Greathead, 1976; Noyes, 2002). The encyrtid<br />
Ooencyrtus kuvanae, an Asian egg parasitoid of the gypsy moth, Lymantria dispar L.,<br />
was released <strong>in</strong> many European countries. It is has not been recorded from <strong>Switzerland</strong>,<br />
but s<strong>in</strong>ce it is present <strong>in</strong> all neighbour<strong>in</strong>g countries, there is no doubt that it also occurs<br />
<strong>in</strong> <strong>Switzerland</strong> (Greathead, 1976; Noyes, 2002).<br />
Other parasitic Hymenoptera of exotic orig<strong>in</strong> are used as augmentative biological<br />
control agents <strong>in</strong> Swiss greenhouses. The most commonly used <strong>species</strong>, such as the<br />
whitefly parasitoid Encarsia formosa, are mentioned <strong>in</strong> our list because they have<br />
Advisor:<br />
Bernhard Seifert, Staatliches<br />
Museum für Naturkunde, Görlitz,<br />
Germany (Formicidae)
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 78<br />
become part of greenhouse ecosystems although, <strong>in</strong> most cases, they cannot survive<br />
w<strong>in</strong>ter conditions outdoors.<br />
Many exotic seed chalcids of the genus Megastigmus (Torymidae) have been accidentally<br />
<strong>in</strong>troduced <strong>in</strong>to Europe with seed trad<strong>in</strong>g, some of them hav<strong>in</strong>g become pests <strong>in</strong><br />
European seed orchards (Roques and Skrzypczynska, 2003). Only M. spermothrophus,<br />
the North American Douglas fir seed chalcid, is reported from <strong>Switzerland</strong>, but many<br />
other <strong>species</strong> are known from neighbour<strong>in</strong>g countries and, thus, are probably present <strong>in</strong><br />
<strong>Switzerland</strong>.<br />
Two sphecid wasps have recently <strong>in</strong>vaded <strong>Switzerland</strong>. The American Isodontia<br />
mexicana has been found <strong>in</strong> the Tic<strong>in</strong>o and the Lemanic region s<strong>in</strong>ce the early 1990s<br />
(Vernier, 1995, 2000). It feeds on crickets and, given its abundance, its impact on<br />
native ecosystems would merit further attention. Sceliphron curvatum is an Asian<br />
<strong>species</strong> which arrived <strong>in</strong> Europe <strong>in</strong> the 1970s. The first specimens were found recently<br />
<strong>in</strong> <strong>Switzerland</strong> (Gonseth et al., 2001). Its nests are commonly found <strong>in</strong> houses and other<br />
build<strong>in</strong>gs.<br />
Five exotic ant <strong>species</strong> are recorded for the Swiss fauna (Freitag et al., 2000; Neumeyer<br />
and Seifert, 2005). These are ‘tramp’ ants of tropical and subtropical orig<strong>in</strong> but transported<br />
by human activities to most regions of the world. The most common is the<br />
Asian <strong>species</strong> Monomorium pharaonis, very abundant <strong>in</strong> build<strong>in</strong>gs and often considered<br />
as an urban pest. Hypoponera schau<strong>in</strong>slandi is found <strong>in</strong> greenhouses and other<br />
heated build<strong>in</strong>gs <strong>in</strong> Central Europe (Seifert, 2004). L<strong>in</strong>epithema humile is the famous<br />
Argent<strong>in</strong>e ant, which has <strong>in</strong>vaded a vast territory along the Mediterranean coast and<br />
has been occasionally recorded <strong>in</strong> <strong>Switzerland</strong> (e.g. Kutter, 1981). Neumeyer and<br />
Seifert (2005) state that it could soon become established <strong>in</strong> outdoor situations because<br />
of global warm<strong>in</strong>g. Tap<strong>in</strong>oma melanocephalum and Paratrech<strong>in</strong>a longicornis have<br />
been observed <strong>in</strong> few occasions <strong>in</strong>doors <strong>in</strong> <strong>Switzerland</strong> and their establishment is not<br />
firmly established (Dorn et al., 1997; Freitag et al., 2000). Another ant, Lasius neglectus<br />
Van Loon, Boomsma & Andrasfalvy, an Asian <strong>species</strong>, is rapidly <strong>in</strong>vad<strong>in</strong>g Europe,<br />
caus<strong>in</strong>g some damage <strong>in</strong> urban areas (Seifert, 2000). It is not yet recorded from <strong>Switzerland</strong>,<br />
but its occurrence <strong>in</strong> neighbour<strong>in</strong>g countries (France, Italy and Germany)<br />
suggests that it may be already present.<br />
F<strong>in</strong>ally, the chestnut gallwasp, Dryocosmus kuriphilus Yasumatsu (Hym.: Cynipidae),<br />
is worth mention<strong>in</strong>g as a potential threat for the European chestnut, Castanea sativa, <strong>in</strong><br />
<strong>Switzerland</strong> and the rest of Europe. This Ch<strong>in</strong>ese <strong>species</strong> was found for the first time <strong>in</strong><br />
Northern Italy <strong>in</strong> 2002 and is considered as the most serious <strong>in</strong>sect pest of Castanea<br />
spp. worldwide (Bosio, 2004). Attacks of Dryocosmus kuriphilus reduce growth of<br />
young chestnuts and fruit<strong>in</strong>g. Yield reductions of 50–70 % are observed and severe<br />
<strong>in</strong>festations may result <strong>in</strong> the dead of the tree.
4 > Insects – Insecta 79<br />
4.5 Diptera<br />
Checklists of Swiss Diptera have been published recently (Merz et al., 1998, 2001).<br />
These do not conta<strong>in</strong> <strong>in</strong>formation on the exotic or <strong>in</strong>vasive status of the <strong>species</strong> but the<br />
authors (Drs. B. Merz, J.-P. Haenni and G. Baechli) were contacted and k<strong>in</strong>dly reviewed<br />
the list to extract the <strong>species</strong> which are thought to be exotic.<br />
Less than 20 dipteran <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> are of <strong>alien</strong> orig<strong>in</strong>. Some are agricultural<br />
or horticultural pests. The Nearctic agromyzids Liriomyza huidobrensis and L. trifolii<br />
are pests of vegetables, ma<strong>in</strong>ly <strong>in</strong> greenhouses but occasionally found outdoors. Another<br />
agromyzid fly, Napomyza gymnostoma, was recently found <strong>in</strong> <strong>Switzerland</strong> (Eder<br />
and Bauer, 2003). N. gymnostoma, previously known as a harmless <strong>in</strong>sect from some<br />
European countries, but not <strong>Switzerland</strong>, mysteriously became a serious pest of onions<br />
and leeks one or two decades ago <strong>in</strong> several Central and western European countries.<br />
Its orig<strong>in</strong> is not known, although it may be a virulent ecotype of a European <strong>species</strong><br />
that unexpectedly adapted itself to agricultural systems and crops. Its sudden pest status<br />
and spread may also be due to changes <strong>in</strong> pest control methods <strong>in</strong> leek and onion crops,<br />
or to global warm<strong>in</strong>g.<br />
Two of the world’s most serious fruit flies (Tephritidae), the olive fly Bactrocera oleae<br />
and the Mediterranean fruit fly, Ceratitis capitata, both orig<strong>in</strong>at<strong>in</strong>g from the Mediterranean<br />
region, are also established <strong>in</strong> <strong>Switzerland</strong>, where they have reached their<br />
climatic limit and, thus, cause little damage. Two North American tephritids have<br />
recently <strong>in</strong>vaded the country. Rhagoletis completa feeds on walnut and is considered a<br />
pest <strong>in</strong> its native region. It was discovered <strong>in</strong> the Tic<strong>in</strong>o <strong>in</strong> 1991 (Merz, 1991) and,<br />
s<strong>in</strong>ce then, has spread rapidly to many regions north of the Alps (see Fact Sheet).<br />
Populations are <strong>in</strong>creas<strong>in</strong>g and severe damage has been observed <strong>in</strong> <strong>Switzerland</strong> and<br />
Italy. Another American <strong>species</strong>, R. c<strong>in</strong>gulata, was found at the same time, <strong>in</strong> cherries<br />
(Merz, 1991, as R. <strong>in</strong>differens). Until now, populations have rema<strong>in</strong>ed very low and it<br />
does not seem to spread as quickly as R. completa.<br />
The North American Drosophilidae Chymomyza amoena attacks various fruits of<br />
broadleaved trees, such as apple, walnut, plum, acorn, etc. It is present <strong>in</strong> high numbers<br />
<strong>in</strong> both urban and forest environments. Its spread and ecology <strong>in</strong> <strong>Switzerland</strong> have been<br />
studied <strong>in</strong> detail (e.g. Burla and Baechli, 1992; Band, 1995; Band et al., 1998, 1999).<br />
Another drosophilid, Drosophila curvisp<strong>in</strong>a, orig<strong>in</strong>at<strong>in</strong>g from east Asia, is present <strong>in</strong><br />
the Tic<strong>in</strong>o, Vaud and Valais (Bächli et al., 2002; Bächli, pers. comm.).<br />
Other <strong>alien</strong> Diptera occurr<strong>in</strong>g <strong>in</strong> <strong>Switzerland</strong> are saprophagous or coprophagous. These<br />
<strong>species</strong> are often of cosmopolitan distribution. The calliphorid Chrysomya albiceps and<br />
the muscid Hydrotaea aenescens are found on human cadavers and other decay<strong>in</strong>g<br />
matters. The black soldier fly, Hermetia illucens (Stratiomyidae), was found <strong>in</strong> very<br />
high numbers <strong>in</strong> composted plants <strong>in</strong> the Tic<strong>in</strong>o (Sauter, 1989). In North America, this<br />
very common <strong>species</strong> is often associated with poultry houses and other farm<strong>in</strong>g or<br />
animal-rear<strong>in</strong>g activities. Only one milichiid, Desmometopa varipalpis, is mentioned <strong>in</strong><br />
our list as clearly exotic. However, the Milichiidae are small, often cosmopolitan and<br />
Advisors:<br />
Bernhard Merz, Muséum<br />
d’Histoire Naturelle, Genève;<br />
Jean-Paul Haenni, Muséum<br />
d’Histoire Naturelle, Neuchâtel;<br />
Gerhard Baechli, Zoologisches<br />
Museum, Zürich; Wolfgang Billen,<br />
Amt für Landwirtschaft, Lörrach,<br />
Germany
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 80<br />
poorly known <strong>in</strong>sects, and it is possible that other milichiid members of the Swiss<br />
fauna are also of exotic orig<strong>in</strong> (B. Merz, pers. comm.)<br />
Exotic mosquitos (Culicidae) represent a major threat to human health worldwide. The<br />
tiger mosquito, Aedes albopictus, was found <strong>in</strong> <strong>Switzerland</strong> for the first time <strong>in</strong> the<br />
Tic<strong>in</strong>o <strong>in</strong> 2003 (see Fact Sheet). Besides caus<strong>in</strong>g human nuisance through its bites, A.<br />
albopticus is also a potential vector of various illnesses. In Asia, its region of orig<strong>in</strong>, it<br />
is a natural vector of dengue fever and other arboviruses, as well as filaria for both<br />
human and domestic animals. In North America, it is a vector of the West Nile virus.<br />
4.6 Hemiptera<br />
This order probably encompasses the highest number of exotic pests worldwide. Small<br />
Hemiptera, particularly aphids, scales, whiteflies and psyllids, are very easily carried<br />
around the globe on plant material. Many are pests of worldwide distribution and it is<br />
sometimes difficult to assess their region of orig<strong>in</strong>. Among the cosmopolitan <strong>species</strong> of<br />
doubtful orig<strong>in</strong> and listed as exotic <strong>in</strong> <strong>Switzerland</strong>, for example, are the aphids Myzus<br />
persicae, Aphis gossypii and C<strong>in</strong>ara cupressi, the whitefly Bemisia tabaci, and the<br />
scales Dynaspidiotus britannicus, Quadraspidiotus pyri and Planococcus citri. Many<br />
other cosmopolitan <strong>species</strong> are not <strong>in</strong>cluded <strong>in</strong> this list because they are supposed to be<br />
of European orig<strong>in</strong> (e.g. the aphids Acyrthosiphon pisum, Brevicoryne brassicae and<br />
Rhopalosiphum padi), although their orig<strong>in</strong> is unclear and the possibility that they<br />
entered <strong>Switzerland</strong> a long time ago cannot be ruled out.<br />
Another problem, particularly prevalent <strong>in</strong> Hemiptera, is the high number of tropical or<br />
subtropical pest <strong>species</strong> <strong>in</strong>troduced <strong>in</strong>to greenhouses with their host plants. The most<br />
abundant <strong>species</strong> have been <strong>in</strong>cluded <strong>in</strong> the list because there is no doubt that they have<br />
become established <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> <strong>in</strong>door conditions. For example, the aphid Aphis<br />
gossypii, the whiteflies Bemisia tabaci and Trialeurodes vaporarium and the scale<br />
Planococcus citri are recurrent problems <strong>in</strong> protected crops and plants <strong>in</strong> <strong>Switzerland</strong>.<br />
Many other tropical and subtropical <strong>species</strong> are occasionally reported from greenhouses<br />
and <strong>in</strong>door plants, but it is difficult to say whether their presence results from a<br />
permanent establishment or from regular <strong>in</strong>troductions. In addition to those <strong>species</strong><br />
<strong>in</strong>cluded <strong>in</strong> our list, Kozar et al. (1994) mentions several <strong>species</strong> of scale <strong>in</strong>sects that<br />
have been recovered from greenhouses and <strong>in</strong>door plants <strong>in</strong> <strong>Switzerland</strong>. Interest<strong>in</strong>gly,<br />
three of these scale <strong>species</strong> previously known only from greenhouses (Diaspidiotus<br />
dist<strong>in</strong>ctus, Coccus hesperidum and Icerya purchasi) are now found established <strong>in</strong> the<br />
field (Kozar et al., 1994). Other ‘greenhouse’ Hemiptera of exotic orig<strong>in</strong> and not<br />
mentioned <strong>in</strong> our list are bug <strong>species</strong> of the genera Orius and Xylocoris (Anthocoridae),<br />
commonly used as biological control agents aga<strong>in</strong>st thrips, spider mites or aphids.<br />
Many hemipteran <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> have migrated from neighbour<strong>in</strong>g countries,<br />
especially the Mediterranean region. In most cases these have probably been <strong>in</strong>troduced<br />
with their host plant. For example, most Psyllidae and Triozidae, <strong>in</strong>cluded <strong>in</strong> the<br />
list feed specifically on plants of Mediterranean orig<strong>in</strong> which are used <strong>in</strong> <strong>Switzerland</strong><br />
Advisors:<br />
Daniel Burckhardt,<br />
Naturhistorisches Museum Basel<br />
(Psyll<strong>in</strong>a); Gerolf Lampel, Pensier,<br />
and Yves Gonseth, Centre Suisse<br />
de Cartographie de la Faune,<br />
Neuchâtel (Aphid<strong>in</strong>a); Ralf<br />
Heckmann, Konstanz, Germany<br />
(Heteroptera)
4 > Insects – Insecta 81<br />
as ornamentals. However, other southern European <strong>species</strong> may have entered <strong>Switzerland</strong><br />
<strong>in</strong>dependently. In particular, four true bugs, the lygaeids Arocatus longiceps,<br />
Orsillus depressus and Oxycarenus lavaterae, and the mirid Deraeocoris flavil<strong>in</strong>ea are<br />
clearly expand<strong>in</strong>g their range from southern to Central Europe (e.g. Adlbauer and<br />
Rabitsch, 2000; Rabitsch, 2002). Whether the spread is purely natural or not rema<strong>in</strong>s<br />
unclear.<br />
Most of the known exotic Hemiptera <strong>in</strong>cluded <strong>in</strong> the list are recognized as pests, feed<strong>in</strong>g<br />
on crops, ornamentals and forest trees <strong>in</strong> <strong>Switzerland</strong>. It is likely that many other<br />
exotic Hemiptera rema<strong>in</strong> undetected because they feed on non-commercial plants.<br />
Among aphids, one of the best-known cases of exotic pests <strong>in</strong> Europe is the grape<br />
phylloxera, Viteus vitifoliae which <strong>in</strong>vaded Europe from North America <strong>in</strong> the 19 th<br />
century, caus<strong>in</strong>g serious damage to v<strong>in</strong>eyards and endanger<strong>in</strong>g the European w<strong>in</strong>e<br />
<strong>in</strong>dustry. The problem was solved by graft<strong>in</strong>g European cultivars on less susceptible<br />
American rootstocks; however, the level of damage has <strong>in</strong>creased aga<strong>in</strong> <strong>in</strong> recent years,<br />
<strong>in</strong>clud<strong>in</strong>g <strong>in</strong> <strong>Switzerland</strong>. Other aphid <strong>species</strong>, such as Myzus persicae, Macrosiphum<br />
euphorbiae and Aphis gossypii, attack a wide range of vegetable crops, both <strong>in</strong>doors<br />
and outdoors. They are also vectors of serious viral diseases. The Russian wheat aphid,<br />
Diuraphis noxia (Kurdjumov), a serious pest of cereals, has not yet been reported from<br />
<strong>Switzerland</strong>, but has already <strong>in</strong>vaded France, Italy and Austria (CABI, 2001; Lethmayer<br />
and Rabitsch, 2002). Orchard trees <strong>in</strong> <strong>Switzerland</strong>, <strong>in</strong> particular apple trees, can<br />
be severely damaged by the North American woolly aphid Eriosoma lanigerum and the<br />
Asian Aphis spiraecola. In forestry, the most serious exotic aphid is certa<strong>in</strong>ly the<br />
woolly aphid Dreyfusia nordmannianae, a pest of firs (Abies) of Caucasian orig<strong>in</strong>, but<br />
other <strong>species</strong> such as Gilletteella cooleyi on Pseudotsuga and Elatobium abiet<strong>in</strong>um on<br />
Picea, both from North America, may cause some concern to foresters.<br />
Only two exotic whitefly <strong>species</strong> (Aleyrodidae) are known to occur <strong>in</strong> <strong>Switzerland</strong>,<br />
Bemisia tabaci and Trialeurodes vaporarium, but these are among the ma<strong>in</strong> pests of<br />
vegetables <strong>in</strong> greenhouses. Control is achieved us<strong>in</strong>g aphel<strong>in</strong>id parasitoids <strong>in</strong> the<br />
genera Encarsia and Eretmocerus.<br />
A number of exotic scale <strong>in</strong>sects are also known as pests <strong>in</strong> <strong>Switzerland</strong>, particularly on<br />
orchard and ornamental trees. The San José scale, Quadraspidiotus perniciosus, is an<br />
Asian <strong>species</strong> that has <strong>in</strong>vaded nearly all cont<strong>in</strong>ents, <strong>in</strong>clud<strong>in</strong>g Europe, where it is still<br />
expand<strong>in</strong>g its range. It is already distributed <strong>in</strong> most parts of <strong>Switzerland</strong>, where it<br />
causes serious damage to orchards, particularly apple, peach and plum (see Fact Sheet).<br />
Another congeneric <strong>species</strong>, Q. pyri, also attacks fruit trees, but its orig<strong>in</strong> rema<strong>in</strong>s<br />
uncerta<strong>in</strong>, and it may be <strong>in</strong>digenous. In recent years, serious damage by scale <strong>in</strong>sects<br />
has been observed on urban trees. The diaspidid Pseudaulacaspis pentagona, an <strong>in</strong>sect<br />
of Asian orig<strong>in</strong> and known to be polyphagous attack<strong>in</strong>g ma<strong>in</strong>ly mulberry and peach<br />
trees, has caused severe damage to ornamental trees such as Sophora, Aesculus and<br />
Catalpa <strong>in</strong> Swiss cities (Mani et al., 1997). Similarly, the coccid Pulv<strong>in</strong>aria regalis<br />
(see Fact Sheet) is a polyphagous exotic <strong>species</strong> of uncerta<strong>in</strong> orig<strong>in</strong> that has recently<br />
caused heavy damage to Tilia and Aesculus <strong>in</strong> Zürich (Hippe and Frey, 1999).
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 82<br />
Only seven <strong>species</strong> of Auchenorrhyncha are listed as exotic <strong>in</strong> <strong>Switzerland</strong>. However,<br />
some of them are of economic importance. Metcalfa pru<strong>in</strong>osa, a flatid planthopper of<br />
American orig<strong>in</strong>, has recently <strong>in</strong>vaded the Tic<strong>in</strong>o through Italy (see Fact Sheet). This<br />
polyphagous <strong>species</strong> is found on many trees and shrubs, but also on crops such as<br />
soyabean. It can be particularly harmful to fruits such as grape, pear, apple, citrus and<br />
peach. It is expected that M. pru<strong>in</strong>osa will expand its distribution to most of <strong>Switzerland</strong>.<br />
Another North American <strong>species</strong>, the v<strong>in</strong>e leafhopper Scaphoideus titatus, recently<br />
entered <strong>Switzerland</strong>. It does not cause direct damage to v<strong>in</strong>e, but it transmits a<br />
severe mycoplasma disease, the ‘flavescence dorée’ (Günthart, 1987). Two other North<br />
American <strong>species</strong> are present <strong>in</strong> <strong>Switzerland</strong>, the cicadellid leafhopper Graphocephala<br />
fennahi on Rhododendron, and the membracid treehopper Stictocephala bisonia, a<br />
polyphagous <strong>species</strong> found ma<strong>in</strong>ly <strong>in</strong> orchards, but neither of them causes serious<br />
damage.<br />
The only heteropteran bugs of non-European orig<strong>in</strong> known to occur <strong>in</strong> <strong>Switzerland</strong> are<br />
the two North American lace bugs Corythucha ciliata and Corythucha arcuata.<br />
C. ciliata is a serious pest of Platanus <strong>in</strong> urban areas <strong>in</strong> Europe. Heavy <strong>in</strong>festations<br />
cause discolouration of leaves and premature leaf fall. It was first observed <strong>in</strong> <strong>Switzerland</strong><br />
<strong>in</strong> 1983 and its distribution now covers most of western <strong>Switzerland</strong> (see Fact<br />
Sheet). Corythucha arcuata is a similar <strong>species</strong>, feed<strong>in</strong>g on oak. It was first found <strong>in</strong><br />
Italy <strong>in</strong> 2000 and <strong>in</strong> 2003 <strong>in</strong> <strong>Switzerland</strong> (Tic<strong>in</strong>o) (see Fact Sheet). Similarly, another<br />
American bug, the coreid Leptoglossus occidentalis Heidemann, has recently been<br />
found <strong>in</strong> high numbers <strong>in</strong> northern Italy. It feeds on various conifer seeds and is considered<br />
a serious pest <strong>in</strong> seed orchards <strong>in</strong> North America (Villa et al., 2001).<br />
4.7 Orthoptera<br />
The Orthoptera of <strong>Switzerland</strong> and their distributions are listed <strong>in</strong> Thorens and Nadig<br />
(1997). Only two <strong>species</strong> established <strong>in</strong> the country are clearly exotic. The house<br />
cricket Acheta domesticus is a cosmopolitan <strong>species</strong>, probably of North African orig<strong>in</strong>.<br />
In Central Europe, it lives ma<strong>in</strong>ly <strong>in</strong> build<strong>in</strong>gs, although <strong>in</strong> summer it is also commonly<br />
found outdoors, particularly <strong>in</strong> the Valais. The house cricket is omnivorous, feed<strong>in</strong>g<br />
ma<strong>in</strong>ly on refuse and, occasionally, on stored products.<br />
The greenhouse camel cricket, Tachyc<strong>in</strong>es asynomorus, probably orig<strong>in</strong>ates from east<br />
Asia, but is now found worldwide. It was <strong>in</strong>troduced <strong>in</strong>to Europe <strong>in</strong> the 19 th century. In<br />
<strong>Switzerland</strong>, it lives ma<strong>in</strong>ly <strong>in</strong> greenhouses, where it feeds on fruits, seedl<strong>in</strong>gs and<br />
<strong>in</strong>sects. It is also occasionally found outdoors, e.g. <strong>in</strong> botanical gardens.<br />
Some Mediterranean <strong>species</strong> such as the Egyptian grasshopper Anacridium aegyptium<br />
(L.) are occasionally caught <strong>in</strong> <strong>Switzerland</strong>. Specimens found <strong>in</strong> the Tic<strong>in</strong>o may have<br />
migrated from the South, however, it seems that at least some catches <strong>in</strong> various regions<br />
may be related to the importation of food products. (Nadig and Thorens, 1991;<br />
Rezbanyai-Reser, 1993).<br />
Advisor:<br />
Hannes Bauer, Naturhistorisches<br />
Museum, Bern
4 > Insects – Insecta 83<br />
4.8 Dictyoptera<br />
At least five exotic cockroach <strong>species</strong> are known to be established <strong>in</strong> <strong>Switzerland</strong>.<br />
Others may be <strong>in</strong>troduced occasionally with imported goods. Unlike the native cockroaches,<br />
the exotic <strong>species</strong> are synanthropic, i.e. they are always found <strong>in</strong> association<br />
with humans. These cockroaches are of tropical and subtropical orig<strong>in</strong> but they now<br />
occur worldwide. Some <strong>species</strong>, such as Blatta orientalis and Periplaneta americana,<br />
have been cosmopolitan for such a long time that their orig<strong>in</strong> is uncerta<strong>in</strong>. B. orientalis,<br />
P. americana, P. australasiae and B. germanica have been established <strong>in</strong> <strong>Switzerland</strong><br />
for a long time, whereas Supella longipalpa is a recent <strong>in</strong>troduction.<br />
All these <strong>species</strong> are considered as serious urban pests. They are gregarious, nocturnal,<br />
and difficult to eradicate <strong>in</strong> houses. They can eat almost anyth<strong>in</strong>g, e.g. human food and<br />
animal feed, bookb<strong>in</strong>d<strong>in</strong>gs, wallpaper, excrement, leather products, etc. They can carry<br />
organisms caus<strong>in</strong>g human diseases, have a repulsive odour, and can cause allergic<br />
reaction and anxiety <strong>in</strong> some <strong>in</strong>dividuals. In <strong>Switzerland</strong>, the most problematic <strong>species</strong><br />
are B. germanica, S. longipalpa and B. orientalis, whereas Periplaneta spp. are of<br />
m<strong>in</strong>or importance (Landau et al., 1999).<br />
4.9 Isoptera<br />
There is no record of establishment of exotic termites <strong>in</strong> <strong>Switzerland</strong>. However, <strong>in</strong><br />
recent years, subterranean termites of the genus Reticulitermes have <strong>in</strong>vaded new areas<br />
(see Fact Sheet). The North American R. flavipes (Kollar) is <strong>in</strong>troduced <strong>in</strong> some German<br />
and Austrian cities. The southern European R. grassei Clément occurs now <strong>in</strong><br />
southern England and R. santonensis de Feytaud, a <strong>species</strong> of uncerta<strong>in</strong> orig<strong>in</strong> (and<br />
perhaps synonymous with R. flavipes) has spread from south-western to northern<br />
France. These termites undoubtedly have the potential to <strong>in</strong>vade the warmest areas <strong>in</strong><br />
<strong>Switzerland</strong>. They live <strong>in</strong> colonies <strong>in</strong> the soil, ma<strong>in</strong>ly <strong>in</strong> urban areas. They are particularly<br />
harmful to wooden elements <strong>in</strong> build<strong>in</strong>g but can also attack liv<strong>in</strong>g trees. The<br />
spread of subterranean termites <strong>in</strong> France is caus<strong>in</strong>g major concern. New regulations<br />
have been set up to limit the spread.<br />
4.10 Thysanoptera<br />
Many exotic thrips <strong>species</strong> are found, regularly or occasionally, <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong><br />
greenhouses. However, it is not always clear whether they are firmly established <strong>in</strong> the<br />
country. At least four <strong>species</strong> are considered permanently established, ma<strong>in</strong>ly <strong>in</strong> greenhouses.<br />
Three of them, Frankl<strong>in</strong>iella <strong>in</strong>tonsa, F. occidentalis and Heliothrips haemorrhoidalis<br />
are polyphagous, cosmopolitan pests on ornamentals, vegetables, and fruits.<br />
F. occidentalis, the western flower thrips, is among the most destructive greenhouse<br />
pests <strong>in</strong> <strong>Switzerland</strong> (Ebener et al., 1989; Schmidt and Frey, 1995). It is also a vector of<br />
various viruses. The pest status of F. occidentalis is relatively recent. It occurs naturally<br />
<strong>in</strong> North America on wild flowers and emerged as a greenhouse pest <strong>in</strong> the Neth-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 84<br />
erlands <strong>in</strong> 1983. S<strong>in</strong>ce then, it has spread all around the world (CABI, 2001). In <strong>Switzerland</strong>,<br />
F. <strong>in</strong>tonsa is particularly associated with strawberry crops (L<strong>in</strong>der et al.,<br />
1998).<br />
The gladiolus thrips, Thrips simplex, orig<strong>in</strong>ates from South Africa and was <strong>in</strong>troduced<br />
<strong>in</strong> many regions with gladiolus corms. It reproduces only on gladiolus corms and<br />
leaves but adults are found on many other plants (CABI, 2001). In <strong>Switzerland</strong>, it does<br />
not survive outdoors.<br />
4.11 Psocoptera<br />
Psocoptera, or psocids, are rather poorly known <strong>in</strong>sects. Most <strong>species</strong> live <strong>in</strong> woodlands,<br />
but some are domestic, i.e. they <strong>in</strong>habit houses, warehouses, etc., feed<strong>in</strong>g on<br />
organic matter. The domestic <strong>species</strong> can become a nuisance, although they rarely<br />
cause economic damage. Partial European and Swiss lists of Psocoptera exist, but the<br />
orig<strong>in</strong> of the <strong>species</strong> is often unclear. Lienhard (1994) lists 29 <strong>species</strong> that are totally,<br />
essentially or occasionally domestic <strong>in</strong> <strong>Switzerland</strong>. Most of them are not, or very<br />
rarely, found <strong>in</strong> the field, suggest<strong>in</strong>g that they are not <strong>in</strong>digenous. We have <strong>in</strong>cluded <strong>in</strong><br />
our list the <strong>species</strong> that Lienhard (1994) mentions as exclusively or essentially domestic<br />
<strong>in</strong> <strong>Switzerland</strong>, keep<strong>in</strong>g <strong>in</strong> m<strong>in</strong>d that some of them might be <strong>in</strong>digenous but poorly<br />
known <strong>in</strong> their natural habitat.<br />
Three <strong>species</strong> of the genus Dorypteryx are most likely of exotic orig<strong>in</strong>. Their spread <strong>in</strong><br />
Europe is illustrated <strong>in</strong> Lienhard (1994). D. domestica, described from Zimbabwe <strong>in</strong><br />
1958, was first found <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> 1973. From there, it has spread rapidly to most<br />
European countries where it has become one of the most frequent domestic <strong>species</strong>. D.<br />
longipennis, a <strong>species</strong> of unknown orig<strong>in</strong>, was first found <strong>in</strong> Luxembourg <strong>in</strong> 1988. It<br />
reached <strong>Switzerland</strong> <strong>in</strong> 1992 and is now spread<strong>in</strong>g very rapidly. D. pallida is an older<br />
<strong>in</strong>troduction. It was first described from North America, and found <strong>in</strong> Germany <strong>in</strong><br />
1890. Its spread seems to be slower than that of the other two <strong>species</strong>. Other <strong>species</strong><br />
that are most probably exotic <strong>in</strong>clude Liposcelis mendax, Ectopsocus pumilis and E.<br />
richardsi, which have been found associated with an importation of dry mushrooms<br />
from Asia. Whether these rare <strong>species</strong> are established <strong>in</strong> <strong>Switzerland</strong> is unclear. Other<br />
<strong>species</strong> are found outdoors <strong>in</strong> the Mediterranean regions but only <strong>in</strong>doors <strong>in</strong> <strong>Switzerland</strong>,<br />
e.g. Trogium pulsatorium and Ectopsocus vachoni.<br />
4.12 Ectoparasites<br />
Ectoparasitic <strong>in</strong>sects of vertebrates <strong>in</strong>clude ma<strong>in</strong>ly Siphonaptera (fleas – on birds and<br />
mammals), Anoplura (suck<strong>in</strong>g lice – on mammals) and Mallophaga (bit<strong>in</strong>g lice – on<br />
birds and mammals). Undoubtedly, several of the ectoparasites present <strong>in</strong> <strong>Switzerland</strong><br />
are allochtonous, hav<strong>in</strong>g arrived <strong>in</strong> the region with their hosts. However, these <strong>in</strong>sects<br />
are rather poorly studied, particularly <strong>in</strong> <strong>Switzerland</strong>, and their orig<strong>in</strong> is often unclear.<br />
In their work on the Siphonaptera of France and the western Mediterranean region,<br />
Advisor:<br />
Yves Gonseth, Centre Suisse de<br />
Cartographie de la Faune,<br />
Neuchâtel
4 > Insects – Insecta 85<br />
Beaucournu and Launay (1990) provided data for <strong>Switzerland</strong>. At least two fleas<br />
appear to be def<strong>in</strong>itely exotic <strong>in</strong> the country. The cat flea, Ctenocephalides felis felis,<br />
has a cosmopolitan distribution but is probably not of European orig<strong>in</strong> (Beaucournu<br />
and Launay, 1990). The rabbit flea, Spilopsylus cuniculi, is a vector of myxomatosis<br />
and probably orig<strong>in</strong>ates, like its host, from the Iberian Pen<strong>in</strong>sula, while the disease was<br />
<strong>in</strong>troduced <strong>in</strong>to Europe from South America. Some other fleas present <strong>in</strong> <strong>Switzerland</strong><br />
are of doubtful orig<strong>in</strong> and are not <strong>in</strong>cluded <strong>in</strong> the list. For example, the human flea<br />
Pulex irritans L., which feeds on many carnivores, is now cosmopolitan but belongs to<br />
a Nearctic genus. However, its arrival <strong>in</strong> Europe was probably <strong>in</strong> the distant past.<br />
Büttiker and Mahnert (1978) listed 25 Anoplura for <strong>Switzerland</strong>. Only one, Haemodipsus<br />
ventricosus, is very likely to be of exotic orig<strong>in</strong> because it seems to be restricted to<br />
rabbits. Suck<strong>in</strong>g lice of doubtful orig<strong>in</strong> <strong>in</strong>clude the cosmopolitan rat louse, Polyplax<br />
sp<strong>in</strong>ulosa (Burmeister), and the dog suck<strong>in</strong>g louse, L<strong>in</strong>ognathus setosus (v. Olfers).<br />
There is no recent list of Mallophaga for <strong>Switzerland</strong>. Mey (1988) provides a list of<br />
mammalian Mallophaga <strong>in</strong> Europe, but data for <strong>Switzerland</strong> are scarce. Three South<br />
American <strong>species</strong>, Gyropus ovalis Burmeister, Gliricola porcelli (Schrank) and Trimenopon<br />
hispidum (Burmeister), are found on gu<strong>in</strong>ea pigs <strong>in</strong> neighbour<strong>in</strong>g countries<br />
and, thus, are probably present <strong>in</strong> <strong>Switzerland</strong>. Similarly, it is possible that the Nearctic<br />
Trichodectes octomaculatus Pa<strong>in</strong>e and Pitrufquenia coypus Marelli, parasites on<br />
raccoons and coypus, respectively, <strong>in</strong> Central Europe, are present <strong>in</strong> <strong>Switzerland</strong> with<br />
their host.<br />
References<br />
Adlbauer, K. and W. Rabitsch (2000) Orsillus depressus (Mulsant & Rey<br />
1952) <strong>in</strong> Österreich und Liechtenste<strong>in</strong> (Het., Lygaeidae). Heteropteron 8,<br />
19–22.<br />
Bächli, G., Vilela, C.R. and F. Har<strong>in</strong>g (2002) Four new <strong>species</strong> of West<br />
Palaearctic Drosophilidae (Diptera). Mitteilungen der <strong>Schweizer</strong>ischen<br />
Entomologischen Gesellschaft 75, 299–333.<br />
Band, H.T. (1995) Is Chymomyza amoena (Loew) (Diptera: Drosophilidae)<br />
a versatile, coloniz<strong>in</strong>g <strong>species</strong>? Mitteilungen der <strong>Schweizer</strong>ischen<br />
Entomologischen Gesellschaft 68, 23–33.<br />
Band, H., Bächli, G. and R.N. Band (1999) Nearctic Chymomyza amoena<br />
(Loew) (Diptera: Drosophilidae) rema<strong>in</strong>s a domestic <strong>species</strong> <strong>in</strong><br />
<strong>Switzerland</strong>. Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 72, 75–82.<br />
Band, H., Band, R. N. and G. Bächli (1998) Further studies on Nearctic<br />
Chymomyza amoena (Loew) (Diptera, Drosophilidae) <strong>in</strong> <strong>Switzerland</strong>.<br />
Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen Gesellschaft 71,<br />
395–405.<br />
Barbalat, S. and B. Wermel<strong>in</strong>ger (1996) Première capture d'Agrilus<br />
guer<strong>in</strong>i Lac. (Col. Buprestidae) en Suisse. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 69, 201–202.<br />
Barbey, S. (1996) La protection phytosanitaire des conifères et arbustes<br />
d’ornement. Revue Horticole Suisse 69, 120–122.<br />
Beaucournu, J.-C. and Launay, H. (1990) Les puces (Sipohnaptera) de<br />
France et du Bass<strong>in</strong> méditerranéan occidental. Faune de France 76.<br />
Blattner, S. (1959) Sed<strong>in</strong>a buettneri Her<strong>in</strong>g, neu für die Schweiz.<br />
Mitteilungen der Entomologischen Gesellschaft Basel 9, 107–109.<br />
Bonavia, M. and M. Jerm<strong>in</strong>i (1998) La cicadelle Metcalfa pru<strong>in</strong>osa Say au<br />
Tess<strong>in</strong>. Distribution actuelle, dynamique des populations et perspective<br />
de lutte. Revue Suisse de Viticulture, Arboriculure et Horticulture 30,<br />
169–172.<br />
Bosio, G. (2004) Pericolo c<strong>in</strong>ese per il castagno italiano. Informatore<br />
Agrario 60, 71–72.<br />
Burckhardt, D. and J. Freuler (2000) Jump<strong>in</strong>g plant-lice (Hemiptera,<br />
Psylloidea) from sticky traps <strong>in</strong> carrot fields <strong>in</strong> Valais, <strong>Switzerland</strong>.<br />
Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen Gesellschaft 73,<br />
191–209.<br />
Burla, H. and G. Bächli (1992) Chymomyza amoena (Diptera:<br />
Drosophilidae) reared from chestnuts, acorns and fruit collected <strong>in</strong> the<br />
Canton Tic<strong>in</strong>o, <strong>Switzerland</strong>. Mitteilungen der <strong>Schweizer</strong>ischen<br />
Entomologischen Gesellschaft 65, 25–32.<br />
Büttiker, W. and Mahnert, V. (1978) Vorläufige Liste der Anoplura (Insecta<br />
der Schweiz. Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 51, 299–306<br />
CABI (2001) Crop Protection Compendium. CD-ROM. CAB International,<br />
Wall<strong>in</strong>gford, UK.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 86<br />
Dorn, K., Landau, I. and D. Cherix (1997) E<strong>in</strong>schleppung von Tap<strong>in</strong>oma<br />
melanocephalum (Formicidae) <strong>in</strong> der Schweiz. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 70, 242–243.<br />
Ebener, T., Freuler, J. and G. Meylan (1989) Frankl<strong>in</strong>iella occidentalis<br />
Pergande: 'le cas en Suisse romande'. Revue Horticole Suisse 62, 326–<br />
343.<br />
Eder, R. and R. Bauer (2003) La mouche m<strong>in</strong>euse du poireau est arrivée<br />
en Suisse. Der Gemüsebau/le Maraîcher 7/2003, 24–25.<br />
Essl, F. and W. Rabitsch (eds) (2002) Neobiota <strong>in</strong> Österreich.<br />
Umweltbundesamt, Wien.<br />
Fischer, S. (1993) Les m<strong>in</strong>euses (Lepidoptera, Yponomeutidae), <strong>in</strong>sectes<br />
ravageurs des Cupressacees d'ornement en Suisse. Revue Suisse de<br />
Viticulture, d'Arboriculture et d'Horticulture 25, 383–387.<br />
Forster, B. (2002) Auswirkungen e<strong>in</strong>gewanderter oder e<strong>in</strong>geschleppter<br />
Insekten auf die Wald- und Parkbäume der Schweiz. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 75, 135–136<br />
Fowler, S.V. (2004) Biological control of an exotic scale, Orthezia <strong>in</strong>signis<br />
Browne (Homoptera: Ortheziidae), saves the endemic gumwood tree<br />
Commidendrum robustum (Roxb.) DC. (Asteraceae) on the island of St.<br />
Helena. Biological Control 29, 367–374.<br />
Freitag A., Dorn, K. and D. Cherix (2000) First occurrence of the crazy ant<br />
Paratrech<strong>in</strong>a longicornis (Latreille) (Hym., Formicidae: Formic<strong>in</strong>ae) <strong>in</strong><br />
<strong>Switzerland</strong>. Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 73, 301–303.<br />
Geiter, O., Homma, S. and R. K<strong>in</strong>zelbach (2002) Bestandsaufnahme und<br />
Bewertung von Noezoen <strong>in</strong> Deutschland. Umweltforschungsplan des<br />
Bundesm<strong>in</strong>isteriums für Umwelt, Naturschutz und Reaktorsicherheit.<br />
Forschungsbericht 296 89 901/’1 UBA-FB 000215.<br />
Giacalone, I. and G. Lampel (1996) Pucerons (Homoptera, Aphid<strong>in</strong>a) de la<br />
région <strong>in</strong>subrique tess<strong>in</strong>oise d'orig<strong>in</strong>e subméditerranéenne,<br />
méditerranéenne, est-européenne, asiatique et américa<strong>in</strong>e. Mitteilungen<br />
der <strong>Schweizer</strong>ischen Entomologischen Gesellschaft 69, 229–260.<br />
Giacalone, I., Dioli, P. and N. Patocchi (2002) Monitoraggi faunistici alle<br />
Bolle di Magad<strong>in</strong>o (Svizzera meridionale) Eterotteri acquatici e terrestri<br />
(Insecta, Heteroptera: Gerromorpha, Nepomorpha e Geocorisae).<br />
Bollett<strong>in</strong>o della Società Tic<strong>in</strong>ese di Scienze Naturali 90, 81–92.<br />
Gómez, C. and J. Oliveras (2003) Can the Argent<strong>in</strong>e ant (L<strong>in</strong>epithema<br />
humile Mayr) replace native ants <strong>in</strong> myrmecorchory? Acta Oecologica 24,<br />
47–53.<br />
Greathead, D.J. (ed) (1976) A review of biological control <strong>in</strong> western and<br />
southern Europe. Commonwealth Institute of Biological Control. Technical<br />
Communication No 7.<br />
Grimm, K. (1986): Neu für die Schweiz: Sclerocona acutellus EV. –<br />
Mitteilungen der Entomologischen Gesellschaft Basel 36, 171–172<br />
Günthart, H. (1987) Für die Schweiz neue und wenig gesammelte<br />
Zikaden-Arten (Hom., Auchenorrhyncha). 2. Ergänzung. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 60, 83–105.<br />
Günthart, H., R. Mühlethaler and P. Lauterer (2004) Für die Schweiz neue<br />
Zikadenarten und Ergänzungen zu bereits bekannten Arten (Hemiptera<br />
Auchenorrhyncha). Mitteilungen der Entomologischen Gesellschaft Basel<br />
54, 150–160.<br />
Günther, H. and G. Schuster (1990) Checklist of the bugs of central<br />
Europe (Heteroptera). Deutsche Entomologische Zeitschrift 37, 361–396.<br />
Hächler, M., Jerm<strong>in</strong>i, M. and R. Brunetti (1998) Deux nouvelles<br />
noctuelles, ravageurs des cultures de tomate sous abri au Tess<strong>in</strong> et en<br />
Suisse romande. Revue Suisse de Viticulture, d'Arboriculture et<br />
d'Horticulture 30, 281–285<br />
Hättenschwiler, P. (2000) Typhonia beatricis sp. n., e<strong>in</strong>e möglicherweise<br />
aus dem östlichen Mittelmeerraum e<strong>in</strong>geschleppte Psychide (Lepidoptera,<br />
Psychidae) – Mitteilungen der Entomologischen Gesellschaft Basel 50, 2–<br />
17.<br />
Eichhorn, O. (1967) On methods of differentiat<strong>in</strong>g the <strong>species</strong> of the<br />
harmful white woolly aphids (Genus Dreyfusia CB. = Adelges AN.) on fir,<br />
and the consequences for forest protection Technical Bullet<strong>in</strong> of the<br />
Commonwealth Institute of Biological Control 8, 53–82.<br />
Hippe, C. and J.E. Frey (1999) Biology of the horse chestnut scale,<br />
Pulv<strong>in</strong>aria regalis Canard (Hemiptera: Coccoidea: Coccidae), <strong>in</strong><br />
<strong>Switzerland</strong>. Entomologica 33, 305–309.<br />
Hirschheydt, J.V. (1992) Der Amerikanische Nutzholzborkenkäfer<br />
Gnathotrichus materiarius (Fitch) hat die Schweiz erreicht. Mitteilungen<br />
der <strong>Schweizer</strong>ischen Entomologischen Gesellschaft 65, 33–37.<br />
Höhn, H., Höpli, H., Graf, B., Meier, J. and H. Kull (2003) Die grüne<br />
Zitrusblattlaus – auch im <strong>Schweizer</strong> Obstbau. Obst- und We<strong>in</strong>bau 139, 9–<br />
11.<br />
Hoppe, T. (1981) Vorratsschädl<strong>in</strong>ge <strong>in</strong> der Schweiz: Vorkommen und<br />
Resistenz. Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 54, 3–13.<br />
Jenk<strong>in</strong>s, J.C., Aber, J.D. and C.D. Canham (1999) Hemlock woolly adelgid<br />
impacts on community structure and N cycl<strong>in</strong>g rates <strong>in</strong> eastern hemlock<br />
forests. Canadian Journal of Forest Research 29, 630–645.<br />
Jerm<strong>in</strong>i, M., Bonavia, M., Brunetti, R., Mauri, G. and V. Cavalli (1995)<br />
Metcalfa pru<strong>in</strong>osa Say, Hyphantria cunea (Drury) et Dichelomyia<br />
oenophila Haimah, trois curiosités entomologiques ou trois nouveaux<br />
problèmes phytosanitaires pour le Tess<strong>in</strong> et la Suisse? Revue Suisse de<br />
Viticulture, d'Arboriculture et d'Horticulture 27, 57–63.<br />
Jerm<strong>in</strong>i, M., Brunetti, R. and M. Bonavia (2000) Introduzione di<br />
Neodry<strong>in</strong>us typhlocybae per il contenimento biologico di Metcalfa<br />
pru<strong>in</strong>osa: Prime esperienze <strong>in</strong> Svizzera. Sherwood – Foreste ed Alberi<br />
Oggi, Supplemento 55, 18–20.<br />
Kenis, M. and B. Forster (1998) Die Rosskastanien-M<strong>in</strong>iermotte: neu <strong>in</strong><br />
der Schweiz. Der Gartenbau 39, 16–17.<br />
Karsholt O. and J. Razowski (1996) The Lepidoptera of Europe: a<br />
distributional checklist. Stenstrup, Denmark: Apollo Books.
4 > Insects – Insecta 87<br />
Klausnitzer, B. (2004) Harmonia axyridis (Pallas, 1773) <strong>in</strong> Basel-Stadt<br />
(Coleoptera, Cocc<strong>in</strong>ellidae). Mitteilungen der Entomologischen<br />
Gesellschaft Basel 54, 115–122.<br />
Kosztarab, M. and F. Kozár (1988) Scale <strong>in</strong>sects of Central Europe.<br />
Budapest, Hungary: Akadémiai Kiadó.<br />
Kozar F., Guignard, E., Bachmann, F., Mani, E. and C. Hippe (1994) The<br />
scale <strong>in</strong>sect and whitefly <strong>species</strong> of <strong>Switzerland</strong> (Homoptera: Coccoidea<br />
and Aleyroidea). Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 67, 151–161.<br />
Kutter, H. (1981) Iridomyrmex humilis (Hym., Formicidae), Gattung und<br />
Art neu für die Schweiz. Mitteilungen der <strong>Schweizer</strong>ischen<br />
Entomologischen Gesellschaft 54, 171–172.<br />
Lampel, G. (1974) Für die Schweiz neue Blattlaus-Arten (Homoptera,<br />
Aphid<strong>in</strong>a) I Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 47, 273–306.<br />
Lampel,. G. and W. Meier (2003) Hemiptera: Sternorrhyncha – Aphid<strong>in</strong>a<br />
Volume 1. Fauna Helvetica 8.<br />
Lienhard, C. (1994) Staubläuse (Psocoptera) – ungebetene Gäste <strong>in</strong> Haus<br />
und Vorrat. Mitteilungen der Entomologischen Gesellschaft Basel 44,<br />
122–160<br />
L<strong>in</strong>der, C., Anton<strong>in</strong>, P., Mittaz, C. and R. Terrettaz (1998) Les thrips des<br />
fraisiers en Suisse romande. Especes, dynamique des populations,<br />
nuisibilite. Revue Suisse de Viticulture, d'Arboriculture et d'Horticulture<br />
30, 161–166.<br />
Liston, A.D. (1981) A provisional list of Swiss sawflies. Deutsche<br />
Entomologische Zeitschrift N.F. 28, 165–181.<br />
Lethmayer, C. and W. Rabitsch (2002) Pflanzenläuse (Sternorrhyncha). In:<br />
Essl, F. & Rabitsch, W. (eds) Neobiota <strong>in</strong> Österreich. Umweltbundesamt,<br />
Wien, pp. 316–323.<br />
Lucht, W.H. (1987) Die Käfer Mitteleuropas. Katalog. Goeke & Evers,<br />
Krefeld.<br />
Mani, E. and C. Baroffio (1997) Biologische Bekämpfung der San-José-<br />
Schildlaus im Kanton Zug mit der Schlupfwespe Encarsia perniciosi.<br />
<strong>Schweizer</strong> Zeitschrift für Obst- und We<strong>in</strong>bau 133, 392–394.<br />
Mani E., Merz, B., Brunetti, R., Schaub, L., Jerm<strong>in</strong>i, M. and F. Schwaller<br />
(1994) Zum Auftreten der beiden amerikanischen Fruchtfliegenarten<br />
Rhagoletis completa Cresson und Rhagoletis <strong>in</strong>differens Curran <strong>in</strong> der<br />
Schweiz (Diptera: Tephritidae). Mitteilungen der <strong>Schweizer</strong>ischen<br />
Entomologischen Gesellschaft 67, 177–182.<br />
Mani E., Kozar, F., Schwaller F. and C. Hippe (1997) Auftreten und<br />
Biologie der Maulbeerschildlaus Pseudaulacasois pentagona (Targioni-<br />
Tozzetti) <strong>in</strong> der Schweiz (Homoptera: Diaspididae). Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 70, 399–408.<br />
Mario, B., Jacques, D., Luigi C. and B. Roberto (2001) Update of<br />
monitor<strong>in</strong>g data of Diabrotica virgifera virgifera <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> 2001.<br />
XXI IWGO Conference, VIII Diabrotica Subgroup Meet<strong>in</strong>g, Legnaro – Padua<br />
– Venice – Italy, Oct. 27 – Nov. 3, 2001. Proceed<strong>in</strong>gs, pp. 169–173.<br />
Meier, F., R. Engesser, R., B. Forster, B. and O. Odermatt (2004)<br />
Forstschutz-Überblick 2003. Report. Eidg. Forschungsanstalt für Wald,<br />
Schnee und Landschaft, Birmensdorf. 22 pp<br />
Meier, W. Ergänzungen zur Blattlausfauna der Schweiz I (Homoptera,<br />
Aphididae). Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 45, 1–30.<br />
Meier, W. Ergänzungen zur Blattlausfauna der Schweiz II (Homoptera,<br />
Aphididae). Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 48, 405–435.<br />
Merz, B. (1991) Rhagoletis completa Cresson und Rhagoletis <strong>in</strong>differens<br />
Curran, zwei wirtschaftlich bedeutende nordamerikanische Fruchtfliegen,<br />
neu für Europa (Diptera: Tephritidae). Mitteilungen der <strong>Schweizer</strong>ischen<br />
Entomologischen Gesellschaft 64, 55–47.<br />
Merz, B., Bächli, G., Haenni, J.-P. and Y. Gonseth (eds) (1998) Diptera.<br />
Checklist. Fauna Helvetica 1, 266–267.<br />
Merz, B., Baechli, G. and J.P. Haenni (2001) Erster Nachtrag zur<br />
Checkliste der Diptera der Schweiz. 2001 Mitteilungen der<br />
Entomologischen Gesellschaft Basel 51, 110–140.<br />
Mey, E. (1988) Uebersicht über die Säugtiere-Mallophagen Europas.<br />
Angewandte Parasitologie 29, 113–126<br />
Nadig, A. and P. Thorens (1991) Liste faunistique commentée des<br />
Orthoptères de Suisse (Insecta, Orthoptera, Saltatoria). Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 64, 281–291.<br />
Neuenschwander, P. (1984) Erster Nachweis der Olivenfliege Dacus oleae<br />
(Gmel.) (Dipt. Tephritidae) <strong>in</strong> der Schweiz. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 57, 286<br />
Neumeyer, R. and B. Seifert (2005) Kommentierte Liste der frei lebenden<br />
Ameisen (Hymenoptera: Formicidae) <strong>in</strong> der Schweiz. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 78, 1–17.<br />
Noyes, J.S. (2001) Interactive catalogue of world Chalcidoidea 2001. CD-<br />
ROM Taxapad, Vancouver, Canada.<br />
Prescher, S., Moretti, M. and P. Duelli (2002) Scuttle flies (Diptera,<br />
Phoridae) <strong>in</strong> Castanea sativa forests <strong>in</strong> the southern Alps (Tic<strong>in</strong>o,<br />
<strong>Switzerland</strong>), with thirteen <strong>species</strong> new to <strong>Switzerland</strong>. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 75, 289–298.<br />
Pschorn-Walcher, H. and A. Taeger (1995) Blattwespen (Hymenoptera:<br />
Symphyta) aus Zeltfallen-Fängen im Kanton Jura. Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 68, 373–385.<br />
Rabitsch, W. (2002) Deraeocoris flavil<strong>in</strong>ea (A. Costa, 1862) erstmals <strong>in</strong><br />
Österreich festgestellt (Heteroptera, Miridae). Beiträge zur<br />
Entomofaunistik 3, 181–183.<br />
Rezbanyai-Reser, L. (1983) Über Caradr<strong>in</strong>a-Arten, <strong>in</strong>sbesondere über C.<br />
<strong>in</strong>grata Staud<strong>in</strong>ger, 1897, e<strong>in</strong>e für die Schweiz und für Mitteleuropa neue<br />
mediterrane Art (Lep., Noctuidae). Entomologische Berichte Luzern 10,<br />
99–109.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 88<br />
Rezbanyai-Reser, L. (1984) Ueber Heliothis armigera Hbn. ihr<br />
Wanderflugjahr 1983, sowie Angaben über ihr Ersche<strong>in</strong>en <strong>in</strong> der Schweiz<br />
<strong>in</strong> früheren Jahren (Lep. Noctuidae). Mitteilungen der Entomologischen<br />
Gesellschaft Basel 34, 71–91.<br />
Rezbanyai-Reser, L. (1991) Hyphantria cunea Drury, 1773, und Noctua<br />
tirrenica Bieb<strong>in</strong>ger, Speidel & Hanigk, 1983, im Südtess<strong>in</strong>, neu für die<br />
Schweiz (Lep.: Arctiidae, Noctuidae). Entomologische Berichte Luzern 26,<br />
94–96, 135–152.<br />
Rezbanyai-Reser, L. (1993) Erneut e<strong>in</strong> Heuschrecken-Fremdl<strong>in</strong>g, die<br />
Aegyptische Knarrschrecke, Anacridium aegyptium, bei Luzern,<br />
Zentralschweiz, gefunden (Saltatoria). Entomologische Berichte Luzern<br />
29, 67–68.<br />
Rezbanyai-Reser, L. (2000) Zur Nachtgrossfalterfauna der Magad<strong>in</strong>o-<br />
Ebene, 196–210m, Kanton Tess<strong>in</strong>, 1980–1995 (Lepidoptera:<br />
“Macroheterocera”). Entomologische Berichte Luzern 43, 17–179.<br />
Rezbanyai-Reser, L., Schäffer, E. and M. Hächler (1997) Platyperigea<br />
<strong>in</strong>grata (Staud<strong>in</strong>ger, 1897) zum ersten Mal <strong>in</strong> der Zentralschweiz, sowie<br />
weitere Fundangaben aus der Südwestschweiz (Lepidoptera: Noctuidae).<br />
Mitteilungen der Entomologischen Gesellschaft Basel 47, 2–8.<br />
Rezbanyai-Reser, L., Blöchl<strong>in</strong>ger, H., Hoppe, H., Schäffer, E. and J.<br />
Schmid (1998) Zur Weiterverbreitung von Eupithecia s<strong>in</strong>uosaria<br />
(Eversmann, 1848) <strong>in</strong> der Schweiz (Lepidoptera, Geometridae). Atalanta<br />
28, 309–314.<br />
Rognes K. (1997) Additions to the Swiss fauna of blowflies with an<br />
analysis of the systematic position of Calliphora stylifera (Pokorny, 1889)<br />
<strong>in</strong>clud<strong>in</strong>g a description of the female (Diptera, Calliphoridae). Mitteilungen<br />
der <strong>Schweizer</strong>ischen Entomologischen Gesellschaft 70, 63–76.<br />
Roques, A. and M. Skrzypczynska (2003) Seed-<strong>in</strong>fest<strong>in</strong>g chalcids of the<br />
genus Megastigmus Dalman, 1820 (Hymenoptera: Torymidae) native and<br />
<strong>in</strong>troduced to the West Palearctic region: taxonomy, host specificity and<br />
distribution. Journal of Natural History 37, 127–238.<br />
Sauter, W. (1983) Die Schmetterl<strong>in</strong>ge der Schweiz. 8. Nachtrag :<br />
Microlepidopteren. Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 56, 107–124.<br />
Sauter, W. (1989) Interessante neue Insektenfunde aus der Schweiz<br />
(Dipt., Lep.) Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen<br />
Gesellschaft 62, 147–149.<br />
Schaefer, H.A. (1949) Beiträge zur Kenntnis der Psylliden der Schweiz.<br />
Mitteilungen der <strong>Schweizer</strong>ischen Entomologischen Gesellschaft 22, 1–<br />
96.<br />
Schmidt, M.E. and J.E. Frey (1995) Monitor<strong>in</strong>g of the western flower<br />
thrips Frankl<strong>in</strong>iella occidentalis <strong>in</strong> greenhouses. Mededel<strong>in</strong>gen Faculteit<br />
Landbouwkundige en Toegepaste Biologische Wetenschappen,<br />
Universiteit Gent 60, 847–850.<br />
Schwenke, W. (ed) (1972) Die Forstschädl<strong>in</strong>ge Europas, I. Band. Würmer,<br />
Schnecken, Sp<strong>in</strong>nentier, Tausendfüssler und Hemimetabole Insecten.<br />
Paul Parey, Hamburg and Berl<strong>in</strong>.<br />
Schwenke, W. (ed) (1982) Die Forstschädl<strong>in</strong>ge Europas, 4. Band.<br />
Hautflüger und Zweiflüger. Paul Parey, Hamburg and Berl<strong>in</strong>.<br />
Seifert, B. (2000) Rapid range expansion <strong>in</strong> Lasius neglectus<br />
(Hymenoptera: Formicidae) – an Asian <strong>in</strong>vader swamps Europe.<br />
Mitteilungen aus dem<br />
Museum für Naturkunde <strong>in</strong> Berl<strong>in</strong>. Deutsche Entomologische Zeitschrift<br />
47, 173–179.<br />
Seifert, B. (2004) Hypoponera punctatissima (Roger) and H. schau<strong>in</strong>slandi<br />
(Emery) – tow morphologically and biologically dist<strong>in</strong>ct <strong>species</strong><br />
(Hymenoptera: Formicidae). Abhandlungen und Berichte des<br />
Naturkundemuseums Görlitz 75, 61–81.<br />
Skuhrava, M. and V. Skuhravy (1997) Gall midges (Diptera:<br />
Cecidomyiidae) of <strong>Switzerland</strong>. Mitteilungen der <strong>Schweizer</strong>ischen<br />
Entomologischen Gesellschaft 70, 133–176.<br />
Stary, P. (1995) Natural enemy spectrum of Aphis spiraephaga (Hom.,<br />
Aphididae), an exotic immigrant aphid <strong>in</strong> Central Europe. Entomophaga<br />
40, 29–34.<br />
Thorens, P. and A. Nadig (1997) Atlas de distribution des orthopteres de<br />
Suisse: sauterelles, grillons, criquets (Orthoptera), mante religieuse<br />
(Mantodea). Centre Suisse de Cartographie de la Faune, Neuchâtel.<br />
Vernier, R. (1995) Isodontia mexicana (Sauss.), un Sphec<strong>in</strong>i américa<strong>in</strong><br />
naturalisé en Suisse (Hymenoptera, Sphecidae). Mitteilungen der<br />
<strong>Schweizer</strong>ischen Entomologischen Gesellschaft 68, 169–177.<br />
Vernier, R. (2000) Isodontia mexicana (Sauss.) (Hymenoptera, Sphecidae)<br />
est apparu dans le Bass<strong>in</strong> lémanique. Bullet<strong>in</strong> Roman d’Entomologie 18,<br />
143–145.<br />
Villa, M., Tescari, G. and S.J. Taylor (2001) Nuovi dati sulla presenza <strong>in</strong><br />
Italia di Leptoglossus occidentalis (Heteroptera Coreidae). Bollet<strong>in</strong>o della<br />
Societa Entomologica Italiana 133, 103–112.<br />
Watson, G.W., Voegtl<strong>in</strong>, D.J. Murphy, S.T. and R.G. Foottit (1999)<br />
Biogeography of the C<strong>in</strong>ara cupressi complex (Hemiptera: Aphididae) on<br />
Cupressaceae, with description of a pest <strong>species</strong> <strong>in</strong>troduced <strong>in</strong>to Africa.<br />
Bullet<strong>in</strong> of Entomological Research 89, 271–283.<br />
Whitebread, S. (1997) Platyperigea <strong>in</strong>grata (Staud<strong>in</strong>ger, 1897): Die Raupe<br />
als unerwünschter Gast im Hause (Lepidoptera: Noctuidae). Mitteilungen<br />
der Entomologischen Gesellschaft Basel 47, 9–11.
4 > Insects – Insecta 89<br />
Tab. 4.1 > Established <strong>alien</strong> Insects <strong>in</strong> <strong>Switzerland</strong>: Coleoptera.<br />
Species Habitat – life traits Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
COLEOPTERA<br />
Anobiidae<br />
Lasioderma serricorne (F.) Stored products, especially tobacco Tropics and subtropics C.Besuchet, pers. comm.<br />
Stegobium paniceum (L.) Stored products, very polyphagous Cosmopolitan Lucht, 1987<br />
Oligomerus ptil<strong>in</strong>oides (Wollaston) Feeds on wood products Mediterranean region, possibly C. Besuchet, pers. comm.<br />
native<br />
Anthicidae<br />
Stricticomus tobias Marseul Feeds on rotten plant tissue Asia M<strong>in</strong>or, Central Asia, India C. Besuchet, pers. comm.<br />
Apionidae<br />
Alocentron curvirostre (Gyllenhal) In stems of Alcea rosea L. Asia, Middle East C. Besuchet, pers. comm.<br />
Aspidapion validum (Germar) In fruits of Alcea rosea Asia, Middle East C. Besuchet, pers. comm.<br />
Rhopalapion longirostre (Olivier) In seeds of Alcea rosea Asia, Middle East C. Besuchet, pers. comm.<br />
Bostrichidae<br />
Rhyzoperta dom<strong>in</strong>ica (F.) Stored products, ma<strong>in</strong>ly cereals Tropics and subtropics C. Besuchet, pers. comm.<br />
Bruchidae<br />
Acanthoscelides obtectus (Say) In legum<strong>in</strong>ous seeds, ma<strong>in</strong>ly beans South and Central America Lucht, 1987<br />
Acanthoscelides pallidipennis (Motschulsky) In seeds of Amorpha fruticosa North America C. Besuchet, pers. comm.<br />
Bruchus pisorum (L.) In peas North America or Near East Lucht, 1987<br />
Callosobruchus ch<strong>in</strong>ensis (L.) In legum<strong>in</strong>ous seeds East Asia C. Besuchet, pers. comm.<br />
Buprestidae<br />
Agrilus guer<strong>in</strong>i Lacordaire Xylophagous, on Salix Eastern Europe, Russia? Barbalat and Wermel<strong>in</strong>ger,<br />
1996<br />
Carabidae<br />
Perigona nigriceps (Dejean) Predator, <strong>in</strong> various environments South Asia Lucht, 1987<br />
Cerambycidae<br />
Gracilia m<strong>in</strong>uta (F.) Xylophagous, polyphagous, often found <strong>in</strong> wicker Mediterranean region Lucht, 1987<br />
Nathrius brevipennis (Mulsant) Xylophagous, polyphagous, often found <strong>in</strong> wicker Mediterranean region Lucht, 1987<br />
Neoclytus acum<strong>in</strong>atus (F) Xylophagous, on Frax<strong>in</strong>us, Tic<strong>in</strong>o North America C. Besuchet, pers. comm.<br />
Phymatodes lividus (Rossi) Xylophagous, on broadleaved trees Mediterranean region C. Besuchet, pers. comm.<br />
Xylotrechus stebb<strong>in</strong>gi Gahan Xylophagous, polyphagous, on aspen <strong>in</strong> Tic<strong>in</strong>o Himalayas C. Besuchet, pers. comm.<br />
Cerylonidae<br />
Murmidius ovalis (Beck) Stored products, especially mouldy cereals, hay, Cosmopolitan C. Besuchet, pers. comm.<br />
etc.<br />
Chrysomelidae<br />
Diabrotica virgifera virgifera LeConte Pest on Maize, root feeder, Tic<strong>in</strong>o Mexico, Central America Mario et al., 2001<br />
Lept<strong>in</strong>otarsa deceml<strong>in</strong>eata (Say) Defoliator, pest on Solanaceae North America Lucht, 1987<br />
Cleridae<br />
Korynetes caeruleus (De Geer) Predator on anobiids <strong>in</strong> wood Cosmopolitan Lucht, 1987<br />
Necrobia ruficollis (F.) Domestic, <strong>in</strong>sect predator Cosmopolitan Lucht, 1987<br />
Necrobia rufipes (De Geer) Domestic, <strong>in</strong> stored products, also predator Tropics, subtropics Lucht, 1987<br />
Necrobia violacea (L.) Ma<strong>in</strong>ly domestic, also on cadavers Cosmopolitan Lucht, 1987<br />
Tarsostenus univittatus (Rossi) Predator <strong>in</strong> wood, found <strong>in</strong> Genève only Cosmopolitan C. Besuchet, pers. comm.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 90<br />
Species<br />
Habitat – life traits Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
Cocc<strong>in</strong>ellidae<br />
Harmonia axyridis (Pallas) Polyphagous predator (perhaps not yet established) East Asia Klausnitzer, 2004<br />
Rodolia card<strong>in</strong>alis (Mulsant)<br />
Colydiidae<br />
Predator of scale <strong>in</strong>sects, released and established<br />
<strong>in</strong> Tic<strong>in</strong>o<br />
Australia C. Besuchet, pers. comm.<br />
Myrmecoxenus vaporariorum Guer<strong>in</strong>-<br />
Meneville<br />
Cryptophagidae<br />
In greenhouses, manure, compost, etc. Unclear Lucht, 1987<br />
Atomaria lewisi Reitter In decay<strong>in</strong>g plant material East Asia Lucht, 1987<br />
Caenoscelis subdeplanata Brisout de<br />
Barneville<br />
Cucujidae<br />
In decay<strong>in</strong>g wood and plant material North America C. Besuchet, pers. comm.<br />
Cryptolestes ferrug<strong>in</strong>eus (Stephens) Stored products, ma<strong>in</strong>ly cereal gra<strong>in</strong>s Cosmopolitan Lucht, 1987<br />
Cryptolestes spartii (Curtis) Stored products Cosmopolitan C. Besuchet, pers. comm.<br />
Cryptolestes turcicus (Grouvelle)<br />
Curculionidae<br />
Domestic, on plant products Perhaps Turkey C. Besuchet, pers. comm.<br />
Pentarthrum huttoni Wollaston Decay<strong>in</strong>g wood South-western Europe (native?) C. Besuchet, pers. comm.<br />
Sitophilus granarius (L.) Stored products India Lucht, 1987<br />
Sitophilus oryzae (L.) Stored products Asia Lucht, 1987<br />
Sitophilus zeamais Motschulsky<br />
Dermestidae<br />
Stored products Asia C. Besuchet, pers. comm.<br />
Anthrenus festivus Rosenhauer Domestic Mediterranean region C. Besuchet, pers. comm.<br />
Anthrenus flavipes LeConte Domestic, feeds on furnitures, fabrics, etc. Cosmopolitan (Mediterranean<br />
region?)<br />
C. Besuchet, pers. comm.<br />
Attagenus brunneus Faldermann Domestic Cosmopolitan (native?) C. Besuchet, pers. comm.<br />
Attagenus quadrimaculatus Kraatz Domestic Cosmopolitan (native?) C. Besuchet, pers. comm.<br />
Attagenus rossi Ganglbauer Domestic Cosmopolitan (native?) C. Besuchet, pers. comm.<br />
Attagenus smirnovi Zhantiev Domestic Cosmopolitan (native?) C. Besuchet, pers. comm.<br />
Attagenus unicolor (Brahm) Domestic, feeds ma<strong>in</strong>ly on fabrics Africa Lucht, 1987<br />
Dermester ater DeGeer Domestic, on animal products, fabrics, etc. Cosmopolitan (southern Europe?) C. Besuchet, pers. comm.<br />
Dermestes maculatus DeGeer Domestic, on animal products Cosmopolitan (North America?) Lucht, 1987<br />
Dermestes peruvianus La Porte de C. Domestic, on animal products, fabrics, etc. Central and South America C. Besuchet, pers. comm.<br />
Reesa vespulae (Milliron) Domestic and <strong>in</strong> museum collections North America C. Besuchet, pers. comm.<br />
Trogoderma angustum (Solier) Domestic and <strong>in</strong> museum collections South America C. Besuchet, pers. comm.<br />
Trogoderma glabrum Herbst Domestic and <strong>in</strong> nests of solitary wasps Cosmopolitan (native?) Lucht, 1987<br />
Trogoderma granarium Everts Stored products, especially cereals India C. Besuchet, pers. comm.<br />
Trogoderma versicolor (Creutzer)<br />
Histeridae<br />
Domestic and <strong>in</strong> museum collections Cosmopolitan Lucht, 1987<br />
Carc<strong>in</strong>ops pumilio Erichson<br />
Hydrophilidae<br />
Predator on Diptera Cosmopolitan Lucht, 1987<br />
Cercyon lam<strong>in</strong>atus Sharp In various humid environments East Asia Lucht, 1987<br />
Cryptopleurum subtile Sharp<br />
Languriidae<br />
In various humid environments East Asia Lucht, 1987<br />
Cryptophilus <strong>in</strong>teger (Heer) On decay<strong>in</strong>g plant material Cosmopolitan Lucht, 1987
4 > Insects – Insecta 91<br />
Species<br />
Habitat – life traits Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
Latridiidae<br />
Adistemia watsoni (Wollaston) Feeds on fungus, found <strong>in</strong> herbarium Cosmopolitan Lucht, 1987<br />
Corticaria ferrug<strong>in</strong>ea Marsham On fungus, on decay<strong>in</strong>g plant material Cosmopolitan C. Besuchet, pers. comm.<br />
Corticaria fulva (Comolli) On fungus, on decay<strong>in</strong>g plant material Cosmopolitan Lucht, 1987<br />
Corticaria pubescens Gyllenhal On fungus, on decay<strong>in</strong>g plant material Cosmopolitan Lucht, 1987<br />
Corticaria serrata Paykull On fungus, on decay<strong>in</strong>g plant material Cosmopolitan Lucht, 1987<br />
Dienerella filum (Aubé) On fungus, on decay<strong>in</strong>g plant material Central America Lucht, 1987<br />
Lathridius m<strong>in</strong>utus (L.) On fungus, on decay<strong>in</strong>g plant material Cosmopolitan Lucht, 1987<br />
Migneauxia orientalis Reitter On fungus, on decay<strong>in</strong>g plant material Mediterranean region C. Besuchet, pers. comm.<br />
Stephostetus (= Aridius) bifasciatus (Reitter) Feeds on fungus <strong>in</strong> various environments Australia C. Besuchet, pers. comm.<br />
Stephostetus nodifer (Westwood) Feeds on fungus <strong>in</strong> various environments New Zealand Lucht, 1987<br />
Thes bergrothi (Reitter)<br />
Lyctidae<br />
On fungus, on decay<strong>in</strong>g plant material North-eastern Europe C. Besuchet, pers. comm.<br />
Lyctus africanus Lesne Domestic, <strong>in</strong> wood Africa C. Besuchet, pers. comm.<br />
Lyctus brunneus (Stephens) Domestic, <strong>in</strong> tropical wood South-east Asia Lucht, 1987<br />
Lyctus cavicollis Le Conte<br />
Merophysiidae<br />
Domestic, <strong>in</strong> wood North America C. Besuchet, pers. comm.<br />
Holoparamecus caularum (Aubé)<br />
Mycetophagidae<br />
On fungus, on decay<strong>in</strong>g plant material Cosmopolitan C. Besuchet, pers. comm.<br />
Berg<strong>in</strong>us tamarisci Wollaston Found on p<strong>in</strong>e trees <strong>in</strong> <strong>Switzerland</strong> Canary Islands C. Besuchet, pers. comm.<br />
Litargus balteatus Le Conte On fungus, on decay<strong>in</strong>g plants, e.g. cereals North America C. Besuchet, pers. comm.<br />
Typhaea stercorea (L.)<br />
Nitidulidae<br />
On fungus, on decay<strong>in</strong>g plants, e.g. cereals North America Lucht, 1987<br />
Carpophilus dimidiatus (F.) On stored products and cultivated fields, ma<strong>in</strong>ly<br />
cereals<br />
North America C. Besuchet, pers. comm.<br />
Carpophilus hemipterus (L.) Feeds on fruits, dry fruits, cereals Cosmopolitan Lucht, 1987<br />
Carpophilus marg<strong>in</strong>ellus Motschulsky Ma<strong>in</strong>ly domestic, on cereals South-east Asia C. Besuchet, pers. comm.<br />
Carpophilus quadrisignatus Erichson Feeds on dry fruits Probably America C. Besuchet, pers. comm.<br />
Glischrochilus fasciatus (Olivier) Feeds on vegetables, fruits, etc. North America C. Besuchet, pers. comm.<br />
Glischrochilus quadrisignatus (Say) Feeds on vegetables, fruits, etc. North and Central America C. Besuchet, pers. comm.<br />
Urophorus rubripennis (Heer)<br />
Orthoperidae<br />
Under oak bark and on Apiaceae Perhaps Mediterranean region C. Besuchet, pers. comm.<br />
Orthoperus aequalis Sharp<br />
Ostomidae<br />
In compost, <strong>in</strong> Tic<strong>in</strong>o Hawaii C. Besuchet, pers. comm.<br />
Tenebroides mauritanicus (L.)<br />
Ptiliidae<br />
On stored products, especially cereals Africa Lucht, 1987<br />
Acrotrichis <strong>in</strong>sularis (Maekl<strong>in</strong>) In organic matters, from Vaud and Weissenste<strong>in</strong> North America C. Besuchet, pers. comm.<br />
Acrotrichis sanctaehelenae Johnson<br />
Ptilodactylidae<br />
In organic matters, Tic<strong>in</strong>o St Helena, Africa? C. Besuchet, pers. comm.<br />
Ptilodactyla exotica Chap<strong>in</strong><br />
Pt<strong>in</strong>idae<br />
Domestic, <strong>in</strong>door plants North America C. Besuchet, pers. comm.<br />
Gibbium psylloides (Czemp<strong>in</strong>ski) Domestic, on stored products Cosmopolitan Lucht, 1987<br />
Niptus hololeucus (Faldermann) Domestic, feeds on fabrics Asia M<strong>in</strong>or, southern Russia Lucht, 1987
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 92<br />
Species<br />
Habitat – life traits Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
Pt<strong>in</strong>us tectus Boieldieu Domestic, on stored products Australia, New Zealand Lucht, 1987<br />
Epauloecus (=Tipnus) unicolor (Piller & Mitt.) In barns, cowshed, animal burrows, etc<br />
Scolytidae<br />
Cosmopolitan Lucht, 1987<br />
Gnathotrichus materiarius (Fitch) Xylophagous, on conifers North America Hirschheydt, 1992<br />
Phloeos<strong>in</strong>us aubei (Perris) Xylophagous, on Cupressaceae Mediterranean region Lucht, 1987<br />
Tripodendron laeve Eggers Xylophagous, on Picea East Asia, Scand<strong>in</strong>avia C. Besuchet, pers. comm.<br />
Xylebor<strong>in</strong>us alni (Niisima) Xylophagous, on broadleaved trees East Asia C. Besuchet, pers. comm.<br />
Xyleborus punctulatus Kurentzov Xylophagous, on broadleaved trees Siberia C. Besuchet, pers. comm.<br />
Xylosandrus germanus (Blandford)<br />
Silvanidae<br />
Xylophagous, polyphagous East Asia Lucht, 1987<br />
Ahasverus advena (Waltl) Feeds on fungus on rotten stored products South America Lucht, 1987<br />
Oryzaephilus mercator (Fauvel) On stored products, polyphagous Tropics C. Besuchet, pers. comm.<br />
Oryzaephilus sur<strong>in</strong>amensis (L.)<br />
Staphyl<strong>in</strong>idae<br />
On stored products, polyphagous Cosmopolitan Lucht, 1987<br />
Acrotona pseudotenera (Cameron) Mouldy hay East Asia C. Besuchet, pers. comm.<br />
Carpelimus zealandicus (Sharp) Sandy banks New Zealand C. Besuchet, pers. comm.<br />
Edaphus beszedesi Reitter Compost, rott<strong>in</strong>g plant material Pehaps Mediterranean region Lucht, 1987<br />
Leptoplectus remyi Jeannel Unclear, found <strong>in</strong> Tic<strong>in</strong>o Asia C. Besuchet, pers. comm.<br />
Lithocharis nigriceps (Kraatz) Compost, rott<strong>in</strong>g plant material Asia Lucht, 1987<br />
Micropeplus marietti Jaquel<strong>in</strong> Du Val Waste land, fallow land Southern Europe, Caucasus C. Besuchet, pers. comm.<br />
Oligota parva Kraatz Domestic, <strong>in</strong> compost South America Lucht, 1987<br />
Oxytelus migrator (Fauvel) In compost, stable litter, etc. South East Asia C. Besuchet, pers. comm.<br />
Paraphloeostiba gayndahensis MacLeay Ferment<strong>in</strong>g plant matters Australia C. Besuchet, pers. comm.<br />
Philonthus parcus Sharp In compost, stable litter, ferment<strong>in</strong>g plant material,<br />
etc.<br />
East Asia C. Besuchet, pers. comm.<br />
Philonthus rectangulus Sharp In decompos<strong>in</strong>g matters East Asia Lucht, 1987<br />
Philonthus sp<strong>in</strong>ipes Sharp In stable litter, cadavers, etc. East Asia C. Besuchet, pers. comm.<br />
Thecturota marchii (Dodero) Waste land, compost Southern Europe Lucht, 1987<br />
Trichiusa immigrata Lohse<br />
Tenebrionidae<br />
In compost and manure North America C. Besuchet, pers. comm.<br />
Alphitobius diaper<strong>in</strong>us (Panzer) In stored products, polyphagous Tropics C. Besuchet, pers. comm.<br />
Alphitophagus bifasciatus (Say) Ma<strong>in</strong>ly domestic, In rotten fruits Cosmopolitan C. Besuchet, pers. comm.<br />
Gnatocerus cornutus (F.) Stored products Central America Lucht, 1987<br />
Latheticus oryzae Waterhouse Stored products, cereals India C. Besuchet, pers. comm.<br />
Tenebrio molitor L., polyphagous Stored products Cosmopolitan Lucht, 1987<br />
Tribolium castaneum (Herbst) Stored products Cosmopolitan Lucht, 1987<br />
Tribolium confusum Jacquel<strong>in</strong> du Val Stored products Perhaps America, cosmopolitan Lucht, 1987<br />
Tribolium destructor Uyttenboogaart Stored products South America C. Besuchet, pers. comm.<br />
Tribolium madens Charpentier Stored products Cosmopolitan C. Besuchet, pers. comm.
4 > Insects – Insecta 93<br />
Tab. 4.2 > Established <strong>alien</strong> <strong>in</strong>sects <strong>in</strong> <strong>Switzerland</strong>: Lepidoptera.<br />
Species Habita – Life traits Orig<strong>in</strong> References for <strong>Switzerland</strong><br />
LEPIDOPTERA<br />
Arctiidae<br />
Hyphantria cunea (Drury) Polyphagous defoliator, <strong>in</strong> Tic<strong>in</strong>o North America Rezbanyai-Reser, 1991; Jerm<strong>in</strong>i<br />
et al., 1995<br />
Gelechiidae<br />
Scrobipalpa ocellatella (Boyd) Defoliator/ borer on Chenopodiaceae, particularly on beet Southern Europe Karsholt and Razowski, 1996;<br />
CABI, 2001<br />
Sitotroga cerealella (Olivier) On stored products North America CABI, 2001<br />
Geometridae<br />
Eupithecia s<strong>in</strong>uosaria Eversmann On Chenopodium (not sure if established <strong>in</strong> <strong>Switzerland</strong>) Eastern Europe Rezbanyai-Reser et al., 1998<br />
Gracillariidae<br />
Cameraria ohridella Deschka & Dimic Leaf m<strong>in</strong>er on Aesculus Unknown Kenis and Förster, 1998<br />
Caloptilia azaleella (Brants) Leaf m<strong>in</strong>er on Rhododendron, <strong>in</strong> greenhouses EastAsia CSCF, unpublished list<br />
Parectopa rob<strong>in</strong>iella Clemens Leaf m<strong>in</strong>er on Rob<strong>in</strong>ia North America Sauter, 1983<br />
Phyllonorycter leucographella (Zeller) Leaf m<strong>in</strong>er on Pyracantha and Crataegus East Mediterranean Region Sauter, 1983<br />
P. platani (Staud<strong>in</strong>ger) Leaf m<strong>in</strong>er on Platanus Balkans, Asia M<strong>in</strong>or M. Kenis, personal observation<br />
P. rob<strong>in</strong>iella (Clemens) Leaf m<strong>in</strong>er on Rob<strong>in</strong>ia North America M. Kenis, personal observation<br />
Lycaenidae<br />
Cacyreus marshalli (Butler) On Pelargonium, <strong>in</strong> Tic<strong>in</strong>o South Africa Y. Gonzeth, pers. comm.<br />
Noctuidae<br />
Caradr<strong>in</strong>a <strong>in</strong>grata Staud<strong>in</strong>ger Defoliator (not sure if established) East Mediterranean Region Rezbanyai-Reser L. 1983,<br />
Withebread, 1997<br />
Chrysodeixis chalcites Esper Vegetables, <strong>in</strong> glasshouses, migrant (not sure if established)<br />
Helicoverpa armigera (Hübner) Polyphagous defoliator, ma<strong>in</strong>ly <strong>in</strong> glasshouses (not sure if<br />
established)<br />
Mediterranean Region? Hächler et al., 1998<br />
Africa? Rezbanyai-Reser L. 1984,<br />
Hächler et al., 1998<br />
Sed<strong>in</strong>a buettneri (Her<strong>in</strong>g) Defoliator, ma<strong>in</strong>ly on Carex Siberia Blattner (1959)<br />
Psychidae<br />
Typhonia beatricis Hättenschwiler Polyphagous, <strong>in</strong> particular on mosses Eastern Mediterranean Hättenschwiler (2000)<br />
Pyralidae<br />
Cadra cautella (Walker) On stored products Africa Hoppe (1981)<br />
Ephestia elutella (Hübner) On stored products Cosmopolitan Hoppe (1981)<br />
E. kuehniella (Zeller) On stored products North and Central America Hoppe (1981)<br />
Plodia <strong>in</strong>terpunctella (Hübner) On stored products Cosmopolitan Hoppe (1981)<br />
Sclerocona acutella (Eversmann) On Phragmites Siberia Grimm (1986)<br />
Saturniidae<br />
Samia cynthia walkeri (C. and R. Felder) Defoliator on Ailanthus East Asia CSCF, unpublished list<br />
T<strong>in</strong>eidae<br />
Monopis crocicapitella (Clemens) On fabrics Cosmopolitan CSCF, unpublished list<br />
Opogona sacchari (Bojer) Polyphagous pest <strong>in</strong> glasshouses Africa CABI, 2001<br />
Tortricidae<br />
Cydia molesta (Busck) Orchard pest, on Rosaceae East Asia CABI, 2001<br />
Yponomeutidae<br />
Argyresthia thuiella (Packard) Leaf m<strong>in</strong>er on Cupressaceae North America Fischer (1993)
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 94<br />
Tab. 4.3 > Established <strong>alien</strong> <strong>in</strong>sects <strong>in</strong> <strong>Switzerland</strong>: Hymenoptera.<br />
Species Habitat – Life traits Orig<strong>in</strong> References for <strong>Switzerland</strong><br />
HYMENOPTERA<br />
Symphyta<br />
Siricidae<br />
Sirex cyaneus Fabricius Feeds on conifer trunks (ma<strong>in</strong>ly Abies) North America Pschorn-Walcher and Taeger, 1995<br />
Tenthred<strong>in</strong>idae<br />
Nematus tibialis Newman Defoliator on Rob<strong>in</strong>ia North America Liston, 1981, Pschorn-Walcher and<br />
Taeger, 1995<br />
Apocrita<br />
Aphel<strong>in</strong>idae<br />
Aphel<strong>in</strong>us mali (Haldeman) Parasitoid of Eriosoma lanigerum (Hausmann), <strong>in</strong>troduced<br />
<strong>in</strong> <strong>Switzerland</strong><br />
North America Greathead, 1976; Noyes, 2002<br />
Aphytis proclia (Walker) Parasitoid of scale <strong>in</strong>sects, <strong>in</strong>troduced <strong>in</strong> Italy Asia Greathead, 1976; Noyes, 2002<br />
Encarsia berlesei (Howard)<br />
Encarsia formosa Gahan<br />
Encarsia lounshuryi (Berlese & Paoli)<br />
Encarsia perniciosi (Tower)<br />
Braconidea<br />
Parasitoid of Pseudaulacaspis pentagona (Targioni-<br />
Tozzetti), <strong>in</strong>troduced <strong>in</strong> Italy<br />
Biocontrol agent aga<strong>in</strong>st whiteflies. Only <strong>in</strong> greenhouses<br />
Parasitoid of Chrysomphalus dictyospermi (Morgan),<br />
<strong>in</strong>troduced <strong>in</strong> Italy<br />
Parasitoid of San José scale, <strong>in</strong>troduced <strong>in</strong> <strong>Switzerland</strong><br />
Aphidius colemani Viereck Biocontrol agent aga<strong>in</strong>st aphids <strong>in</strong> greenhouses India<br />
Dry<strong>in</strong>idae<br />
East Asia<br />
South and Central America<br />
Madeira<br />
Probably Asia<br />
Mani et al., 1997<br />
Greathead, 1976; Noyes, 2002<br />
Greathead, 1976; Noyes, 2002<br />
Mani and Baroffio, 1997<br />
Neodry<strong>in</strong>us typhlocybae (Ashmead) Parasitoid of Metcalfa pru<strong>in</strong>osa Say, <strong>in</strong>troduced <strong>in</strong> Tic<strong>in</strong>o North America Jerm<strong>in</strong>i et al., 2000<br />
Encyrtidae<br />
Metaphycus helvolus (Compere) Biocontrol agent aga<strong>in</strong>st scale <strong>in</strong>sects. Only <strong>in</strong> greenhouses<br />
Ooencyrtus kuvanae (Howard) Parasitoid of Lymantria dispar L., <strong>in</strong>troduced <strong>in</strong> Europe.<br />
Not reported from <strong>Switzerland</strong> but from all adjacent<br />
countries<br />
Formicidae<br />
South Africa Noyes, 2002<br />
East Asia Greathead, 1976; Noyes, 2002<br />
Hypoponera schau<strong>in</strong>slandi (Emery) Antropophilic, <strong>in</strong> greenhouses or other heated build<strong>in</strong>gs. Unclear, tropics Neumeyer & Seifert, 2005<br />
L<strong>in</strong>epithema humile (Mayr) Various habitats <strong>in</strong>doors and outdoors, perhaps not<br />
established <strong>in</strong> <strong>Switzerland</strong><br />
South America Kutter, 1981<br />
Monomorium pharaonis (L.) On stored products, antropophilic, ma<strong>in</strong>ly <strong>in</strong>doors South Asia Freitag et al., 2000<br />
Paratrech<strong>in</strong>a longicornis (Latreille)<br />
Tap<strong>in</strong>oma melanocephalum (F.)<br />
Sphecidae<br />
Isodontia mexicana (Saussure)<br />
Sceliphron curvatum (F. Smith)<br />
Torymidae<br />
Omnivorous, antropophilic, found <strong>in</strong> Zürich airport <strong>in</strong><br />
1999, perhaps not established<br />
On stored products, antropophilic, <strong>in</strong>doors only<br />
Predatory wasp, feeds on criquets, In Tic<strong>in</strong>o<br />
Lemanic region<br />
Predatory wasp, anthropophilic<br />
Old world tropics<br />
Cosmopolitan, tropics<br />
Freitag et al., 2000<br />
Dorn et. al., 1997<br />
and North America Vernier, 1995, 2000<br />
Asia<br />
Gonseth & al. 2001<br />
Megastigmus spermotrophus Wachtl Feeds on seeds of Pseudotsuga North America Roques and Skrzypczynska, 2003<br />
Trichogrammatidae<br />
Trichogramma brassicae Bezdenko Biocontrol agent aga<strong>in</strong>st Lepidoptera Eastern Europe Noyes, 2002
4 > Insects – Insecta 95<br />
Tab. 4.4 > Established <strong>alien</strong> <strong>in</strong>sects <strong>in</strong> <strong>Switzerland</strong>: Diptera.<br />
Species<br />
DIPTERA<br />
Agromyzidae<br />
Liriomyza huidobrensis (Blanchard)<br />
Liriomyza trifolii (Burgess)<br />
Napomyza gymnostoma (Loew)<br />
Calliphoridae<br />
Chrysomya albiceps (Wiedemann)<br />
Cecidomyiidae<br />
Rhopalomyia chrysanthemi (Ahlberg)<br />
Culicidae<br />
Habitat – Life traits Orig<strong>in</strong> References for <strong>Switzerland</strong><br />
Polyphagous leaf m<strong>in</strong>er, pest <strong>in</strong> glasshouses<br />
Polyphagous leaf m<strong>in</strong>er, pest <strong>in</strong> glasshouses <strong>in</strong><br />
Europe, particularly on Chrysanthemum<br />
Leaf m<strong>in</strong>er on Allium spp., especially onion and<br />
leek.<br />
On cadavers<br />
Pest on Chrysanthemum<br />
Aedes albopictus (Skuse) Human nuisance through its bites, and potential<br />
vector of various diseases<br />
Drosophilidae<br />
Chymomyza amoena (Loew)<br />
Drosophila curvisp<strong>in</strong>a Watabe & Toda<br />
Milichiidae<br />
Fruits of various broadleaved trees (apple, walnut,<br />
plum, etc.)<br />
Fruits<br />
Central and South America<br />
North America<br />
Unclear, perhaps <strong>in</strong>digenous<br />
CABI, 2001<br />
CABI, 2001<br />
Eder and Baur (2003)<br />
Cosmopolitan Rognes, 1997<br />
North America Skuhrava and Skuhravi 1997<br />
South-east Asia Unpublished <strong>in</strong>formation<br />
North America<br />
East Asia<br />
Burla and Bächli, 1992<br />
Bächli et al., 2002<br />
Desmometopa varipalpis Malloch Saprophagous/coprophagous Probably Cosmopolitan Merz et al., 2001<br />
Muscidae<br />
Hydrotaea aenescens (Wiedemann) On human or animal cadavers North America Merz et al., 2001<br />
Phoridae<br />
Dohrniphora cornuta (Bigot) Saprophagous, sometimes carnivorous Tropical, Cosmopolitan Prescher et al., 2002<br />
Stratiomyidae<br />
Hermetia illucens (L.)<br />
Tephritidae<br />
Bactrocera oleae (Gmel.)<br />
Ceratitis capitata (Wiedemann)<br />
Rhagoletis c<strong>in</strong>gulata Curran<br />
Rhagoletis completa Cresson<br />
Ulidiidae<br />
Saprophagous, very abundant <strong>in</strong> Tic<strong>in</strong>o.<br />
Fruit fly, on olive, <strong>in</strong> Tic<strong>in</strong>o<br />
On various fruits, e.g. peach, apricot, peer, etc.<br />
Fruit fly, on cherry. First determ<strong>in</strong>ed as R. <strong>in</strong>differens<br />
Curran (B. Merz, pers. comm.)<br />
Fruit fly, on walnut<br />
North and South America ,<br />
Africa<br />
Mediterranean region<br />
Africa<br />
North America<br />
North America<br />
Sauter, 1989<br />
Neuenschwander, 1984<br />
CABI, 2001<br />
Merz 1991, Mani et al., 1994<br />
Merz 1991, Mani et al, 1994<br />
Euxesta pechumani Curran In carrion and dung, <strong>in</strong> Tic<strong>in</strong>o North America B. Merz., pers. comm.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 96<br />
Tab. 4.5 > Established <strong>alien</strong> <strong>in</strong>sects <strong>in</strong> <strong>Switzerland</strong>: Hemiptera.<br />
Species<br />
Host plant Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
HEMIPTERA<br />
Sternorrhyncha<br />
APHIDINA<br />
Adelgidae<br />
Dreyfusia nordmannianae (Eckste<strong>in</strong>) On Abies Caucasus Eichhorn, 1967<br />
Dreyfusia prelli Grosmann On Abies Caucasus Eichhorn, 1967<br />
Eop<strong>in</strong>eus strobus (Hartig) On P<strong>in</strong>us strobus L., Picea spp. Not recorded but<br />
probably <strong>in</strong> <strong>Switzerland</strong><br />
North America Schwenke, 1972<br />
Gilletteella cooleyi (Gillette)<br />
Aphididae<br />
On Picea and Pseudotsuga North America Forster, 2002<br />
Acyrthosiphon caraganae (Choldokovsky) On Caragana and other Fabaceae Siberia CSCF/Lampel, unpublished list<br />
Aphis forbesi Weed On strawberry North America Meier, 1975<br />
Aphis gossypii Glover Polyphagous, ma<strong>in</strong>ly Cucurbitaceae and Malva- Cosmopolitan, tropical CABI, 2001<br />
ceae, <strong>in</strong> greenhouses <strong>in</strong> Central Europe<br />
regions<br />
Aphis spiraecola Patch Polyphagous, e.g. Citrus, apple East Asia Hohn et al., 2003<br />
Aphis spiraephaga F.P. Müller On Spiraea Central Asia Stary, 1995<br />
Appendiseta rob<strong>in</strong>iae (Gillette) On Rob<strong>in</strong>ia North America Lethmayer and Rabitsch, 2002<br />
Aulacorthum circumflexum (Buckton) Polyphagous, <strong>in</strong> greenhouses South East Asia CSCF/Lampel, unpublished list<br />
Elatobium abiet<strong>in</strong>um (Walker) On Picea North America CABI, 2001<br />
Idiopterus nephrelepidis Davis On ferns, mostly <strong>in</strong>doors Neotropics CSCF/Lampel, unpublished list<br />
Ill<strong>in</strong>oia azaleae (Mason) On Rhododendron and other Ericaceae North America CSCF/Lampel, unpublished list<br />
Ill<strong>in</strong>oia lambersi (Mac Gillivray) On Rhododendron and Kalmia North America CSCF/Lampel, unpublished list<br />
Impatient<strong>in</strong>um asiaticum Nevsky On Impatiens Central Asia CSCF/Lampel, unpublished list<br />
Macrosiphoniella sanborni (Gillette) On Chrysanthemum East Asia Meier, 1972<br />
Macrosiphum albifrons Essig On Lup<strong>in</strong>us North America CSCF/Lampel, unpublished list<br />
Macrosiphum euphorbiae (Thomas) Polypgagous, on vegetables North America Derron and Goy, 1995<br />
Megoura lespedezae (Essig & Kuwana) On Lespedeza, Japanese clover East Asia Giacalone & Lampel, 1996<br />
Microlophium primulae (Theobald) On Primula Asia CSCF/Lampel, unpublished list<br />
Myzus ascalonicus Doncaster On Allium spp. Near East CSCF/Lampel, unpublished list<br />
Myzus cymbalariae Stroyan<br />
Polyphagous Not clear. In UK, South Meier, 1972<br />
(=cymbalariellus Str.)<br />
Africa, New Zealand and<br />
Australia<br />
Myzus ornatus La<strong>in</strong>g On Prunus cornuta (Wallich ex Royle) (primary<br />
host) and many herbaceous plants (secondary<br />
hosts<br />
Himalaya CSCF/Lampel, unpublished list<br />
Myzus persicae (Sulzer) Polyphagous Cosmopolitan, probably from<br />
Asia<br />
CABI, 2001<br />
Myzus varians Davidson On Clematis East Asia Giacalone & Lampel, 1996<br />
Nearctaphis bakeri (Cowen) Maloideae (primary hosts) and Fabaceae (secondary<br />
hosts)<br />
North America CSCF/Lampel, unpublished list<br />
Pentatrichopus fragaefolii (Cockerell) On strawberry North America CSCF/Lampel, unpublished list<br />
Rhodobium porosum (Sanderson) On Rosa, <strong>in</strong> greenhouses <strong>in</strong> Central Europe Tropics and subtropics Meier, 1972<br />
Rhopalosiphon<strong>in</strong>us latysiphon (Davidson) Polyphagous North America CSCF/Lampel, unpublished list<br />
Rhopalosiphum maidis (Fitch) On Maize, Sorghum, sugar cane and other<br />
Poaceae<br />
Probably Asia Meier, 1975
4 > Insects – Insecta 97<br />
Species<br />
Host plant Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
Toxoptera aurantii (Boyer de Fonscolombe) Polyphagous, <strong>in</strong> Europe ma<strong>in</strong>ly on Citrus Cosmopolitan, tropics and<br />
subtropics<br />
CSCF/Lampel, unpublished list<br />
Uroleucon erigeronense (Thomas)<br />
Callaphididae<br />
On Asteraceae (Erigeron, Coniza, etc.) North America CSCF/Lampel, unpublished list<br />
Myzocallis (=L<strong>in</strong>eomyzocallis) walshii<br />
(Monell)<br />
On Quercus North America Giacalone and Lampel (1996)<br />
Takecallis arundicolens (Clarke) On bamboo South East Asia Lampel and Meier (2003)<br />
Takecallis arund<strong>in</strong>ariae (Essig) On bamboo South East Asia Giacalone and Lampel (1996)<br />
Takecallis taiwanus (Takahashi) On bamboo South East Asia Giacalone and Lampel (1996)<br />
T<strong>in</strong>ocallis nevskyi Remaudière, Quednau &<br />
Heie<br />
Chaitophoridae<br />
On Ulmus Western Asia Giacalone and Lampel (1996)<br />
Periphyllus californiensis (Sh<strong>in</strong>ji)<br />
Pemphigidae<br />
On Acer East Asia Lampel and Meier (2003)<br />
Eriosoma lanigerum (Hausmann)<br />
Lachnidae<br />
On orchard trees North America CPC<br />
C<strong>in</strong>ara cupressi (Buckton)<br />
Phylloxeridae<br />
On Cupressaceae North America? Taxonomy<br />
confus<strong>in</strong>g<br />
Lampel, 1974. Watson et al.,<br />
1999<br />
Viteus vitifoliae (Fitch)<br />
PSYLLINA<br />
Psyllidae<br />
On grapev<strong>in</strong>e North America CPC<br />
Acizzia jamatonica (Kuwayama) On Albizia julibriss<strong>in</strong> Durazz East Asia D. Burckhardt, pers. comm.<br />
Bactericera trigonica Hodk<strong>in</strong>son On Daucus carota L. Mediterranean region Burckhardt and Freuler, 2000<br />
Cacopsylla fulguralis (Kuwayama) On Elaeagnus East Asia D. Burckhardt, pers. comm.<br />
Cacopsylla pulchella (Löw) On Cercis siliquastrum L. Mediterranean region Burckhardt and Freuler, 2000<br />
Camarotoscena speciosa (Flor) On Populus. Maybe natural spread Probably Mediterranean<br />
region, or Asia<br />
Burckhardt and Freuler, 2000<br />
Ctenaryta<strong>in</strong>a eucalypti (Maskell) On Eucalyptus Australia D. Burckhardt, pers. comm.<br />
Homotoma ficus (L.) On Ficus carica L. Mediterranean region and<br />
Middle East<br />
Burckhardt and Freuler, 2000<br />
Livilla spectabilis (Flor) On Spartium junceum L. Mediterranean region Schaefer, 1949<br />
Livilla variegata (Löw) On Laburnum Southern Europe Schaefer, 1949<br />
Phyllopecta trisignata (Löw) On Rubus fruticosus L. Maybe natural spread Southern Europe, Near East Schaefer, 1949<br />
Spanioneura fonscolombii Foerster<br />
Triozidae<br />
On Buxus sempervirens L. Maybe natural spread Mediterranean region Schaefer, 1949<br />
Trioza alacris Flor On Laurus nobilis L. Mediterranean region to<br />
Caucasus<br />
Schaefer, 1949<br />
Trioza centranthi (Vallot)<br />
ALEYRODINA<br />
Aleyrodidae<br />
On Centranthus rubber (L.) Mediterranean region Schaefer, 1949<br />
Bemisia tabaci (Gennadius) Polyphagous, <strong>in</strong> greenhouse Cosmopolitan, probably Asia CABI, 2001<br />
Trialeurodes vaporariorum Westwood<br />
COCCINA<br />
Diaspididae<br />
Polyphagous, <strong>in</strong> greenhouse Central America CABI, 2001<br />
Aonidia lauri (Bouché) Polyphagous, on ornamentals Mediterranean region Kozar et al., 1994
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 98<br />
Species<br />
Host plant Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
Diaspidiotus dist<strong>in</strong>ctus (Leonardi) On Corylus, Matricaria and Quercus Mediterranean region. Maybe<br />
<strong>in</strong>digenous<br />
Kosztarab and Kozar, 1988<br />
Diaspidiotus osborni (Newell & Cockerell) On Platanus, Corylus and Gleditsia North America Kozar et al., 1994<br />
Dynaspidiotus britannicus (Newstead) Polyphagous, <strong>in</strong>doors and outdoors Unclear Kozar et al., 1994<br />
Pseudaulacaspis pentagona (Targioni-<br />
Tozzetti)<br />
Polyphagous, on ornamental and orchard trees East Asia Mani et al., 1997<br />
Quadraspidiotus labiatarum (Marshal) Polyphagous Mediterranean region. Maybe<br />
<strong>in</strong>digenous<br />
Kozar et al., 1994<br />
Quadraspidiotus lenticularis (L<strong>in</strong>d<strong>in</strong>ger) Polyphagous on broadleaved trees Mediterranean region. Maybe<br />
<strong>in</strong>digenous<br />
Kozar et al., 1994<br />
Quadraspidiotus perniciosus (Comstock) Polyphagous, pest <strong>in</strong> orchards East Asia Mani and Baroffio, 1997<br />
Quadraspidiotus pyri (Lichtenste<strong>in</strong>)<br />
Ortheziidae<br />
Polyphagous, pest of fruit trees Mediterranean region. Maybe<br />
<strong>in</strong>digenous<br />
Kozar et al., 1994<br />
Orthezia <strong>in</strong>signis Browne<br />
Pseudococcidae<br />
Polyphagous, <strong>in</strong> greenhouses Neotropics Kozar et al., 1994<br />
Peliococcus multisp<strong>in</strong>us (Siraiwa) On Thymus East Asia, Caucasus? Kozar et. al, 1994<br />
Planococcus citri (Risso) Polyphagous, <strong>in</strong> greehouses and on <strong>in</strong>door plants Tropics and subtropics Kozar et. al, 1994<br />
Trionymus penium (Williams)<br />
Coccidae<br />
On Pseudosasa South-east Asia Kozar et. al, 1994<br />
Chloropulv<strong>in</strong>aria floccifera (Westwood) Polyphagous Mediterranean region or East<br />
Asia?<br />
Kozar et al., 1994<br />
Coccus hesperidum L. Polyphagous, <strong>in</strong>doors and outdoors Cosmopolitan, but prob. not<br />
from Europe<br />
Kozar et al., 1994<br />
Eupulv<strong>in</strong>aria hydrangeae (Ste<strong>in</strong>weden) Polyphagous on broadleaved trees Perhaps East Asia Kozar et al., 1994<br />
Pulv<strong>in</strong>aria regalis Canard<br />
Margarodidae<br />
Polyphagous on broadleaved trees Perhaps East Asia Kozar et al., 1994<br />
Icerya purchasi Maskell<br />
Auchenorrhyncha<br />
Cicadellidae<br />
Polyphagous, <strong>in</strong>doors and outdoors <strong>in</strong> <strong>Switzerland</strong> Australia Kozar et al., 1994<br />
Edwardsiana platanicola (Vidano) On Platanus Unknown. Introduced from<br />
Northen Italy<br />
Günthart, 1987<br />
Eupteryx decemnotata Rey On Salvia Southern Europe Günthart, 1987<br />
Graphocephala fennahi Young On Rhododendron North America Günthart, 1987<br />
Orientus ishidae (Matsumura) On Salix and Betula East Asia Günthart et al., 2004<br />
Scaphoideus titanus Ball<br />
Flatidae<br />
On v<strong>in</strong>e, vector of ‘flavescence dorée’ North America Günthart, 1987<br />
Metcalfa pru<strong>in</strong>osa Say<br />
Membracidae<br />
Polyphagous North America Bonavia and Jerm<strong>in</strong>i, 1998<br />
Stictocephala bisonia Kopp & Yonke<br />
Heteroptera<br />
Lygaeidae<br />
Polyphagous North America Günthart, 1987<br />
Arocatus longiceps Stal On Platanus Mediterranean region,<br />
perhaps expand<strong>in</strong>g naturally<br />
Giacalone et al., 2002<br />
Orsillus depressus Dallas On Cupressaceae Mediterranean region,<br />
perhaps expand<strong>in</strong>g naturally<br />
R. Heckmann, pers. comm.
4 > Insects – Insecta 99<br />
Species<br />
Host plant Orig<strong>in</strong> Reference for <strong>Switzerland</strong><br />
Oxycarenus lavaterae (F.)<br />
Miridae<br />
On Malvaceae and Tiliaceae Mediterranean region,<br />
perhaps expand<strong>in</strong>g naturally<br />
R. Heckmann, pers. comm.<br />
Deraeocoris flavil<strong>in</strong>ea (A. Costa)<br />
T<strong>in</strong>gidae<br />
Predator on aphids on broead-leaved trees Mediterranean Region,<br />
perhaps expand<strong>in</strong>g naturally<br />
Rabitsch, 2002<br />
Corythucha arcuata (Say) On Quercus North America Meier et al., 2004<br />
Corythucha ciliata (Say) On Platanus North America Barbey, 1996<br />
Tab. 4.6 > Established <strong>alien</strong> <strong>in</strong>sects <strong>in</strong> <strong>Switzerland</strong>: Orthoptera, Dictyoptera, Thysanoptera, Psocoptera, Syphonaptera and Anoplura.<br />
Species<br />
Habitat – Life traits Orig<strong>in</strong> References for <strong>Switzerland</strong><br />
ORTHOPTERA<br />
Gryllidae<br />
Acheta domesticus (L.)<br />
Rhaphidophoridae<br />
Omnivorous, synanthropic, also ouside <strong>in</strong> the<br />
Valais<br />
North Africa, Cosmopolitan Thorens and Nadig, 1997<br />
Tachyc<strong>in</strong>es asynamorus Adelung<br />
DICTYOPTERA<br />
Blattellidae<br />
Omnivorous, greenhouses and botanical gardens Cosmopolitan, probably from<br />
East Asia<br />
Thorens and Nadig, 1997<br />
Blattella germanica (L.) Omnivorous, synanthropic Cosmopolitan Landau et al., 1999<br />
Supella longipalpa (F.)<br />
Blattidae<br />
Omnivorous, synanthropic Africa Landau et al., 1999<br />
Blatta orientalis L. Omnivorous, synanthropic Cosmopolitan, possibly<br />
<strong>in</strong>digenous<br />
Landau et al., 1999<br />
Periplaneta americana (L.) Omnivorous, synanthropic Africa, cosmopolitan, Landau et al., 1999<br />
Periplaneta australasiae (F.)<br />
THYSANOPTERA<br />
Thripidae<br />
Omnivorous, synanthropic Cosmopolitan, tropics and<br />
subtropics<br />
Landau et al., 1999<br />
Frankl<strong>in</strong>iella <strong>in</strong>tonsa (Trybom) Polyphagous, ma<strong>in</strong>ly <strong>in</strong> greenhouses East Asia CABI, 2001, L<strong>in</strong>der et al., 1998<br />
Frankl<strong>in</strong>iella occidentalis (Pergande) Polyphagous, ma<strong>in</strong>ly <strong>in</strong> greenhouses North America. Now cosmopolitan<br />
CABI, 2001, Ebener et al., 1989<br />
Heliothrips haemorrhoidalis (Bouché) Polyphagous, <strong>in</strong> greenhouses Probably tropical America.<br />
Now cosmopolitan<br />
CABI, 2001<br />
Thrips simplex (Morison)<br />
PSOCOPTERA<br />
Trogiomorpha<br />
Trogiidae<br />
Develops on Gladiolus, but found also on many<br />
other plants, <strong>in</strong> greenhouses<br />
Probably South Africa CABI, 2001<br />
Cerobasis annulata (Hagen) Domestic Unclear (see text) Lienhard, 1994<br />
Lep<strong>in</strong>otus <strong>in</strong>quil<strong>in</strong>us von Heyden Domestic, rarely outdoors Unclear (see text) Lienhard, 1994
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 100<br />
Species<br />
Habitat – Life traits Orig<strong>in</strong> References for <strong>Switzerland</strong><br />
Lep<strong>in</strong>otus patruelis Pearman Domestic, rarely outdoors Unclear (see text) Lienhard, 1994<br />
Lep<strong>in</strong>otus reticulatus Enderle<strong>in</strong> Domestic, occasionally outdoors Unclear (see text) Lienhard, 1994<br />
Trogium pulsatorium (L.) Domestic Unclear, perhaps Mediterranean<br />
region (see text)<br />
Psyllipsocidae<br />
Dorypteryx domestica (Smithers)<br />
Dorypteryx longipennis Smithers<br />
Dorypteryx pallida Aaron<br />
Psyllipsocus ramburii Sélys-Longchamps<br />
Troctomorpha<br />
Liposcelididae<br />
Domestic<br />
Domestic<br />
Domestic<br />
Domestic and <strong>in</strong> caves<br />
Unclear, perhaps Africa (see<br />
text)<br />
Unclear (see text)<br />
Unclear, perhaps North<br />
America (see text)<br />
Unclear (see text)<br />
Lienhard, 1994<br />
Lienhard, 1994<br />
Lienhard, 1994<br />
Lienhard, 1994<br />
Lienhard, 1994<br />
Liposcelis bostrychophila Badonnel Domestic, occasionally outdoors Unclear (see text) Lienhard, 1994<br />
Liposcelis brunnea Motschulsky Domestic, occasionally outdoors Unclear (see text) Lienhard, 1994<br />
Liposcelis corrodens (Heymons) Domestic, occasionally outdoors Unclear (see text) Lienhard, 1994<br />
Liposcelis decolor (Pearman) Domestic, occasionally outdoors Unclear (see text) Lienhard, 1994<br />
Liposcelis entomophila (Enderle<strong>in</strong>) Domestic Unclear (see text) Lienhard, 1994<br />
Liposcelis mendax Pearman<br />
Liposcelis pearmani Lienhard<br />
Domestic<br />
Domestic, occasionally outdoors<br />
Unclear, perhaps Asia (see<br />
text)<br />
Unclear (see text)<br />
Lienhard, 1994<br />
Lienhard, 1994<br />
Liposcelis pubescens Broadhead Domestic Unclear (see text) Lienhard, 1994<br />
Sphaeropsocidae<br />
Badonnelia titei Pearman Domestic Unclear (see text) Lienhard, 1994<br />
Psocomorpha<br />
Ectopsocidae<br />
Ectopsocus pumilis (Banks)<br />
Ectopsocus richardsi (Pearman)<br />
Ectopsocus vachoni Badonnel<br />
SIPHONAPTERA<br />
Pulicidae<br />
Ctenocephalides felis felis<br />
Spilopsylus cuniculi (Dale)<br />
ANOPLURA<br />
Hoplopleuridae<br />
Domestic<br />
Domestic<br />
Domestic<br />
(Bouché) Ectoparasite on cat and, occasionally, other<br />
carnivores<br />
Ectoparasite on rabbit and, occasionally, other<br />
mammals<br />
Unclear, perhaps Asia (see<br />
text)<br />
Unclear, perhaps Asia (see<br />
text)<br />
Unclear, perhaps Mediterranean<br />
region (see text)<br />
Cosmopolitan, probably not<br />
Europe<br />
Probably Iberian Pen<strong>in</strong>sula<br />
Lienhard, 1994<br />
Lienhard, 1994<br />
Lienhard, 1994<br />
Beaucournu and Launay (1990)<br />
Beaucournu and Launay (1990)<br />
Haemodipsus ventricosus (Denny) Ectoparasite on rabbit Probably Iberian Pen<strong>in</strong>sula Büttiker and Mahnert (1978)
5 > Spiders and Allies – Arachnida 101<br />
5 > Spiders and Allies – Arachnida<br />
Prepared by Theo Blick, Ambros Haenggi and Rüdiger Wittenberg<br />
5.1 Introduction<br />
This chapter summarizes the available <strong>in</strong>formation on Arachnida, except Acari, cover<strong>in</strong>g<br />
<strong>species</strong>’ distribution, biology, and potential harm to the environment and economy.<br />
S<strong>in</strong>ce knowledge of the natural distribution, orig<strong>in</strong>s and movement for the Arachnida is<br />
very limited, it was decided to use specific def<strong>in</strong>itions for the terms described below,<br />
so that coverage of <strong>in</strong>vasive <strong>species</strong> is expanded to <strong>in</strong>clude native <strong>species</strong> which are<br />
spread<strong>in</strong>g, thereby not discrim<strong>in</strong>at<strong>in</strong>g between natural spread and human-mediated<br />
expansion. The follow<strong>in</strong>g def<strong>in</strong>itions characterize the framework for this chapter, and<br />
expla<strong>in</strong> which <strong>species</strong> are covered and which are not.<br />
> Neozoa (Geiter et al., 2002): A neozoan animal has been <strong>in</strong>troduced, by direct or<br />
<strong>in</strong>direct human mediation, to a region to which it is not native and has established a<br />
population there.<br />
> <strong>Invasive</strong> Species (Geiter et al., 2002): This term does not discrim<strong>in</strong>ate between<br />
natural and human-mediated colonization of a new territory and focuses on <strong>species</strong><br />
caus<strong>in</strong>g problems.<br />
The spiders discussed <strong>in</strong> this report are categorized ma<strong>in</strong>ly by habitat and biology, as<br />
follows:<br />
> Species of natural habitats: spiders and their relatives which live <strong>in</strong> natural, nearnatural<br />
or human-<strong>in</strong>fluenced habitats (e.g. crop fields), but not <strong>in</strong> close proximity to<br />
human build<strong>in</strong>gs. The report focuses on <strong>species</strong> which have changed their distribution,<br />
ma<strong>in</strong>ly dur<strong>in</strong>g the last two decades because of a lack of older data. It is based<br />
on pitfall trap results, s<strong>in</strong>ce cont<strong>in</strong>uous and standardized <strong>in</strong>formation on orb web<br />
<strong>species</strong> is not available.<br />
> Species <strong>in</strong>side, and <strong>in</strong> close proximity, to human build<strong>in</strong>gs: spiders and their<br />
relatives which typically <strong>in</strong>habit walls of build<strong>in</strong>gs or live <strong>in</strong> direct contact with humans,<br />
and which have expanded their distribution range <strong>in</strong>to Central Europe dur<strong>in</strong>g<br />
recent decades.<br />
> House-dwell<strong>in</strong>g <strong>species</strong>: spiders and their relatives which exclusively occur <strong>in</strong><br />
build<strong>in</strong>gs, and no populations <strong>in</strong> natural habitats are known.<br />
> Greenhouse-<strong>in</strong>habit<strong>in</strong>g <strong>species</strong>: spiders and their relatives which, <strong>in</strong> Central<br />
Europe, exclusively <strong>in</strong>habit greenhouses and other similarly warm build<strong>in</strong>gs. They<br />
have established populations <strong>in</strong> these warm environments, but cannot survive out-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 102<br />
side due to their climatic requirements. They can also be <strong>in</strong>troduced <strong>in</strong>to houses, e.g.<br />
with flowers (e.g. Eperigone eschatological (Crosby)).<br />
> ‘Banana spiders’: spiders which are <strong>in</strong>troduced with fruit commodities, especially<br />
bananas. They are often rather spectacular <strong>in</strong>dividuals, but they are not able to establish<br />
<strong>in</strong> our climate.<br />
> Terrarium <strong>species</strong>: spiders, mostly tarantulas/theraphosids (so-called ‘bird-eat<strong>in</strong>g<br />
spiders’) from warm regions, which escape captivity, but cannot breed <strong>in</strong> the Central<br />
European climate and at the most survive until the follow<strong>in</strong>g w<strong>in</strong>ter.<br />
The orders Araneae (spiders), Opiliones (harvestmen) and Pseudoscorpiones (false<br />
scorpions) with<strong>in</strong> the class Arachnida are covered <strong>in</strong> this report. Acari (ticks and<br />
mites), although very important for the agricultural and the health sectors, are not<br />
<strong>in</strong>cluded <strong>in</strong> this review, because of the difficulties of prepar<strong>in</strong>g comprehensive lists and<br />
their m<strong>in</strong>or relevance for the environmental sector. Species which were <strong>in</strong>troduced to<br />
<strong>Switzerland</strong> more than a few decades ago are not <strong>in</strong>cluded, because of their largely<br />
unresolved status. Thus, the wasp spider (Argiope bruennichi (Scopoli)) is not <strong>in</strong>cluded<br />
<strong>in</strong> the list. Additionally, <strong>species</strong> which besides <strong>in</strong>habit<strong>in</strong>g houses and basements also<br />
live <strong>in</strong> caves, rock screes, walls <strong>in</strong> v<strong>in</strong>eyards or quarries, as for example the genus<br />
Pholcus or the jump<strong>in</strong>g spider Salticus scenicus (Clerck), are not discussed here.<br />
The ma<strong>in</strong> general literature sources for the report are Thaler and Knoflach (1995),<br />
Geiter et al. (2002) and Komposch (2002).<br />
Knowledge about synanthropic spiders and their relatives <strong>in</strong> <strong>Switzerland</strong> is extremely<br />
limited. Therefore many observations discussed <strong>in</strong> the report are based on knowledge,<br />
although likewise rudimentary, accumulated <strong>in</strong> other Central European countries, and<br />
extrapolated to <strong>Switzerland</strong>. The situation <strong>in</strong> these countries will be similar to that <strong>in</strong><br />
<strong>Switzerland</strong> and <strong>in</strong>clud<strong>in</strong>g the <strong>in</strong>formation gives a more comprehensive picture of the<br />
<strong>alien</strong> spider fauna of Central Europe.<br />
Generally, synanthropic spiders seem to attract less attention than <strong>species</strong> <strong>in</strong>habit<strong>in</strong>g<br />
natural habitats even among arachnologists or, rather, little is published about them.<br />
The spitt<strong>in</strong>g spider Scytodes thoracica (Latr.) is an example of a spider almost exclusively<br />
found <strong>in</strong> houses <strong>in</strong> Central Europe; its distribution is fairly well-known by<br />
arachnologists, but there are few publications about it. The same is true for <strong>species</strong> of<br />
the genus Araneus, which frequently occur <strong>in</strong> houses and gardens.<br />
5.2 List of <strong>species</strong><br />
Table 5.1 <strong>in</strong>troduces the <strong>species</strong> mentioned <strong>in</strong> this report.
5 > Spiders and Allies – Arachnida 103<br />
Tab. 5.1 > Species mentioned <strong>in</strong> this report, named after Platnick (2004).<br />
Species name Author, Year Family Habitat<br />
Achaearanea tabulata * Levi, 1980 Theridiidae In houses<br />
Artema atlanta * Walckenaer, 1837 Pholcidae In houses<br />
Astrobunus laevipes * (Canestr<strong>in</strong>i, 1872) Phalangiidae Natural<br />
Chelifer cancroides (L., 1758) Cheliferidae In houses<br />
Cicur<strong>in</strong>a japonica (Simon, 1886) Dictynidae Natural<br />
Coleosoma floridanum Banks, 1900 Theridiidae Greenhouses<br />
Coll<strong>in</strong>sia <strong>in</strong>errans (O. P.-Cambridge, 1885) L<strong>in</strong>yphiidae Natural<br />
Dasylobus graniferus (Canestr<strong>in</strong>i, 1871) Phalangiidae Natural<br />
Dictyna civica (Lucas, 1850) Dictynidae On build<strong>in</strong>gs<br />
Diplocephalus graecus * (O. P.-Cambridge, 1872) L<strong>in</strong>yphiidae Natural<br />
Eperigone eschatologica (Crosby, 1924) L<strong>in</strong>yphiidae Greenhouses<br />
Eperigone trilobata (Emerton, 1882) L<strong>in</strong>yphiidae Natural<br />
Erigone autumnalis Emerton, 1882 L<strong>in</strong>yphiidae Natural<br />
Harpactea rubicunda (C.L. Koch, 1838) Dysderidae In houses, but also natural<br />
Hasarius adansoni (Audou<strong>in</strong>, 1826) Salticidae Greenhouses<br />
Heteropoda venatoria (L., 1767) Sparassidae Greenhouses<br />
Holocnemus pluchei (Scopoli, 1763) Pholcidae In houses<br />
Micropholcus fauroti * (Simon, 1887) Pholcidae In houses<br />
Nesticus eremita Simon, 1879 Nesticidae Natural<br />
Oecobius maculatus Simon, 1870 Oecobiidae Natural<br />
Opilio canestr<strong>in</strong>ii (Thorell, 1876) Phalangiidae On build<strong>in</strong>gs<br />
Ostearius melanopygius (O. P.-Cambridge, 1879) L<strong>in</strong>yphiidae Natural<br />
Pseudeuophrys lanigera (Simon, 1871) Salticidae On build<strong>in</strong>gs<br />
Psilochorus simoni (Berland, 1911) Pholcidae In houses<br />
Thanatus vulgaris * Simon, 1870 Philodromidae Greenhouses<br />
Uloborus plumipes Lucas, 1846 Uloboridae Greenhouses<br />
Zodarion italicum (Canestr<strong>in</strong>i, 1868) Zodariidae Natural<br />
Zodarion rubidum Simon, 1914 Zodariidae Natural<br />
Zoropsis sp<strong>in</strong>imana (Dufour, 1820) Zoropsidae In houses<br />
*Species not yet recorded from <strong>Switzerland</strong>.<br />
5.3 Species of natural habitats<br />
Eperigone trilobata (Emerton)<br />
This spider has a wide distribution range <strong>in</strong> North America (Millidge, 1987) and was<br />
first recorded from Europe <strong>in</strong> the 1980s at Karlsruhe, Germany. A catalogue of Swiss<br />
spiders (Maurer and Hänggi, 1990) noted several records from the Swiss cantons Jura<br />
and Tic<strong>in</strong>o. This <strong>species</strong> is a common member of the spider fauna <strong>in</strong> all open habitats<br />
(e.g. Blick et al., 2000) and was first recorded <strong>in</strong> the Jura <strong>in</strong> 1999 at about 800 m above<br />
sea level (T. Blick, unpubl.).
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 104<br />
Fig. 5.1 > Records of Eperigone trilobata (left) and Zodarion italicum (right) <strong>in</strong> Germany<br />
and north-west <strong>Switzerland</strong>.<br />
After Staudt (2004).<br />
(green dots: s<strong>in</strong>ce 2000, grey dots: 1990–1999, red dots: 1980–1989)<br />
In Germany, this <strong>species</strong> has spread from Baden-Württemberg, where it was first<br />
found, to Hessen and Rhe<strong>in</strong>land-Pfalz and <strong>in</strong>to north-western Bayern (Staudt, 2004: see<br />
Fig. 5.1). There are also records from outside this area, i.e. <strong>in</strong> the east of Bayern and<br />
the south of Niedersachsen (T. Blick, unpubl.; pers. comm. of various arachnologists –<br />
these data are not <strong>in</strong>cluded <strong>in</strong> Fig. 5.1). The data presented above <strong>in</strong>dicate that this<br />
<strong>species</strong> will colonize Central Europe <strong>in</strong> the foreseeable future. However, it is unclear<br />
what the altitude limit will be or whether its frequency <strong>in</strong> samples will <strong>in</strong>crease. Dur<strong>in</strong>g<br />
past years, there has been no <strong>in</strong>dication that the frequency of the <strong>species</strong> is <strong>in</strong>creas<strong>in</strong>g –<br />
to the contrary, <strong>in</strong>vestigations on the same field site <strong>in</strong> 1994 and 1999 showed a decrease<br />
<strong>in</strong> numbers of the <strong>species</strong> (Baur et al., 1996; Hänggi and Baur, 1998; Blick,<br />
unpubl.). In most samples the frequency of the <strong>species</strong> reached only about 5 % – the<br />
maximum was 30 % at Basel railway station <strong>in</strong> 2002 (Hänggi and Heer, unpubl.).<br />
Displacement of native spider <strong>species</strong> has not been recorded and would be very difficult<br />
to prove. Such studies would <strong>in</strong>volve standardized experiments at the same field<br />
site over decades, with year-round sampl<strong>in</strong>g and also Swiss-wide sites for comparative<br />
work. Fund<strong>in</strong>g for this k<strong>in</strong>d of research seems unlikely, although the samples could be<br />
used for other arthropod research.
5 > Spiders and Allies – Arachnida 105<br />
Zodarion italicum (Canestr<strong>in</strong>i)<br />
The orig<strong>in</strong> of the aptly named Zodarion italicum is the south of Europe. It has expanded<br />
its distribution s<strong>in</strong>ce the publication of the catalogue of Swiss spiders (Maurer<br />
and Hänggi, 1990 – cited as sub Z. gallicum), and has s<strong>in</strong>ce reached the south of <strong>Switzerland</strong>.<br />
It is possible that it is even native to that region (see, e.g., Lessert, 1910). This<br />
spider is a highly specialized predator of ants (Pekar and Kral, 2002) and occurs ma<strong>in</strong>ly<br />
<strong>in</strong> open habitats. The rapid expansion of its range is probably attributable to humanmediated<br />
transport to new areas and to global warm<strong>in</strong>g, which allows <strong>species</strong> of southern<br />
orig<strong>in</strong> to survive north of the Alps. However, this probable relationship would be<br />
difficult to demonstrate. The <strong>species</strong> is also expand<strong>in</strong>g its range <strong>in</strong> Germany, as is its<br />
sister <strong>species</strong> Z. rubidium Simon (see Bosmans, 1997; Staudt, 2004; also see Figure<br />
5.1).<br />
Other <strong>species</strong> found <strong>in</strong> natural habitats<br />
Besides the two <strong>species</strong> mentioned above, a number of other <strong>species</strong> are spread<strong>in</strong>g <strong>in</strong><br />
Central Europe, or their spread is likely with<strong>in</strong> the next few years. A selection of these<br />
<strong>species</strong> is discussed below.<br />
> Coll<strong>in</strong>sia <strong>in</strong>errans (O.P.-Cambridge) (syn. Milleriana <strong>in</strong>errans, C. submissa) has<br />
been found locally <strong>in</strong> <strong>Switzerland</strong> dur<strong>in</strong>g the last 50 years. Currently, the <strong>species</strong> is<br />
expand<strong>in</strong>g its range <strong>in</strong> western Germany (Klapkarek and Riecken, 1995), and has<br />
reached the north-east of Bayern (Blick, 1999). However, it has not yet reached the<br />
abundance of Eperigone trilobata, despite their similarities <strong>in</strong> size and ecological<br />
niche. The future expansion of this <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> should be monitored.<br />
> Ostearius melanopygius (O.P.-Cambridge): Ruzicka (1995) portrays the spread of<br />
this cosmopolitan <strong>species</strong> of unknown orig<strong>in</strong> (cryptogenic) <strong>in</strong> Europe. In <strong>Switzerland</strong>,<br />
a similar pattern as for C. <strong>in</strong>errans has been observed. However, occasionally<br />
the <strong>species</strong> exhibits mass outbreaks, which can be a nuisance to humans although it<br />
does no actual harm (Sacher, 1978); for cases <strong>in</strong> <strong>Switzerland</strong> see, e.g., Benz et al.<br />
(1983). The normal sampl<strong>in</strong>g techniques us<strong>in</strong>g traps at ground level are not effective<br />
under these circumstances and generally catch only s<strong>in</strong>gle <strong>in</strong>dividuals. The reasons<br />
for the mass outbreaks are not yet understood.<br />
> The distribution of Harpactea rubicunda (C.L. Koch) described by Wiehle (1953)<br />
was restricted to the eastern part of Germany at that time. S<strong>in</strong>ce then, the <strong>species</strong> has<br />
expanded its range considerably westwards, most probably by human-mediated<br />
transport. It occurs <strong>in</strong> houses as well as <strong>in</strong> other synanthropic habitats. In <strong>Switzerland</strong>,<br />
the <strong>species</strong> was found near Zurl<strong>in</strong>den (Hänggi, 1988) and <strong>in</strong> a disused railway<br />
area at Basel (Hänggi and Weiss, 2003).<br />
> Erigone autumnalis Emerton, like Eperigone trilobata, orig<strong>in</strong>ates <strong>in</strong> North America.<br />
The <strong>species</strong> has been found at several locations <strong>in</strong> <strong>Switzerland</strong> (Maurer and<br />
Hänggi, 1990; Hänggi, unpubl.). However, it seems to be less abundant and covers a<br />
smaller range than E. trilobata.<br />
> Nesticus eremita Simon can be found outdoors around Basel (Hänggi and Weiss,<br />
2003; Hänggi, unpubl.), whereas further north the <strong>species</strong> is restricted to underground<br />
canals and artificial caves (Jäger, 1995, 1998; Blick, unpubl.). It is very
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 106<br />
likely that this southern European <strong>species</strong> will expand its range further <strong>in</strong> the near<br />
future.<br />
> Cicur<strong>in</strong>a japonica (Simon): this spider of Japanese–Ch<strong>in</strong>ese orig<strong>in</strong> was not accepted<br />
as an established spider <strong>species</strong> <strong>in</strong> Germany by Platen et al. (1995), s<strong>in</strong>ce the <strong>in</strong>troduction<br />
of the <strong>species</strong> near Kehl am Rhe<strong>in</strong> was thought to be temporary. S<strong>in</strong>ce then,<br />
fairly large populations have been found <strong>in</strong> the area around the railway at Basel<br />
(Hänggi and Heer, unpubl.). This <strong>in</strong>dicates that the <strong>species</strong> is able to establish <strong>in</strong><br />
Europe and the further development of its populations should be monitored.<br />
> Diplocephalus graecus (O.P.-Cambridge): Blick et al. (2000) presents a record of<br />
this <strong>species</strong> of Mediterranean orig<strong>in</strong> from agricultural areas near Paris, France. Today<br />
the <strong>species</strong> has reached Belgium (Bonte et al., 2002), so it is very likely that it<br />
will expand its range <strong>in</strong>to most of Central Europe, <strong>in</strong>clud<strong>in</strong>g <strong>Switzerland</strong>, <strong>in</strong> the near<br />
future.<br />
> Dasylobus graniferus (Canestr<strong>in</strong>i) (syn. Eudasylobus nicaeensis): Martens (1978)<br />
mentions records of this harvestman <strong>species</strong> from southern <strong>Switzerland</strong>. However, <strong>in</strong><br />
1997 more than 100 <strong>in</strong>dividuals were collected near Liestal (canton Baselland) (I.<br />
Weiss, unpubl.). Thus, an expansion <strong>in</strong>to Central Europe of this southern European<br />
<strong>species</strong> is expected, but will probably be difficult to document, s<strong>in</strong>ce almost no research<br />
on harvestmen distribution is be<strong>in</strong>g carried out <strong>in</strong> <strong>Switzerland</strong> or <strong>in</strong> adjacent<br />
areas of Germany.<br />
> Another harvestman, Astrobunus laevipes (Canestr<strong>in</strong>i), is already <strong>in</strong> an expansion<br />
phase <strong>in</strong> Central Europe (see Höfer and Spelda, 2001), especially along rivers. It<br />
reached the Netherlands recently (Wijnhoven, 2003). There are no records for <strong>Switzerland</strong><br />
yet. However, its taxonomic differentiation from A. bernard<strong>in</strong>us Simon,<br />
known from the Jura, would merit attention (see Höfer and Spelda, 2001).<br />
Any estimation of potential impacts of the <strong>species</strong> discussed above would only be<br />
guesswork. Economic damage can reasonably be excluded, but it is possible that native<br />
<strong>species</strong> would be replaced or at least populations reduced due to the <strong>in</strong>vaders. However,<br />
test<strong>in</strong>g this hypothesis would necessitate long-term monitor<strong>in</strong>g studies <strong>in</strong> the field<br />
on established plots. These <strong>in</strong>vestigations are not be<strong>in</strong>g carried out to the knowledge of<br />
the authors.<br />
5.4 Species <strong>in</strong>, and <strong>in</strong> close proximity to, human build<strong>in</strong>gs<br />
Only a few members of the Arachnida are restricted to the outside of houses and other<br />
human-made structures, although there are native <strong>species</strong> naturally occurr<strong>in</strong>g on rocks<br />
and tree bark. The <strong>species</strong> which have expanded their range <strong>in</strong> recent decades are<br />
discussed below:<br />
> Dictyna civica (Lucas) <strong>in</strong>habits walls of houses particularly <strong>in</strong> warm climates<br />
(Braun, 1952; Billaudelle, 1957; van Keer and van Keer, 1987). The <strong>species</strong> is considered<br />
to be a nuisance by owners of houses <strong>in</strong> the lowland parts of <strong>Switzerland</strong> as<br />
well as <strong>in</strong> the Rh<strong>in</strong>e valley <strong>in</strong> Baden-Württemberg (Stächele, 2002).
5 > Spiders and Allies – Arachnida 107<br />
> Pseudeuophrys lanigera (Simon) (syn. Euophrys lanigera) is a good example of a<br />
spider which has been cont<strong>in</strong>uously expand<strong>in</strong>g its range <strong>in</strong> Central Europe <strong>in</strong> recent<br />
decades (Braun, 1960; Wijnhoven, 1997; Staudt 2004). Although Maurer and<br />
Hänggi (1990) listed only a few sites for this <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>, it is likely to<br />
expand its range <strong>in</strong> the near future. It is not known whether the <strong>species</strong> is replac<strong>in</strong>g<br />
or <strong>in</strong>fluenc<strong>in</strong>g the population of native jump<strong>in</strong>g spiders with similar ecological<br />
niches on house walls, e.g. Salticus scenicus.<br />
> The harvestman Opilio canestr<strong>in</strong>ii (Thorell) has established populations on walls of<br />
houses <strong>in</strong> Central Europe (Enghoff, 1987; Bliss, 1990; Gruber, 1988; Malten, 1991;<br />
van der Weele, 1993). However, records of natural populations, i.e. on bark of trees,<br />
have been documented s<strong>in</strong>ce then (e.g. Staudt, 2004). As the monitor<strong>in</strong>g of harvestmen<br />
distribution and ecology <strong>in</strong> <strong>Switzerland</strong> is m<strong>in</strong>imal, there are almost no data<br />
available from <strong>Switzerland</strong> on this <strong>species</strong> (Martens, 1978: sub O. ravennae).<br />
House-dwell<strong>in</strong>g <strong>species</strong><br />
Spiders liv<strong>in</strong>g <strong>in</strong> residential houses have either adapted from native natural habitats<br />
such as tree bark, caves and cliffs, or have been <strong>in</strong>troduced from southern Europe and<br />
become established. Sacher (1983) gives an overview of spiders liv<strong>in</strong>g <strong>in</strong> houses. Some<br />
additional house-dwell<strong>in</strong>g <strong>species</strong> are occasionally observed (Hänggi, 2003), e.g.<br />
Achaearanea tabulata Levi <strong>in</strong> Austria and Germany (see Knoflach, 1991; Thaler and<br />
Knoflach, 1995), and therefore this <strong>species</strong> is likely to be found <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> the<br />
future. However, with the exception of Zoropsis sp<strong>in</strong>imana (Dufour), which is discussed<br />
below, no house-dwell<strong>in</strong>g <strong>species</strong> is show<strong>in</strong>g signs of be<strong>in</strong>g <strong>in</strong>vasive.<br />
Zoropsis sp<strong>in</strong>imana (Dufour)<br />
The first record of Z. sp<strong>in</strong>imana was an <strong>in</strong>dividual caught <strong>in</strong> 1994 <strong>in</strong> a residential house<br />
<strong>in</strong> Basel (Hänggi, 2003). S<strong>in</strong>ce then other records have been reported from houses <strong>in</strong><br />
the south of <strong>Switzerland</strong> (Tic<strong>in</strong>o). Observations <strong>in</strong> Austria (Thaler and Knoflach, 1998)<br />
suggest this <strong>species</strong> could cause problems, as it is one of the very few spiders <strong>in</strong> Central<br />
Europe which can penetrate the human sk<strong>in</strong> with its cheliceres and produce a<br />
pa<strong>in</strong>ful bite (Hansen, 1996). This <strong>species</strong> has not (yet) been recorded from Germany<br />
(Blick et al., 2002).<br />
Other <strong>species</strong> of house-dwell<strong>in</strong>g spiders and their relatives<br />
Some other spider <strong>species</strong> liv<strong>in</strong>g <strong>in</strong> houses are currently found more regularly <strong>in</strong> Central<br />
Europe, e.g. Psilochorus simoni (Berland). These <strong>species</strong> do expand their range,<br />
but are found only <strong>in</strong> small numbers or as <strong>in</strong>dividuals. More <strong>in</strong>vaders can be expected,<br />
<strong>in</strong> particular <strong>in</strong> the family Pholcidae (daddy long legs spiders). This is <strong>in</strong>dicated by<br />
observations <strong>in</strong> the harbour at Antwerpen, Belgium (van Keer and van Keer, 2001),<br />
where the <strong>in</strong>troduced Artema atlanta Walckenaer and Micropholcus fauroti (Simon)
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 108<br />
have become established, and from German cities, where stable populations of<br />
Holocnemus pluchei (Scopoli) have been found (Jäger, 1995, 2000).<br />
Interest<strong>in</strong>gly, <strong>in</strong> contrast to the expand<strong>in</strong>g nature of many house-dwell<strong>in</strong>g <strong>species</strong>, the<br />
false scorpion Chelifer cancroides (L.) seems to be decreas<strong>in</strong>g <strong>in</strong> abundance, although<br />
the distribution and population sizes of this group are very poorly <strong>in</strong>vestigated <strong>in</strong><br />
<strong>Switzerland</strong>, especially <strong>in</strong> synanthropic areas. Apparently, the <strong>in</strong>creased hygiene and<br />
the changed climate <strong>in</strong> houses caused by central heat<strong>in</strong>g affects this <strong>species</strong>. The<br />
considerable alteration <strong>in</strong> temperature, and changes <strong>in</strong> daily temperature and humidity,<br />
might favour some <strong>species</strong>, especially <strong>species</strong> adapted to a warm environment, and<br />
negatively affect others.<br />
5.5 Greenhouse-<strong>in</strong>habit<strong>in</strong>g <strong>species</strong><br />
> Hasarius adansoni (Audou<strong>in</strong>) is a cosmopolitan <strong>species</strong>, which is widespread <strong>in</strong><br />
European greenhouses (Simon, 1901; Holzapfel, 1932; König and Pieper, 2003).<br />
Records from <strong>Switzerland</strong> are summarized by Hänggi (2003). Information on synanthropic<br />
spiders <strong>in</strong> general, and this <strong>species</strong> <strong>in</strong> particular, is too limited to come to any<br />
conclusion about the status of <strong>species</strong>, e.g. whether populations are <strong>in</strong>creas<strong>in</strong>g.<br />
> Uloborus plumipes Lucas is another greenhouse <strong>species</strong>, but is much more common<br />
than the <strong>species</strong> above (Jonsson, 1993, 1998; Thaler and Knoflach, 1995). However,<br />
misidentifications <strong>in</strong> some records aris<strong>in</strong>g from confusion with the congeneric<br />
U. glomosus (Walckenaer) cannot be ruled out.<br />
> The first record of Coleosoma floridanum Banks <strong>in</strong> <strong>Switzerland</strong> was reported by<br />
Knoflach (1999) from the tropical greenhouse of the Old Botanical Garden at Basel.<br />
This pantropical <strong>species</strong> is occasionally reported from greenhouses <strong>in</strong> Europe<br />
(Hillyard, 1981; Broen et al., 1998; Knoflach, 1999). However, its status cannot be<br />
evaluated yet.<br />
> Eperigone eschatologica (Crosby) was first recorded <strong>in</strong> Europe from Germany and<br />
Belgium ( see Kle<strong>in</strong> et al., 1995; Bosmans and Vanuytven, 1998) but very recently it<br />
was found <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong> a private flat, most probably carried on a plant from a<br />
German garden centre (Hänggi, unpubl.). It rema<strong>in</strong>s to be seen whether it will become<br />
established <strong>in</strong> greenhouses <strong>in</strong> <strong>Switzerland</strong>.<br />
> Heteropoda venatoria (L.), a member of the family Thomisidae (crab spiders) of<br />
South-east Asian orig<strong>in</strong>, was repeatedly reported from heated build<strong>in</strong>gs, e.g. <strong>in</strong> zoos<br />
(Jäger, 2000). Surveys and research on the spider faunas of heated build<strong>in</strong>gs and<br />
greenhouses would be a useful exercise, s<strong>in</strong>ce this <strong>species</strong> is also able to penetrate<br />
human sk<strong>in</strong> (c.f. Zoropsis sp<strong>in</strong>imana).<br />
> Thanatus vulgaris Simon, a Mediterranean sister <strong>species</strong> of the native T. atratus<br />
Simon, would most probably be found <strong>in</strong> Swiss greenhouses if an <strong>in</strong>tensive survey<br />
were undertaken (see Jones, 1997; Jäger, 2002).
5 > Spiders and Allies – Arachnida 109<br />
5.6 «Banana spiders» and terrarium <strong>species</strong><br />
S<strong>in</strong>ce the <strong>species</strong> <strong>in</strong>troduced by trade <strong>in</strong> bananas and other tropical fruits (see e.g.<br />
Schmidt, 1971) cannot establish populations <strong>in</strong> our climate, they are only of medical<br />
<strong>in</strong>terest, because they <strong>in</strong>clude some dangerous members of the family Ctenidae from<br />
South America. Pesticides used before or dur<strong>in</strong>g transport will often lead to the death<br />
of spiders en route or shortly afterwards (pers. obs. by T. Blick of a ctenid which<br />
arrived <strong>in</strong> Bayreuth, Germany <strong>in</strong> a consignment of bananas from Brazil).<br />
However, some of the many stable populations of spider <strong>species</strong> <strong>in</strong> greenhouses are<br />
probably the result of such <strong>in</strong>troductions with trade. Thus, it is possible that poisonous<br />
<strong>species</strong>, especially small ones, <strong>in</strong>clud<strong>in</strong>g some dangerous for humans could be accidentally<br />
<strong>in</strong>troduced and established <strong>in</strong> greenhouses (see, e.g., Huhta, 1972).<br />
Spiders appropriately kept <strong>in</strong> terrariums are of no concern, but escaped specimens<br />
should be handled with care, s<strong>in</strong>ce the bite of some <strong>species</strong> (although not many) can be<br />
dangerous for humans. If they escape <strong>in</strong>to the wild they will die when temperatures<br />
drop, s<strong>in</strong>ce all <strong>species</strong> are of tropical or subtropical orig<strong>in</strong>.<br />
5.7 Discussion and recommendations<br />
Monitor<strong>in</strong>g spider <strong>species</strong> currently expand<strong>in</strong>g their range after <strong>in</strong>troduction, and<br />
surveys at places prone to <strong>in</strong>troductions, are recommended as elements of an early<br />
warn<strong>in</strong>g system. This would allow the spread of the <strong>alien</strong> <strong>species</strong> to be followed and<br />
document any displacement of native <strong>species</strong>. Some <strong>species</strong> are <strong>in</strong>vad<strong>in</strong>g, but there is<br />
no way of evaluat<strong>in</strong>g their potential threat to native biodiversity. The data collected on<br />
synanthropic spiders <strong>in</strong> <strong>Switzerland</strong>, as well as other Central European countries, are<br />
too limited to allow any conclusions to be drawn. Some cases of established tropical<br />
<strong>species</strong> and <strong>in</strong>creas<strong>in</strong>g trade <strong>in</strong>dicate the possibility of venomous spider <strong>species</strong> arriv<strong>in</strong>g<br />
<strong>in</strong> <strong>Switzerland</strong>. If a venomous spider becomes established, the public would need to<br />
be well-<strong>in</strong>formed about how to handle the situation and anti-venom kits should be<br />
made available.<br />
Furthermore, to document threats to native <strong>species</strong> <strong>in</strong> natural habitats, the establishment<br />
of long-term surveys <strong>in</strong> specific habitats is recommended. Without further studies<br />
on <strong>alien</strong> spider <strong>species</strong>, potential impacts are only guesswork.<br />
Surveys for synanthropic <strong>alien</strong> spiders and their relatives are of lesser importance, as<br />
long as no dangerous spiders are <strong>in</strong>troduced and the <strong>species</strong> that do arrive cannot<br />
become established outside. Thus, the costs of conduct<strong>in</strong>g surveys for these would<br />
probably not be justified. However, monitor<strong>in</strong>g of some selected <strong>species</strong> (e.g. Oecobius<br />
maculatus Simon and Zoropsis sp<strong>in</strong>imana) to document their spread would be both<br />
worthwhile and manageable.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 110<br />
In particular, we recommend an assessment of <strong>alien</strong> spiders <strong>in</strong> greenhouses and other<br />
heated build<strong>in</strong>gs. Greenhouses <strong>in</strong> nurseries and garden centres are the most likely to be<br />
colonized by <strong>alien</strong> spider <strong>species</strong>, and from there these can be spread to households on<br />
plant material (e.g. Eperigone eschatologica). Research and monitor<strong>in</strong>g are needed,<br />
because of the potential economic impact of some <strong>species</strong>. The first encounter with<br />
Uloborus plumipes is an example: a nursery started an <strong>in</strong>quiry to f<strong>in</strong>d out what <strong>species</strong><br />
had <strong>in</strong>fested their property after this spider had covered all plants with webs, so that<br />
plants became difficult to sell. Moreover, the possibility of accidental <strong>in</strong>troductions of<br />
poisonous spiders should not be underrated. Nurseries would be the ma<strong>in</strong> targets for<br />
these monitor<strong>in</strong>g programmes, but other heated build<strong>in</strong>gs <strong>in</strong> botanical gardens and zoos<br />
should be <strong>in</strong>cluded.<br />
The follow<strong>in</strong>g are considered to have the greatest potential for economic impacts:<br />
> Mass outbreaks of nuisance <strong>species</strong> (see Ostearius melanopygius), but without<br />
caus<strong>in</strong>g real damage.<br />
> A dramatic <strong>in</strong>crease <strong>in</strong> the population of the wall-<strong>in</strong>habit<strong>in</strong>g spider Dictyna civica.<br />
> The potential medical costs for treatments of bites of <strong>in</strong>troduced poisonous spiders,<br />
see ‘banana spiders’, and escaped terrarium <strong>species</strong>.<br />
Hav<strong>in</strong>g noted the potential threats to humans, it has to be stressed that broad use of<br />
pesticides aga<strong>in</strong>st spiders is not a reasonable reaction, because of the non-target effects<br />
of these chemicals and the fact that publicity of such measures will exaggerate arachnophobia,<br />
already well-grounded <strong>in</strong> the human population.<br />
In conclusion, only a very small number of spiders and their relatives are considered as<br />
problem <strong>species</strong> on a global scale, <strong>in</strong>clud<strong>in</strong>g Central Europe (e.g. Welch et al., 2001 do<br />
not list any <strong>alien</strong> spiders for Scotland). A reasonable explanation could be that many<br />
phytophagous <strong>in</strong>sects live <strong>in</strong> closer association with their host plants, e.g. eggs, larvae<br />
and pupae are firmly attached to host plants or <strong>in</strong>side them. This would facilitate their<br />
transport with plant material. Another factor is that spiders with their predatory behaviour<br />
are less obvious than phytophagous <strong>in</strong>sects that damage their host plants. Moreover,<br />
as po<strong>in</strong>ted out earlier, the group is rather neglected and many <strong>in</strong>formation gaps<br />
still exist. However, after successful <strong>in</strong>troduction <strong>in</strong>to a new area, many spiders are<br />
highly capable of a rapid expansion <strong>in</strong> range either by natural means such as balloon<strong>in</strong>g<br />
or by hitch<strong>in</strong>g a ride on vehicles.
5 > Spiders and Allies – Arachnida 111<br />
References<br />
Baur, B., Joshi, J., Schmid, B., Hänggi, A., Borcard, D., Stary, J., Pedroli-<br />
Christen, A., Thommen, G.H., Luka, H., Rusterholz, H.-P., Oggier, P.,<br />
Ledergerber S. and A. Erhardt (1996) Variation <strong>in</strong> <strong>species</strong> richness of<br />
plants and diverse groups of <strong>in</strong>vertebrates <strong>in</strong> three calcareous grasslands<br />
of the Swiss Jura mounta<strong>in</strong>s. Rev. Suisse Zool. 103 (4), 801–833.<br />
Benz, G., Nyffeler M. and R. Hug (1983) Ostearius melanopygius (O.P.-<br />
Cambridge) (Aran., Micryphantidae) neu für die Schweiz. Über e<strong>in</strong><br />
Massenauftreten der Sp<strong>in</strong>ne <strong>in</strong> Zürich und die Zerstörung der Population<br />
durch Schneefall. Mitt. schweiz. Ent. Ges. 56, 201–204.<br />
Billaudelle, H. (1957) Zur Biologie der Mauersp<strong>in</strong>ne Dictyna civica (H.<br />
Luc.) (Dictynidae, Araneida). Zeitschr. Angew. Entomol. 41, 475–512.<br />
Blick, T. (1999) Sp<strong>in</strong>nen auf Kopfsalatfeldern bei Kitz<strong>in</strong>gen (Unterfranken,<br />
Bayern). Arachnol. Mitt. 17, 45–50.<br />
Blick, T., Pfiffner L. and H. Luka (2000) Epigäische Sp<strong>in</strong>nen auf Äckern<br />
der Nordwest-Schweiz im mitteleuropäischen Vergleich (Arachnida:<br />
Araneae). Mitt. Dt. Ges. allg. angew. Entomol. 12, 267–276<br />
Blick, T., Hänggi A. and K. Thaler (2002) Checkliste der Sp<strong>in</strong>nentiere<br />
Deutschlands, der Schweiz, Österreichs Belgiens und der Niederlande<br />
(Arachnida: Araneae, Opiliones, Pseudoscorpiones, Scorpiones,<br />
Palpigradi). Version 1. June 2002.<br />
Internet: http://www.AraGes.de/checklisten.html<br />
Bliss, P. (1990) Zur Verbreitung von Opilion canestr<strong>in</strong>ii (Thorell) <strong>in</strong> der<br />
Deutschen Demokratischen Republik (Arachnida: Opiliones,<br />
Phalangiidae). Acta Zool. Fennica 190, 41–44.<br />
Bonte, D., Criel, P., Baert L. and D. De Bakker (2002) The <strong>in</strong>vasive<br />
occurrence of the Mediterranean dwarfspider Diplocephalus graecus (O.<br />
P.-Cambridge, 1872) <strong>in</strong> Belgium (Araneae: L<strong>in</strong>yphiidae). Belgian Journal<br />
of Zoology 132, 171–173.<br />
Bosmans, R. and H. Vanuytven (1998) Eperigone eschatologica een<br />
Amerikaanse immigrant <strong>in</strong> West Europa (Araneae, Erigonidae). Nwsbr.<br />
Belg. Arachnol. Ver. 13 (1), 15–16.<br />
Bosmans, R. (1997) Revision of the genus Zodarion Walckenaer, 1833,<br />
part II. Western and Central Europe, <strong>in</strong>clud<strong>in</strong>g Italy (Araneae: Zodariidae).<br />
Bullet<strong>in</strong> of the British Arachnological Society 10, 265–294.<br />
Braun, R. (1952) “Maserung“ von Wänden durch Sp<strong>in</strong>nen. Natur u. Volk<br />
82, 230–233.<br />
Braun, R (1960) E<strong>in</strong>e für Deutschland neue Spr<strong>in</strong>gsp<strong>in</strong>ne, Euophrys<br />
lanigera (E. Simon 1871) (Araneae; Salticidae, Heliophan<strong>in</strong>ae,<br />
Euophryeae) mit Bemerkungen zu ihren Männchenvarianten. Nachr.<br />
Naturwiss. Mus. Aschaffenburg 64, 77–85, Taf. I-II.<br />
Broen, B. von, Thaler-Knoflach B. and K. Thaler (1998) Nachweis von<br />
Coleosoma floridanum <strong>in</strong> Deutschland (Araneae: Theridiidae). Arachnol.<br />
Mitt. 16, 31–32.<br />
Enghoff, H. (1987) Opilio canestr<strong>in</strong>ii (Thorell, 1876) – en ny<strong>in</strong>vandret<br />
mejer i Danmark (Opiliones). Ent. Meddr. 55, 39–42.<br />
Geiter, O., Homma S. and R. K<strong>in</strong>zelbach (2002) Bestandsaufnahme und<br />
Bewertung von Neozoen <strong>in</strong> Deutschland. Untersuchung der Wirkung von<br />
Biologie und Genetik ausgewählter Neozoen auf Ökosysteme und<br />
Vergleich mit den potenziellen Effekten gentechnisch veränderter<br />
Organismen. Texte Umweltbundesamt 25/2002, Berl<strong>in</strong>, 290 pp.<br />
Gruber, J. (1988) Neunachweise und Ergänzungen zur Verbreitung von<br />
Opilio canestr<strong>in</strong>ii (Thorell) und Opilio transversalis Roewer. Ann. Naturhist.<br />
Mus. Wien 90B, 361–365.<br />
Hänggi, A. (2003) Nachträge zum ”Katalog der schweizerischen<br />
Sp<strong>in</strong>nen“. 3. Neunachweise von 1999 bis 2002 und Nachweise<br />
synanthroper Sp<strong>in</strong>nen. Arachnol. Mitt. 26.<br />
Hänggi, A. and B. Baur (1998) The effect of forest edge on ground-liv<strong>in</strong>g<br />
arthropods <strong>in</strong> a remnant of unfertilized calcareous grassland <strong>in</strong> the Swiss<br />
Jura mounta<strong>in</strong>s. Mitt. schweiz. entomol. Ges. 71, 343–354.<br />
Hänggi, A. and I. Weiss (2003) Sp<strong>in</strong>nen (Araneae) und Weberknechte<br />
(Opiliones). In: Fauna und Flora auf dem Eisenbahngelände im Norden<br />
Basels. Monogr. Entomol. Ges. Basel 1, 74–79 and 204–205 (Anhang 7).<br />
Hansen, H. (1996) L‘importenza medica di alcuni ragni viventi negli<br />
ambienti urbani di Venezia. Boll. Mus. civ. St. nat. Venezia 45 (1994), 21–<br />
32.<br />
Hillyard, P.D. (1981) Coleosoma floridanum Banks (Araneae: Theridiidae)<br />
and Boeorix manducus Thorell (Opiliones: Assamiidae): two tropical<br />
arachnids <strong>in</strong> botanical gardens. Newsletter of the British Arachnological.<br />
Society 31, 3–4.<br />
Höfer, A.M. and J. Spelda (2001) On the distribution of Astrobunus<br />
laevipes Canestr<strong>in</strong>i, 1872 (Arachnida: Opiliones) <strong>in</strong> Central Europe.<br />
Arachnol. Mitt. 22, 42–49.<br />
Holzapfel, M. (1932) Die Gewächshausfauna des Berner Botanischen<br />
Gartens. Rev. suisse Zool. 39, 325–371.<br />
Huhta, V. (1972) Loxosceles laeta (Nicolet) (Araneae, Loxoscel<strong>in</strong>ae), a<br />
venomous spider established <strong>in</strong> a build<strong>in</strong>g <strong>in</strong> Hels<strong>in</strong>ki, F<strong>in</strong>land, and notes<br />
on some other synanthropic spiders. Ann. Ent. Fennici 38, 152–156.<br />
Jäger, P. (1995) Erstnachweis von Holocnemus pluchei und zweiter<br />
Nachweis von Nesticus eremita für Deutschland <strong>in</strong> Köln (Araneae:<br />
Pholcidae, Nesticidae). Arachnol. Mitt. 10, 23–24.<br />
Jäger, P. (1998) Weitere Funde von Nesticus eremita (Araneae:<br />
Nesticidae) <strong>in</strong> Süddeutschland mit Angaben zur Taxonomie im Vergleich<br />
zu N. cellulanus. Arachnol. Mitt. 15, 13–20.<br />
Jäger, P. (2000) Selten nachgewiesene Sp<strong>in</strong>nenarten aus Deutschland<br />
(Arachnida: Araneae). Arachnol. Mitt. 19, 49–57.<br />
Jäger, P. (2002) Thanatus vulgaris Simon, 1870 – e<strong>in</strong> Weltenbummler<br />
(Araneae: Philodromidae). Mit Anmerkungen zur Term<strong>in</strong>ologie der<br />
weiblichen Genit<strong>alien</strong>. Arachnol. Mitt. 23, 49–57.<br />
Jones, D. (1997) Thanatus vulgaris Simon, 1870 a further British record.<br />
Newsletter of the British Arachnological Society 80, 6–7.<br />
Jonsson, L.J. (1993) Nachweis von Uloborus plumipes <strong>in</strong> e<strong>in</strong>em<br />
Gewächshaus <strong>in</strong> Niedersachsen. Arachnol. Mitt. 6, 42–43.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 112<br />
Jonsson. L.J. (1998) Toftspideln – en sp<strong>in</strong>delart som <strong>in</strong>vanderat Europas<br />
växthus. Fauna och Flora 93, 119–124.<br />
Klapkarek, N. and U. Riecken (1995) Zur Verbreitung und Autökologie von<br />
Coll<strong>in</strong>sia submissa (Araneae: L<strong>in</strong>yphiidae). Arachnol. Mitt. 9, 49–56.<br />
Kle<strong>in</strong>, W., Stock M. and J. Wunderlich (1995) Zwei nach Deutschland<br />
e<strong>in</strong>geschleppte Sp<strong>in</strong>nenarten (Araneae) – Uloboris plumipes Lucas und<br />
Eperigone eschatologica (Bishop) – als Gegenspieler der Weissen Fliege<br />
im geschützen Zierpflanzenbau? Beitr. Araneol. 4, 301–306.<br />
Knoflach, B. (1991) Achaearanea tabulata Levi, e<strong>in</strong>e für Österreich neue<br />
Kugelsp<strong>in</strong>ne (Arachnida, Aranei: Theridiidae). Ber. nat.-med. Vere<strong>in</strong><br />
Innsbruck 78, 59–64.<br />
Knoflach, B. (1999) The comb-footed spider genera Neottiura and<br />
Coleosoma <strong>in</strong> Europe (Araneae, Theridiidae). Mitt. schweiz. entomol. Ges.<br />
72, 341–371.<br />
Komposch, C. (2002) Sp<strong>in</strong>nentiere: Sp<strong>in</strong>nen, Weberknechte,<br />
Pseudoskorpione, Skorpione (Arachnida: Araneae, Opiliones,<br />
Pseudoscorpiones, Scorpiones, Solifugae). In: Essl, F. and W. Rabitsch<br />
(eds) Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, pp. 250–252.<br />
König, R. and H. Pieper (2003) Notizen zur Taxonomie und<br />
geographischen Verbreitung von Hasarius adansoni (Audou<strong>in</strong>, 1826).<br />
Faun.-Ökol. Mitt. 8, 197–200.<br />
Lessert, R. de (1910) Araignées. Catalogue Invertébrés Suisse 3, 1–639.<br />
Malten, A. (1991) Über Opilio canestr<strong>in</strong>ii. Arachnol. Mitt. 1, 81–83.<br />
Martens, J. (1978) Weberknechte, Opiliones – Sp<strong>in</strong>nentiere, Arachnida.<br />
Tierwelt Deutschlands 64, 464 pp.<br />
Maurer, R. and A. Hänggi (1990) Katalog der schweizerischen Sp<strong>in</strong>nen.<br />
Doc. Faun. Helv. 12: o. Pag.<br />
Millidge, A.F. (1987) The Erigon<strong>in</strong>e spiders of North America. Part 8. The<br />
genus Eperigone Crosby and Bishop (Araneae, L<strong>in</strong>yphiidae). Amer. Mus.<br />
Novit. 2885, 1–75.<br />
Pekár, S. and J. Král (2002) Mimicry complex <strong>in</strong> two central European<br />
zodariid spiders (Araneae, Zodariidae): how Zodarion deceives ants.<br />
Biological Journal of the ?L<strong>in</strong>nean Society 75, 517–532.<br />
Platen, R., Blick, T., Bliss, P., Drogla, R., Malten, A., Martens, J., Sacher<br />
P. and J. Wunderlich (1995) Verzeichnis der Sp<strong>in</strong>nentiere (excl. Acarida)<br />
Deutschlands (Arachnida: Araneida, Opilionida, Pseudoscorpionida).<br />
Arachnol. Mitt. Sonderband 1, 1–55.<br />
Platnick, N.I. (2004) The world spider catalog. Version 4.0.<br />
Internet: http://research.amnh.org/entomology/spiders/catalog/<strong>in</strong>dex.html<br />
Ruzicka, V. (1995) The spread<strong>in</strong>g of Ostearius melanopygius (Araneae:<br />
L<strong>in</strong>yphiidae) through Central Europe. European Journal of Entomology 92<br />
(4), 723–726.<br />
Sacher, P. (1978) E<strong>in</strong> Massenvorkommen der Baldach<strong>in</strong>netzsp<strong>in</strong>ne<br />
Ostearius melanopygius (O.P.-Cambridge) <strong>in</strong> Ostthür<strong>in</strong>gen (Araneae:<br />
L<strong>in</strong>yphiidae, Donacochareae). Veröff. Mus. Gera Naturw. R. 6, 53–63.<br />
Sacher, P. (1983) Sp<strong>in</strong>nen (Araneae) an und <strong>in</strong> Gebäuden – Versuch e<strong>in</strong>er<br />
Analyse der synanthropen Sp<strong>in</strong>nenfauna <strong>in</strong> der DDR. Entomol.<br />
Nachrichten Berichte 27, 97–104, 141–152, 197–204, 224.<br />
Schmidt, G.E.W. (1971) Mit Bananen e<strong>in</strong>geschleppte Sp<strong>in</strong>nen. Zool. Beitr.<br />
17 (3), 387–433<br />
Simon, E. (1901) Note sur une Araignée exotique (Hasarius adansoni<br />
Aud.) acclimatée dans les serres chaudes, aux environs de Paris. Bullet<strong>in</strong><br />
de la Société Entomologique de France 1901 (7), 154–155.<br />
Stächele (verfasst von B. Krauss) (2002) Bekämpfung der Mauersp<strong>in</strong>nen<br />
an Hausfassaden. Kle<strong>in</strong>e Anfrage der Abg. Dr. Carm<strong>in</strong>a Brenner CDU und<br />
Antwort des M<strong>in</strong>isteriums für Ernährung und Ländlichen Raum. Landtag<br />
Baden-Württemberg, Drucksache 13/1276, 1–4.<br />
Internet: http://www3.landtagbw.de/WP13/Drucksachen/1000/13_1276_D.PDF<br />
Staudt, A. (2004) Nachweiskarten der Sp<strong>in</strong>nentiere Deutschlands.<br />
Internet: http://www.spiderl<strong>in</strong>g.de.vu<br />
Thaler, K. and B. Knoflach (1995) Adventive Sp<strong>in</strong>nentiere <strong>in</strong> Österreich –<br />
mit Ausblicken auf die Nachbarländer. Stapfia 37, 55–76.<br />
Thaler, K. and B. Knoflach (1998) Zoropsis sp<strong>in</strong>imana (Dufour), e<strong>in</strong>e für<br />
Österreich neue Adventivart (Araneae, Zoropsidae). Ber. nat.-med. Vere<strong>in</strong><br />
Innsbruck 85, 173–185.<br />
Van der Weele, R. (1993) Opilio canestr<strong>in</strong>ii nieuw voor de nederlandse<br />
fauna (Opilionida: Phalangiidae). Ent. Ber. Amsterdam 53, 91.<br />
Van Keer, J. and K. Van Keer (1987) Bevestig<strong>in</strong>g van het voorkommen van<br />
Dictyna civica (Lucas) <strong>in</strong> Belgie en de verspreid<strong>in</strong>g van de soort. Nwsbr.<br />
Belg. Arachnol. Ver. 6, 7–8<br />
Van Keer, K. and J. Van Keer (2001) Ingeburgerde exotische trilsp<strong>in</strong>nen<br />
(Araneae: Pholcidae) <strong>in</strong> Antwerpse haven en enkele algemen<br />
bedenk<strong>in</strong>kgen bij sp<strong>in</strong>nenmigratie. Nwsbr. Belg. Arachnol. Ver. 16 (3),<br />
81–86.<br />
Welch, D., Carss, D.N., Gornall, J., Manchester, S.J., Marquiss, M.,<br />
Preston, C.D., Telfer, M.G., Arnold, H. and J. Holbrook (2001) An audit of<br />
<strong>alien</strong> <strong>species</strong> <strong>in</strong> Scotland. Scottish Natural Heritage Review 139, 1–225.<br />
Wijnhoven, H. (1997) Euophrys lanigera (Simon) met recht op de<br />
nederlandse soortenlist. Nieuwsbrief Sp<strong>in</strong>ed 12, 1–3.<br />
Wijnhoven, H. (2003) De hooiwagen Astrobunus laevipes nieuw voor<br />
Nederland (Opiliones: Phalangiidae). Nederl. Faun. Meded. 19, 73–78.
6 > Molluscs – Mollusca 113<br />
6 > Molluscs – Mollusca<br />
Prepared by Rüdiger Wittenberg<br />
The molluscs are a very large successful group of about 130,000 described <strong>species</strong>,<br />
mak<strong>in</strong>g them second <strong>in</strong> number of <strong>species</strong> only to the arthropods (Remane et al., 1981).<br />
However, of the molluscs’ seven classes only two occur <strong>in</strong> <strong>Switzerland</strong>, s<strong>in</strong>ce the<br />
others are strictly mar<strong>in</strong>e <strong>species</strong>.<br />
In the text below, gastropod and bivalve <strong>species</strong> are discussed separately. For ease of<br />
reference, <strong>in</strong> Table 6.2 the <strong>species</strong> are divided <strong>in</strong>to terrestrial snails (<strong>in</strong>clud<strong>in</strong>g slugs),<br />
freshwater snails and bivalves, while the Fact Sheets are <strong>in</strong> alphabetical order of <strong>species</strong>.<br />
A total of 16 gastropods and three bivalves are regarded as established <strong>alien</strong> <strong>species</strong>.<br />
6.1 Snails and slugs (Gastropoda)<br />
Gastropods make up 70 % of all molluscs and are well represented <strong>in</strong> mar<strong>in</strong>e, freshwater<br />
and terrestrial habitats. About 196 terrestrial and 50 aquatic <strong>species</strong> occur <strong>in</strong> <strong>Switzerland</strong><br />
(Turner et al., 1998).<br />
The exact natural distribution of the gastropod <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> is not well<br />
known, and many current populations may be early relocations (<strong>in</strong>troductions with<strong>in</strong><br />
<strong>Switzerland</strong>). Snails are very successful at hitch<strong>in</strong>g a ride, so that many former barriers<br />
have been easily bridged through human-mediated transport. Snails arrived on plant<br />
material as early as dur<strong>in</strong>g Roman times. The aquatic snail Viviparus ater (Mol<strong>in</strong>a) is<br />
an example of relocation <strong>in</strong> <strong>Switzerland</strong>. Its native range is believed to be south of the<br />
Alps, <strong>in</strong>clud<strong>in</strong>g the Tic<strong>in</strong>o, but today its range north of the Alps encompasses the lower<br />
regions between Lac de Genève and Bodensee. It was released <strong>in</strong>to Lac de Genève<br />
before 1900.<br />
Snails found <strong>in</strong> greenhouses are not covered by this report if they do not occur outside<br />
as well. Turner et al. (1998) provide a discussion of greenhouse snails and <strong>species</strong> that<br />
have been recorded only temporarily, and which are therefore not considered to be<br />
established <strong>in</strong> the wild.<br />
The snail Solatopupa similis (Bruguière) (Chondr<strong>in</strong>idae) was <strong>in</strong>troduced to a location<br />
close to Locarno <strong>in</strong> the 19 th century for unknown reasons. This population thrives, but<br />
the <strong>species</strong>’ impact is considered m<strong>in</strong>imal because of its very localized distribution.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 114<br />
The family Milacidae with two native <strong>species</strong>, has been supplemented <strong>in</strong> <strong>Switzerland</strong><br />
with two more, i.e. Milax gagates (Draparnaud) and Tandonia budapestensis (Hazay),<br />
both from other parts of Europe. M. gagates has been found only <strong>in</strong> gardens, where it<br />
probably arrived with plant material, so it is not known whether it is established <strong>in</strong><br />
<strong>Switzerland</strong>. T. budapestensis is an anthropochorous (disperses as a result of human<br />
activity) <strong>species</strong> that is widely distributed with<strong>in</strong> Europe. It is regarded as a pest,<br />
especially <strong>in</strong> w<strong>in</strong>ter crops (Fischer and Reischütz, 1998), but is rather difficult to<br />
observe because of its cryptic, nocturnal habits.<br />
Another synanthropic <strong>species</strong> (<strong>in</strong> addition to members of the Milacidae) is Limacus<br />
flavus (L.) (Limacidae), which is only rarely encountered <strong>in</strong> <strong>Switzerland</strong>. It is likely<br />
that this <strong>species</strong> will be <strong>in</strong>troduced with plant material <strong>in</strong> the future, but it is probably<br />
not adapted to the Swiss environment, so that it is less likely to become established.<br />
The slugs Deroceras sturanyi (Simroth) and D. panormitanum (Lessona & Pollonera)<br />
belong to the Agriolimacidae. This family is taxonomically challeng<strong>in</strong>g and new<br />
<strong>species</strong> have recently been described (Kerney et al., 1983). These slugs prefer fresh<br />
green plant material and therefore some are recognized as pests. The former <strong>species</strong> has<br />
been considered a pest <strong>in</strong> gardens, while the latter <strong>species</strong> is only rarely found <strong>in</strong><br />
<strong>Switzerland</strong>. However, D. panormitanum is expand<strong>in</strong>g its range <strong>in</strong> Europe and is<br />
<strong>in</strong>creas<strong>in</strong>gly reported as damag<strong>in</strong>g plants of economic importance. This is a <strong>species</strong><br />
that should be monitored, s<strong>in</strong>ce it is likely to establish populations <strong>in</strong> agricultural areas,<br />
with the potential to cause economic damage <strong>in</strong> the near future.<br />
The orig<strong>in</strong> of the slug Boettgerilla pallens Simroth (Boettgerillidae) is assumed to be<br />
the Caucasus, although Jungbluth (1996) argued that it might be a native <strong>species</strong> <strong>in</strong><br />
Central Europe which had not been reported earlier. Most collections of slugs are fairly<br />
recent and the lack of a shell makes it difficult to f<strong>in</strong>d <strong>species</strong> as subfossils. This <strong>species</strong><br />
is a predator of slug eggs and young slugs (Reischütz, 2002), so it is a beneficial<br />
organism <strong>in</strong> agricultural sett<strong>in</strong>gs rather than a pest, as sometimes stated. However, <strong>in</strong><br />
natural localities, it may have a negative impact on native slugs through predation.<br />
Arion lusitanicus Mabille (Arionidae) (see Fact Sheet) is rapidly expand<strong>in</strong>g its range,<br />
and is the most serious <strong>in</strong>vader amongst the snails and slugs. It is not only a pest <strong>in</strong><br />
agriculture and gardens, but also displaces a native congeneric <strong>species</strong> <strong>in</strong> the lowland<br />
parts of <strong>Switzerland</strong> (Turner et al., 1998).<br />
The snail Hygromia c<strong>in</strong>ctella (Draparnaud) (Hygromiidae) has been accidentally<br />
<strong>in</strong>troduced <strong>in</strong>to the northern parts of <strong>Switzerland</strong>, but could be native around Genève or<br />
<strong>in</strong> the Tic<strong>in</strong>o. It was <strong>in</strong>troduced prior to the start of malacological record<strong>in</strong>g and is<br />
mostly found <strong>in</strong> gardens and other anthropogenic sett<strong>in</strong>gs (Kerney et al., 1983).<br />
The only <strong>in</strong>troduced <strong>species</strong> of Helicidae is Cryptomphalus aspersus (O.F. Müller),<br />
which was <strong>in</strong>troduced prior to the start of malacological record<strong>in</strong>g. It is mostly found <strong>in</strong><br />
gardens and other anthropogenic habitats, although it is recorded from natural habitats,<br />
e.g. <strong>in</strong> the Valais. Its environmental impact is not known, but it is regarded as a pest <strong>in</strong><br />
gardens.
6 > Molluscs – Mollusca 115<br />
The freshwater snail Potamopyrgus antipodarum (Gray) (Hydrobiidae) (see Fact<br />
Sheet) is one of the few long-distance <strong>in</strong>vaders <strong>in</strong> this group. It is native to New Zealand<br />
and has been spread around the world probably with ballast water between freshwater<br />
systems and on ornamental aquatic plants. Haynes et al. (1985) have suggested<br />
another pathway as they have shown that P. antipodarum can survive a six-hour passage<br />
through the gut of a trout and produce live young shortly afterwards. It is very<br />
likely to <strong>in</strong>duce ecosystem changes because of its enormous numbers <strong>in</strong> some places.<br />
Two Physella <strong>species</strong>, Physella acuta (Draparnaud) and P. heterostropha (Say) (Physidae),<br />
have been <strong>in</strong>troduced <strong>in</strong>to <strong>Switzerland</strong>. The latter <strong>species</strong> is of North American<br />
orig<strong>in</strong>, but the orig<strong>in</strong> of the former <strong>species</strong> is disputed, although it probably orig<strong>in</strong>ates<br />
<strong>in</strong> south-western Europe. S<strong>in</strong>ce P. heterostropha is cold-tolerant, the expansion of its<br />
range could lead to competition with native snails. While Turner et al. (1998) state that<br />
the two <strong>species</strong> are morphologically dist<strong>in</strong>ct and occur together <strong>in</strong> some parts of <strong>Switzerland</strong>,<br />
Anderson (2003) records the two <strong>species</strong> as synonyms.<br />
Two <strong>alien</strong> <strong>species</strong> of Planorbidae, Gyraulus parvus (Say) and Planorbarius corneus<br />
(L.), probably cause no concern, because the former is a rare <strong>species</strong> of North American<br />
orig<strong>in</strong> and the latter is a native <strong>species</strong> <strong>in</strong> Europe. It is likely that P. corneus cannot<br />
establish populations <strong>in</strong> <strong>Switzerland</strong>, because of unsuitable climatic conditions. However,<br />
it is found frequently, probably released from aquariums or transported by birds<br />
from garden ponds.<br />
6.2 Bivalves (Bivalvia)<br />
There are only three <strong>in</strong>troduced and established bivalves, but they are of concern,<br />
because of their high abundance and feed<strong>in</strong>g behaviour.<br />
The two Corbicula <strong>species</strong> (Corbicula flum<strong>in</strong>ea (O.F. Müller) and C. flum<strong>in</strong>alis (O.F.<br />
Müller)) are very similar, so that they are discussed together (see Fact Sheet). Hakenkamp<br />
and Palmer (1999) have demonstrated the strong <strong>in</strong>fluence Corbicula spp. have on<br />
ecosystem function<strong>in</strong>g by l<strong>in</strong>k<strong>in</strong>g pelagic and benthic processes as a result of their<br />
<strong>in</strong>tense filter feed<strong>in</strong>g activity.<br />
The zebra mussel Dreissena polymorpha (Pallas) (Dreissenidae) (see Fact Sheet) is<br />
one of the most widely cited case studies of a freshwater <strong>in</strong>vader. Whereas the <strong>species</strong><br />
is often considered to be beneficial <strong>in</strong> Europe, especially as a food source for div<strong>in</strong>g<br />
ducks, it is <strong>in</strong>flict<strong>in</strong>g huge costs to the USA and Canada <strong>in</strong> the Great Lakes area. The<br />
transformation of freshwater ecosystems by D. polymorpha is documented by Strayer<br />
et al. (1999) and Karateyev et al. (2002).<br />
The 19 established <strong>alien</strong> mollusc <strong>species</strong> belong to very different groups, i.e. 14 families,<br />
with only one or two <strong>species</strong> per family. However, the <strong>species</strong> can be considered<br />
as belong<strong>in</strong>g to a number of major groups. Seven <strong>species</strong> from four families of slugs<br />
(Milacidae, Limacidae, Agriolimacidae and Arionidae) have been <strong>in</strong>troduced <strong>in</strong>to
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 116<br />
<strong>Switzerland</strong>. The total number of <strong>species</strong> <strong>in</strong> these families found <strong>in</strong> <strong>Switzerland</strong> is 33,<br />
so about 21 % are <strong>in</strong>troduced. This is a rather high percentage, as will be seen below. It<br />
seems that slugs are very good at hid<strong>in</strong>g <strong>in</strong> plant material, decay<strong>in</strong>g material and other<br />
commodities and are widely transported. Today there are three established <strong>in</strong>troduced<br />
bivalves, which is about 11 % of the total of 28 <strong>species</strong> found <strong>in</strong> <strong>Switzerland</strong>. Probably<br />
six out of the 50 freshwater snail <strong>species</strong> are <strong>alien</strong> (12 %). F<strong>in</strong>ally, the large group of<br />
terrestrial snails (terrestrial molluscs exclud<strong>in</strong>g the four families of slugs mentioned<br />
above) comprise approximately 160 <strong>species</strong>, of which a mere three <strong>species</strong> (2 %) are<br />
<strong>in</strong>troduced. With the exception of the slugs, the pattern that emerges might be a random<br />
phenomenon due to small sample sizes; whereby a few <strong>in</strong>troductions of <strong>species</strong> <strong>in</strong><br />
smaller groups form a higher percentage of the total fauna. The extreme would be a<br />
randomly <strong>in</strong>troduced <strong>species</strong> from a group compris<strong>in</strong>g one <strong>species</strong>, which would form<br />
100 % of the fauna of that particular group. The total percentage of established molluscs<br />
is about 6.9 % (19 of 274 <strong>species</strong>).<br />
A comparison of the established mollusc <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> and Austria shows a<br />
very similar picture; the percentage of <strong>in</strong>troduced mollusc <strong>species</strong> is 6.9 % <strong>in</strong> <strong>Switzerland</strong><br />
and 7.6 % <strong>in</strong> Austria (Essl and Rabitsch, 2002). Of course, the total number of<br />
<strong>species</strong> differs because Austria is about twice the size of <strong>Switzerland</strong> (83,855 km 2<br />
compared to 41,285 km 2 ). About 435 mollusc <strong>species</strong> occur <strong>in</strong> Austria, <strong>in</strong>clud<strong>in</strong>g 33<br />
<strong>alien</strong>s. Even more established <strong>alien</strong> mollusc <strong>species</strong> are recorded <strong>in</strong> Germany, about 40,<br />
but five of those are mar<strong>in</strong>e (Geiter et al., 2002). There is a remarkable overlap of the<br />
<strong>alien</strong> <strong>species</strong> <strong>in</strong> the three countries, emphasiz<strong>in</strong>g the regular <strong>in</strong>troduction of some<br />
<strong>species</strong> (<strong>in</strong> some cases us<strong>in</strong>g the same frequent pathways).<br />
Most of the established <strong>alien</strong> mollusc <strong>species</strong> orig<strong>in</strong>ate with<strong>in</strong> Europe (Tab. 5.1),<br />
although the exact orig<strong>in</strong> of some Ponto–Caspian <strong>species</strong> is not known, so whether<br />
these <strong>species</strong> orig<strong>in</strong>ate <strong>in</strong> Europe or Asia may be disputable. Only five (about a quarter)<br />
of the <strong>species</strong> travelled a long distance to <strong>Switzerland</strong>. The majority have apparently<br />
profited from short-distance transport of commodities between European countries.<br />
Tab. 6.1 > Orig<strong>in</strong> of <strong>alien</strong> molluscs established <strong>in</strong> <strong>Switzerland</strong>.<br />
Orig<strong>in</strong><br />
Europe 13<br />
Asia 2<br />
North America 2<br />
<strong>Switzerland</strong> 1<br />
New Zealand 1<br />
Total 19<br />
No. <strong>species</strong>
6 > Molluscs – Mollusca 117<br />
This leads to an evaluation of the pathways by which these 19 <strong>species</strong> arrived <strong>in</strong> <strong>Switzerland</strong>.<br />
It is possible that at least some <strong>species</strong> could have arrived by several pathways,<br />
and for some <strong>species</strong> the pathways by which they were <strong>in</strong>troduced are speculative<br />
rather than proven. The most likely pathways for each <strong>species</strong> are listed <strong>in</strong> Table<br />
6.2. About 74 % (14 <strong>species</strong>) were accidentally <strong>in</strong>troduced, while the others were<br />
released for unknown reasons and from aquariums (often with the good <strong>in</strong>tention of<br />
‘free<strong>in</strong>g’ surplus animals). Five of the eight aquatic <strong>species</strong> were probably transported<br />
by boats and ships, either <strong>in</strong> ballast water or attached to hulls. The accidental terrestrial<br />
<strong>in</strong>troductions were most likely made with imported vegetables and other plant material.<br />
The impacts of the established <strong>alien</strong> molluscs are discussed <strong>in</strong> some detail and further<br />
references are given <strong>in</strong> the Fact Sheets and <strong>in</strong> the text above. Only five out of 19 <strong>species</strong><br />
can be regarded as harmless, based on present knowledge. The terrestrial slugs<br />
(and to a certa<strong>in</strong> extent snails) are ma<strong>in</strong>ly economic pests of agriculture and <strong>in</strong> gardens.<br />
However, research on environmental impacts is largely lack<strong>in</strong>g, with the exception of<br />
Arion lusitanicus displac<strong>in</strong>g the native A. rufus (L.). The situation is different <strong>in</strong> freshwater<br />
ecosystems, with demonstrated dramatic impacts of the <strong>in</strong>troduced bivalves on<br />
native biodiversity and ecosystem function<strong>in</strong>g. The <strong>in</strong>troduced bivalves are a novel life<br />
form <strong>in</strong> their new range, because of their densities and <strong>in</strong>tense filter-feed<strong>in</strong>g activities,<br />
which alter the correlation between benthic and pelagic communities. They may also<br />
have some economic impact because of their colonization of pipes and other artificial<br />
structures.<br />
In general, <strong>alien</strong> <strong>species</strong> should be treated separately from the native fauna and should<br />
not appear on a Red List of endangered <strong>species</strong>, when they are beyond doubt <strong>in</strong>troduced.<br />
This is especially true for the <strong>in</strong>tercont<strong>in</strong>ental <strong>in</strong>vader, s<strong>in</strong>ce some European<br />
<strong>species</strong> could also expand their range <strong>in</strong>to <strong>Switzerland</strong> and dist<strong>in</strong>guish<strong>in</strong>g between the<br />
two categories (<strong>alien</strong>s of European and extra-cont<strong>in</strong>ental orig<strong>in</strong>) is often challeng<strong>in</strong>g.<br />
However, <strong>species</strong> such as Physella heterostropha, a Nearctic <strong>in</strong>vader, should not<br />
appear on the Red List as potentially vulnerable (‘potentiell gefährdet’), as it does.<br />
Thus, the concept of the Red List of <strong>Switzerland</strong> should be re-addressed, with <strong>alien</strong><br />
<strong>species</strong> be<strong>in</strong>g excluded.<br />
Prevention of new mollusc <strong>in</strong>vasions is dependent on identification of the pathways of<br />
<strong>in</strong>troduction. Plant health <strong>in</strong>spection has improved recently and should be vigilant for<br />
new arrivals of slugs and snails, as well as <strong>in</strong>sects and other <strong>in</strong>vertebrates. The ballast<br />
water issue is currently be<strong>in</strong>g addressed for sea-go<strong>in</strong>g ships by the International Maritime<br />
Organization (IMO) whereby methods are evolv<strong>in</strong>g to treat ballast water. Some of<br />
these measures can also be used for ships on <strong>in</strong>land waterways. Another crucial topic is<br />
public education and awareness; when us<strong>in</strong>g a boat, boots or fish<strong>in</strong>g gear, people<br />
should take care not to transport potential hitchhikers. Aquarium owners and pet store<br />
merchants need to be made aware of potential problems aris<strong>in</strong>g from the release of<br />
pets.<br />
Several slug <strong>species</strong> are major pests <strong>in</strong> crops, and management strategies to reduce<br />
harvest losses have been implemented. Slug pellets, a bait that conta<strong>in</strong>s slug attractants<br />
and a molluscicide (e.g. metaldehyde), are frequently used. Alternative methods <strong>in</strong>-
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 118<br />
clude the use of nematodes and runner ducks (a special breed of the mallard) as biological<br />
control agents, as well as hand-pick<strong>in</strong>g. The latter approach, although very<br />
laborious, was successfully used to eradicate the giant African snail (Achat<strong>in</strong>a fulica<br />
Bowdich) <strong>in</strong> Florida, USA (Simberloff, 1996). This was a remarkable achievement.<br />
Slug fences and beer traps are often used <strong>in</strong> gardens. However, these methods are<br />
usually used to protect plants at a specific location and are not designed to reduce<br />
numbers of snails and slugs of environmental concern on a large scale.<br />
In conclusion, the freshwater ecosystems need to receive more attention and the potential<br />
impacts of <strong>alien</strong> <strong>in</strong>vaders should not be underrated. It is generally very difficult to<br />
give conclusive proof of the impacts of <strong>in</strong>vasive <strong>species</strong> on native biodiversity. However,<br />
many of the freshwater <strong>in</strong>vaders reach remarkable densities and will have impacts<br />
on natural ecosystems. These <strong>in</strong>vaders are not only molluscs, but often crustaceans and<br />
fish (see respective chapters). The boom-and-bust phenomenon (Williamson, 1996)<br />
that is frequently observed with many <strong>in</strong>vad<strong>in</strong>g <strong>species</strong> seems often to be caused <strong>in</strong><br />
European freshwater systems by the arrival of yet another <strong>in</strong>vader. Thus, this does not<br />
solve the problem, but simply shifts it to another <strong>in</strong>vader and its impacts. In a terrestrial<br />
context, the slug Arion lusitanicus is probably of greatest concern <strong>in</strong> <strong>Switzerland</strong> (and<br />
Central Europe).<br />
Tab. 6.2 > Established <strong>alien</strong> molluscs <strong>in</strong> <strong>Switzerland</strong>.<br />
Scientific name Family Year Orig<strong>in</strong> Pathway Impact Note<br />
Terrestrial snails<br />
Solatopupa similis Chondr<strong>in</strong>idae 19th Genoa, Italy Released to enrich Probably harmless Only one location near Lo-<br />
(Bruguière) century fauna carno, Tic<strong>in</strong>o<br />
Milax gagates (Drapar- Milacidae 1968 Western and Accidental with vegeta- Pest <strong>in</strong> crops and gardens Not clear whether established<br />
naud) southern<br />
Europe<br />
bles shipment? or only repeatedly <strong>in</strong>troduced<br />
Tandonia budapesten- Milacidae 1935 South-eastern Accidental with vegeta- Pest, <strong>in</strong> particular <strong>in</strong> w<strong>in</strong>ter crops, Anthropochorous – <strong>in</strong>troduced<br />
sis (Hazay) Europe bles shipment? when abundant widely by human-mediated<br />
transport<br />
Limacus flavus (L.) Limacidae 1927 Mediterranean Accidental Harmless Very rare <strong>in</strong> <strong>Switzerland</strong>, mostly<br />
synanthropic<br />
Deroceras sturanyi Agriolimacidae 1963 South-eastern Accidental Damage to plants <strong>in</strong> gardens Mostly secondary habitats<br />
(Simroth) Europe<br />
Deroceras panormita- Agriolimacidae 1982 South-western Accidental Future damage anticipated Very rare <strong>in</strong> <strong>Switzerland</strong>, only <strong>in</strong><br />
num (Lessona &<br />
Pollonera)<br />
Europe gardens and parks<br />
Boettgerilla pallens Boetgerillidae 1960 Caucasus? Accidental Predator of native slugs? Impact not known, but a<br />
Simroth predator of slugs<br />
Arion lusitanicus Mabille Arionidae 1950s Western Accidental Most serious pest <strong>in</strong> gardens and Most problematic terrestrial<br />
Europe agriculture<br />
Displaces native Arion rufus (L.)<br />
snail
6 > Molluscs – Mollusca 119<br />
Scientific name<br />
Hygromia c<strong>in</strong>ctella<br />
(Draparnaud)<br />
Cryptomphalus asperus<br />
(O.F. Müller)<br />
Freshwater snails<br />
Family Year Orig<strong>in</strong> Pathway Impact Note<br />
Hygromiidae 1824? Mediterranean Accidental Harmless Perhaps native <strong>in</strong> southern and<br />
south-western <strong>Switzerland</strong>, but<br />
<strong>in</strong>troduced to the northern parts<br />
Helicidae Before<br />
1789<br />
Viviparus ater (Mol<strong>in</strong>a) Viviparidae Before<br />
1900<br />
Potamopyrgus<br />
antipodarum (Gray)<br />
Physella acuta<br />
(Draparnaud)<br />
Physella heterostropha<br />
(Say)<br />
South-western<br />
Europe<br />
South <strong>Switzerland</strong><br />
Released Garden pest Ma<strong>in</strong>ly synanthropic, but also<br />
found <strong>in</strong> natural habitats<br />
Accidental <strong>in</strong>troduction<br />
with boat traffic<br />
Hydrobiidae 1972 New Zealand Accidental <strong>in</strong>troduction<br />
with boat traffic and<br />
birds<br />
Physidae 1848 South-western<br />
Europe<br />
Physidae Before<br />
1991<br />
Accidental release from<br />
aquariums?<br />
Gyraulus parvus (Say) Planorbidae 1994 North America Accidental with aquatic<br />
plants<br />
Planorbarius corneus<br />
(L.)<br />
Bivalves<br />
Corbicula flum<strong>in</strong>alis<br />
(O.F. Müller)<br />
Corbicula flum<strong>in</strong>ea<br />
(O.F. Müller)<br />
Probably harmless Native <strong>in</strong> southern <strong>Switzerland</strong><br />
Can drastically alter primary<br />
production<br />
Rapid expansion throughout<br />
Europe<br />
Not known Maybe of North American orig<strong>in</strong><br />
North America Accidental Competition with native snails? Expand<strong>in</strong>g <strong>in</strong> Europe<br />
Not known Rare <strong>in</strong> <strong>Switzerland</strong><br />
Planorbidae 1840 Europe Releases from aquariums<br />
Probably harmless Perhaps not established<br />
Corbiculidae 1997 Asia, <strong>in</strong>troduced<br />
via<br />
North America<br />
Corbiculidae 1997 Asia, <strong>in</strong>troduced<br />
via<br />
North America<br />
Probably ballast water Competition with native bivalves The two Corbicula <strong>species</strong> are<br />
very similar<br />
Probably ballast water Competition with native bivalves The two Corbicula <strong>species</strong> are<br />
very similar<br />
Dreissena polymorpha Dreissenidae 1850 Ponto– Probably ballast water Ecosystem eng<strong>in</strong>eer. Species of high concern<br />
(Pallas) Caspian and/or hull foul<strong>in</strong>g Overgrows native mussel <strong>species</strong><br />
region .Costs to prevent them clogg<strong>in</strong>g<br />
pipes, etc. are fairly small<br />
compared with North America
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 120<br />
References<br />
Anderson, R. (2003) Physella (Costatella) acuta Draparnaud <strong>in</strong> Brita<strong>in</strong> and<br />
Ireland – its taxonomy, orig<strong>in</strong>s and relationships to other <strong>in</strong>troduced<br />
Physidae. Journal of Conchology 38, 7–21.<br />
Essl, F. and W. Rabitsch (eds) (2002) Neobiota <strong>in</strong> Österreich. Federal<br />
Environment Agency, 432 pp.<br />
Fischer, W. and P.L. Reischütz (1998) General aspects about the<br />
slugpests. Die Bodenkultur 49 (4), 281–292.<br />
Geiter, O., Homma, S. and R. K<strong>in</strong>zelbach (2002) Bestandsaufnahme und<br />
Bewertung von Neozoen <strong>in</strong> Deutschland. Umweltbundesamt, 308 pp.<br />
Hakenkamp, C.C. and M.A. Palmer (1999) Introduced bivalves <strong>in</strong><br />
freshwater ecosystems: the impact of Corbicula on organic matter<br />
dynamics <strong>in</strong> a sandy stream. Oecologia 119, 445–451.<br />
Haynes, A., Taylor, B. J. R. and M.E. Varley (1985) The <strong>in</strong>fluence of the<br />
mobility of Potamopyrgus jenk<strong>in</strong>si (Smith, E.A.) (Prosobranchia:<br />
Hydrobiidae) on its spread. Arch. Hydrobiol. 103, 497–508.<br />
Jungbluth, J.H. (1996) E<strong>in</strong>wanderer <strong>in</strong> der Molluskenfauna von<br />
Deutschland. I. Der chorologische Befund. In: Gebhardt, H., K<strong>in</strong>zelbach, R.<br />
and S. Schmidt-Fischer (eds) Gebietsfremde Tierarten. Auswirkungen auf<br />
e<strong>in</strong>heimische Arten, Lebensgeme<strong>in</strong>schaften und Biotope;<br />
Situationsanalyse. Ecomed Verlagsgesellschaft, Landsberg, pp. 287–296.<br />
Karateyev, A.Y., Burlakova, L.E. and D.K. Padilla (2002) Impacts of zebra<br />
mussels on aquatic communities and their role as ecosystem eng<strong>in</strong>eers.<br />
In: Leppäkoski, E., Gollasch, S. and Olen<strong>in</strong>, S. (eds) <strong>Invasive</strong> aquatic<br />
<strong>species</strong> of Europe. Distribution, impacts and management. Kluwer<br />
Academic Publishers, Dordrecht, Netherlands, pp. 426–432.<br />
Kerney, M.P., Cameron, R.A.D. and J.H. Jungbluth (1983) Die<br />
Landschnecken Nord- und Mitteleuropas. Paul Parey, Hamburg and<br />
Berl<strong>in</strong>, Germany, 384 pp.<br />
Reischütz, P. (2002) Weichtiere (Mollusca). In: Essl, F. and W. Rabitsch<br />
(eds) Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, Austria, pp. 214–<br />
221.<br />
Remane, A., Storch, V. and U. Welsch (1981) Kurzes Lehrbuch der<br />
Zoologie. Gustav Fischer Verlag, Stuttgart, 537 pp.<br />
Simberloff, D. (1996) Impacts of <strong>in</strong>troduced <strong>species</strong> <strong>in</strong> the United States.<br />
Consequences 2 (2), 13–23.<br />
Strayer, D.L., Caraco, N.F., Cole, J.J., F<strong>in</strong>dlay, S. and M.L. Pace (1999)<br />
Transformation of freshwater ecosystems by bivalves: A case study of<br />
zebra mussels <strong>in</strong> the Hudson River. Bioscience 49, 19–27.<br />
Turner, H., Kuiper, J.G.J., Thew, N., Bernasconi, R., Rüetschi, J.,<br />
Wüthrich, M. and M. Gosteli (1998) Mollusca, Atlas. Fauna Helvetica.<br />
CSCF, Neuchâtel, 527 pp.<br />
Williamson, M (1996) Biological <strong>in</strong>vasions. Chapman & Hall, London,<br />
We<strong>in</strong>heim, New York, Tokyo, Melbourne, Madras, 244 pp.
7 > Other selected <strong>in</strong>vertebrate groups 121<br />
7 > Other selected <strong>in</strong>vertebrate groups<br />
Prepared by Rüdiger Wittenberg and Marc Kenis<br />
This chapter gives only a few examples of <strong>alien</strong> <strong>species</strong> from groups which can cause<br />
problems for the environment not dealt with <strong>in</strong> other chapters of this report. It is not<br />
possible to give comprehensive lists of most of these <strong>in</strong>vertebrate groups. Therefore,<br />
the objective is to give a brief overview of potential problematic <strong>species</strong> for biodiversity<br />
and ecosystems <strong>in</strong> <strong>Switzerland</strong>. As mentioned before, there is a great need for<br />
more taxonomic work on these groups, not only the <strong>alien</strong> <strong>species</strong>, but the native <strong>species</strong><br />
as well. Based on future work, more comprehensive lists of some groups may be added<br />
<strong>in</strong> the future to complete the knowledge of established <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>.<br />
7.1 Nematodes – Nemathelm<strong>in</strong>thes<br />
Nematodes are a large but little-known group of t<strong>in</strong>y worms. There is no comprehensive<br />
checklist of <strong>species</strong> <strong>in</strong> this group for <strong>Switzerland</strong>, but many <strong>species</strong> are of great<br />
economic importance as pests <strong>in</strong> agriculture and forestry. Species exclusively caus<strong>in</strong>g<br />
economic damage are not part of the report, s<strong>in</strong>ce they are the best-known <strong>species</strong> <strong>in</strong><br />
this group; the Swiss Federal Research Stations are actively work<strong>in</strong>g on them, and<br />
there are other sources deal<strong>in</strong>g with them, e.g. CABI (2001) lists the follow<strong>in</strong>g 11<br />
problematic nematodes for <strong>Switzerland</strong>: Globodera pallida (Stone) Behrens, G. rostochiensis<br />
(Wollenweber) Behrens, Heterodera avenae Wollenweber, H. schachtii<br />
A.Schmidt, Longidorus elongatus (de Man) Micoletzky, Meloidogyne arenaria (neal)<br />
Chitwood, M. hapla Chitwood, Pratylenchus penetrans (Cobb) Filipjev & Schuurmans,<br />
Punctodera punctata (Thorne) Mulvey & Stone, Xiph<strong>in</strong>ema diversicaudatum<br />
(Micoletzky) Thorne, and X. <strong>in</strong>dex Thorne & Allen.<br />
The p<strong>in</strong>e wood nematode (Bursaphelenchus xylophilus (Ste<strong>in</strong>er & Buhrer) Nickle)<br />
(see Fact Sheet) is an example of a <strong>species</strong> that has recently been found <strong>in</strong> Europe and<br />
is considered to be a potential threat to P<strong>in</strong>us spp. here. Thus, its wider spread with<strong>in</strong><br />
Europe should be prevented.<br />
The nematode Anguillicola crassus Kuwahara, Niimi & Itagaki (Anguillicolidae) is an<br />
example of a nematode that is hav<strong>in</strong>g an extensive impact by attack<strong>in</strong>g a fish <strong>species</strong>.<br />
This nematode is a parasite of eel <strong>species</strong>. In Europe the native Anguilla anguilla L. is<br />
under threat follow<strong>in</strong>g the <strong>in</strong>vasion of the nematode. The nematode was accidentally<br />
<strong>in</strong>troduced with imports of live Asian eels to Europe <strong>in</strong> 1982 (Konecny et al., 2002).<br />
Besides the negative impact on natural populations of the European eel, it affects<br />
Europe’s important fish<strong>in</strong>g <strong>in</strong>dustries. The European eel is native <strong>in</strong> the river systems
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 122<br />
empty<strong>in</strong>g <strong>in</strong>to the Atlantic, as its life history <strong>in</strong>volves spawn<strong>in</strong>g grounds <strong>in</strong> the Sargasso<br />
Sea. It has itself been <strong>in</strong>troduced to some parts of Europe, e.g. the Danube bas<strong>in</strong>.<br />
The construction of the Rh<strong>in</strong>e–Danube Canal opened the way from the Rh<strong>in</strong>e, where it<br />
is native, to the Danube bas<strong>in</strong>.<br />
Another nematode was accidentally <strong>in</strong>troduced <strong>in</strong>to Europe together with its North<br />
American host, i.e. Baylisascaris procyonis (Stefanski & Zarnowski) with the raccoon,<br />
Procyon lotor (L.) (see mammal chapter). This <strong>species</strong> is of concern for human health.<br />
7.2 Flatworms – Turbellaria, Plathelm<strong>in</strong>thes<br />
In the group Turbellaria (part of the Plathelm<strong>in</strong>thes), one <strong>species</strong> is an abundant <strong>alien</strong><br />
<strong>in</strong>habitant of lakes and rivers, <strong>in</strong>clud<strong>in</strong>g the Rh<strong>in</strong>e <strong>in</strong> <strong>Switzerland</strong>. The predatory<br />
aquatic flatworm Dugesia tigr<strong>in</strong>a (Gerard) is abundant and widespread <strong>in</strong> stand<strong>in</strong>g and<br />
slow-mov<strong>in</strong>g water bodies <strong>in</strong> Europe (Pöckl and Rabitsch, 2002). It was probably<br />
<strong>in</strong>troduced on aquarium plants or fish from North America around 1900. Although this<br />
<strong>species</strong> is found <strong>in</strong> high densities, a negative impact has not (yet) been shown.<br />
Other flatworms of environmental concern are terrestrial. The New Zealand flatworm<br />
(Artioposthia triangulate (Dendy)) (see Fact Sheet) serves as an example of several<br />
predatory flatworm <strong>species</strong> <strong>in</strong>troduced <strong>in</strong>to Europe from New Zealand and Australia.<br />
Native earthworms are a major prey for these flatworms. Earthworms play an important<br />
role <strong>in</strong> nutrient cycl<strong>in</strong>g <strong>in</strong> soil, so that their reduction could lead to ecosystem<br />
changes. They are also a significant element of vertebrate prey.<br />
7.3 Segmented worms – Annelida<br />
The Ponto–Caspian <strong>in</strong>vaders <strong>in</strong> European waters are an <strong>in</strong>terest<strong>in</strong>g group – see the<br />
Crustacea Chapter for more <strong>in</strong>formation. The annelid polychaete worm Hypania<br />
<strong>in</strong>valida (Grube) has become very numerous <strong>in</strong> the sandy sediment of the Rh<strong>in</strong>e, where<br />
it burrows its tubes vertically <strong>in</strong>to the mud (Rey and Ortlepp, 2002; Van der Velde et<br />
al., 2002). The impact of this abundant <strong>species</strong> has not been evaluated. It is approximately<br />
1–2 cm long and can reach densities of about 10,000 <strong>in</strong>dividuals per square<br />
metre. The <strong>species</strong> reached the Rh<strong>in</strong>e after the open<strong>in</strong>g of the Rh<strong>in</strong>e–Danube canal,<br />
probably <strong>in</strong> the ballast tanks of ships.<br />
Another possibly harmless <strong>species</strong> <strong>in</strong> the group Annelida, but an oligochaete worm<br />
(Tubificidae), is the large (up to 20 cm long) Branchiura sowerbyi Beddard, which<br />
orig<strong>in</strong>ates <strong>in</strong> south-eastern Asia. It was probably <strong>in</strong>troduced on tropical aquatic plants<br />
at the beg<strong>in</strong>n<strong>in</strong>g of the 20 th century. At first it occurred only <strong>in</strong> greenhouses, but it<br />
adapted to the colder climate and is now found <strong>in</strong> many slow-mov<strong>in</strong>g rivers (Pöckl and<br />
Rabitsch, 2002). It lives <strong>in</strong> self-made tubes <strong>in</strong> muddy soil and feeds on detritus.
7 > Other selected <strong>in</strong>vertebrate groups 123<br />
A member of a third group of the Annelida is the leech Caspiobdella fadejewi (Epshte<strong>in</strong>)<br />
(Hirund<strong>in</strong>ea; Piscicolidae), which can be found <strong>in</strong> low densities <strong>in</strong> the Rh<strong>in</strong>e<br />
between Basel and the Bodensee. This <strong>species</strong>, also from the Ponto–Caspian system, is<br />
an ectoparasite of various fish <strong>species</strong>.<br />
7.4 Centipedes and millipedes – Myriapoda<br />
A neglected group <strong>in</strong> this report is the Myriapoda, <strong>in</strong>clud<strong>in</strong>g the Diplopoda and the<br />
Chilopoda. It is impossible to give a comprehensive list. In particular the former group,<br />
as plant and detritus feeders, are probably regularly <strong>in</strong>troduced with plant and soil<br />
material. Some tropical or subtropical <strong>species</strong> might be restricted to greenhouses.<br />
However, the number of established <strong>species</strong> <strong>in</strong> the wild is likely to be small (probably<br />
less than ten <strong>species</strong>) and none is of particular threat to native biodiversity and ecosystems.<br />
References<br />
CABI (2001) Crop Protection Compendium. CD-ROM. CAB International,<br />
Wall<strong>in</strong>gford, UK.<br />
Konecny, R., Schabussova, I. and H. Sattmann (2002) ”Helm<strong>in</strong>then“ –<br />
Neozoen der Fische Österreichs. In: Essl, F. and W. Rabitsch (eds)<br />
Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, pp. 221–224.<br />
Pöckl, M. and W. Rabitsch (2002) Ausgewählte aquatische Neozoen<br />
(Cnidaria, Plathelm<strong>in</strong>thes, Kamptozoa, Annelida, Isopoda). In: Essl, F. and<br />
W. Rabitsch (eds) Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, pp.<br />
228–239.<br />
Rey, P. and J. Ortlepp (2002) Koord<strong>in</strong>ierte biologische Untersuchungen<br />
am Hochrhe<strong>in</strong> 2000; Makro<strong>in</strong>vertebraten. BUWAL Schriftenreihe Umwelt<br />
Nr. 345, Gewässerschutz, 98 pp.<br />
Van der Velde, G., Nagelkerken, I., Rajapogal, S. and A. Bij de Vaate<br />
(2002) Invasions by <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>in</strong>land freshwater bodies <strong>in</strong> Western<br />
Europe: the Rh<strong>in</strong>e delta. In: Leppäkoski, E., Gollasch, S. and Olen<strong>in</strong>, S.<br />
(eds) <strong>Invasive</strong> aquatic <strong>species</strong> of Europe. Distribution, impacts and<br />
management. Kluwer Academic Publishers, Dordrecht, pp. 426–432.<br />
Weidema, I.R. (ed) (2000) Introduced <strong>species</strong> <strong>in</strong> the Nordic countries.<br />
Nordic Council of M<strong>in</strong>isters, Copenhagen, 242 pp.<br />
Welch, D., Carss, D.N., Gornall, J., Manchester, S.J., Marquiss, M.,<br />
Preston, C.D., Telfer, M.G., Arnold, H. and J. Holbrook (2001) An audit of<br />
<strong>alien</strong> <strong>species</strong> <strong>in</strong> Scotland. Scottish Natural Heritage Review No 139, 225<br />
pp.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 124<br />
8 > Lichens (Lichen-form<strong>in</strong>g fungi)<br />
Prepared by Rüdiger Wittenberg<br />
There are no lichens known to be <strong>in</strong>troduced and established <strong>in</strong> <strong>Switzerland</strong><br />
(C. Scheidegger, pers. comm.). Lichens, with their extremely slow growth, seem illadapted<br />
to human-mediated transport and the colonization of new regions. There is no<br />
doubt that <strong>species</strong> must be <strong>in</strong>troduced with timber, etc., but the step from <strong>in</strong>troduction<br />
to establishment has apparently not been achieved. Breuss (2002) mentions Anisomeridium<br />
polypori (Ellis & Everh.) M.E. Barr as a neomycete for Austria. However,<br />
recently described new <strong>species</strong> are not necessarily <strong>in</strong>troduced. They could also have<br />
been overlooked <strong>in</strong> the past, <strong>in</strong> particular as floristic research <strong>in</strong> microlichens does not<br />
have a long history and is rather <strong>in</strong>complete. Changes <strong>in</strong> the environment can also<br />
favour some previously rare <strong>species</strong>, so that they become more abundant and widespread,<br />
feign<strong>in</strong>g a new arrival. Aptroot (1999) doubts the neomycete character of A.<br />
polypori, <strong>in</strong>stead assum<strong>in</strong>g a wide natural distribution.<br />
References<br />
Aptroot, A. (1999) Notes on taxonomy, distribution and ecology of<br />
Anisomeridium polypori. Lichenologist 31 (6), 641–642.<br />
Breuss, O. (2002) Flechten. In: Essl, F. and W. Rabitsch (eds) Neobiota <strong>in</strong><br />
Österreich. Umweltbundesamt, Wien, pp. 214–221.
9 > Fungi and a selected bacterium 125<br />
9 > Fungi and a selected bacterium<br />
Prepared by Rüdiger Wittenberg and Marc Kenis<br />
Fungi are an often-neglected group and it is not possible to compile a list of neomycetes<br />
for <strong>Switzerland</strong>, let alone to list the fungi occurr<strong>in</strong>g <strong>in</strong> <strong>Switzerland</strong>. It is a group<br />
where the percentage of undescribed <strong>species</strong> is exceptionally high. In this report,<br />
<strong>species</strong> which are of concern for native biodiversity and economics alike are summarized.<br />
Species exclusively caus<strong>in</strong>g economic damage are not part of the report, s<strong>in</strong>ce<br />
they are the best-known <strong>species</strong>; the Swiss Federal Research Stations are responsible<br />
for work<strong>in</strong>g on them, and there are also other sources, such as university research<br />
departments, deal<strong>in</strong>g with them. Two examples of excellent sources are:<br />
> The Alert List of the European and Mediterranean Plant Protection Organization<br />
(EPPO). The purpose of the Alert List is to draw attention to certa<strong>in</strong> pests that may<br />
present a risk to member countries, thereby function<strong>in</strong>g as an early warn<strong>in</strong>g system:<br />
http://www.eppo.org/QUARANTINE/Alert_List/alert_list.html<br />
> The Crop Protection Compendium (CPC) (see e.g. CABI, 2004) lists, for example,<br />
104 fungi for <strong>Switzerland</strong> and provides Fact Sheets <strong>in</strong>corporat<strong>in</strong>g exhaustive <strong>in</strong>formation<br />
on the <strong>species</strong>. It is also available on the Internet at:<br />
http://www.cabi.org/compendia/cpc/<strong>in</strong>dex.htm<br />
These sources give other l<strong>in</strong>ks as well, so that a wealth of <strong>in</strong>formation is available for<br />
the <strong>species</strong> of economic importance.<br />
Six fungal <strong>species</strong> of immediate threat to the native biodiversity are discussed <strong>in</strong> this<br />
section. There is also one bacterium, which is pathogenic to plants, <strong>in</strong>cluded <strong>in</strong> the text<br />
and a Fact Sheet.<br />
Chestnut blight (Cryphonectria parasitica (Murrill) Barr) (see Fact Sheet) was <strong>in</strong>troduced<br />
<strong>in</strong>to North America and Europe. In North America chestnut blight has been an<br />
ecological disaster. It changed the tree composition of the eastern forests completely by<br />
remov<strong>in</strong>g one of the dom<strong>in</strong>ant trees, Castanea dentata (Marshall). Fortunately, it<br />
seems that the European congeneric is less susceptible to the disease, although it has<br />
suffered and tree composition is alter<strong>in</strong>g, <strong>in</strong> particular <strong>in</strong> the Tic<strong>in</strong>o.<br />
Another tree-attack<strong>in</strong>g fungus, caus<strong>in</strong>g tremendous ecosystem changes <strong>in</strong> Europe, is<br />
Dutch elm disease (Ceratocystis ulmi (Buisman) C.Moreau and C. novo-ulmi (Brassier))<br />
(see Fact Sheet). Mature elm (Ulmus spp.) trees have disappeared from the landscape<br />
<strong>in</strong> many regions. C. novo-ulmi seems to have arrived decades after C. ulmi and is<br />
replac<strong>in</strong>g it <strong>in</strong> many parts of Europe, e.g. Austria (Kirisits et al., 2001). Re<strong>in</strong>hardt et al.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 126<br />
(2003) estimate that Germany <strong>in</strong>curs annual costs of about €5 million for the removal<br />
and replacement of trees, the lost value of dead trees, and additional expenditure of<br />
plant<strong>in</strong>g resistant varieties.<br />
While Phytophthora querc<strong>in</strong>a Jung et al. (see Fact Sheet) is a rather recent <strong>in</strong>vader (or<br />
at least attention has been drawn to it by its recent impact), P. ramorum Werres, de<br />
Cock & Man <strong>in</strong>’t Veld (sudden oak death) (see Fact Sheet) is already well-known<br />
from its devastat<strong>in</strong>g effect <strong>in</strong> North America. In Europe it is still only a problem <strong>in</strong><br />
nurseries, but it can be expected to <strong>in</strong>fest native forests <strong>in</strong> the not-too-distant future.<br />
Many woody hosts have been shown to be susceptible to sudden oak death and it has<br />
been recorded from <strong>Switzerland</strong> (He<strong>in</strong>iger and Stadler, 2003).<br />
The crayfish plague (Aphanomyces astaci (Schikora)) (see Fact Sheet) is one of the<br />
most devastat<strong>in</strong>g fungi attack<strong>in</strong>g European wildlife. The fungus was <strong>in</strong>troduced with<br />
North American crayfish <strong>species</strong>, which are asymptomatic carriers of the disease, <strong>in</strong>to<br />
Europe. The European crayfish <strong>species</strong> are highly susceptible and almost 100 % die<br />
with<strong>in</strong> two weeks of <strong>in</strong>fection. There are regular outbreaks <strong>in</strong> European populations<br />
result<strong>in</strong>g <strong>in</strong> total collapse (Voglmayr and Krisai-Greilhuber, 2002).<br />
The bacterium Erw<strong>in</strong>ia amylovora (Burrill) W<strong>in</strong>slow et al. (see Fact Sheet) is the<br />
causal agent of fire blight, which was first found on Cotoneaster <strong>species</strong> and is now<br />
widely distributed <strong>in</strong> <strong>Switzerland</strong> (Hasler et al., 2002).<br />
An <strong>in</strong>terest<strong>in</strong>g case of a mutual relationship between two <strong>in</strong>vaders may be the association<br />
between mycorrhizal fungi and plant <strong>species</strong>. There is some evidence that <strong>alien</strong><br />
mycorrhiza can help their <strong>alien</strong> host plant to become a weed <strong>in</strong> the <strong>in</strong>troduced range<br />
(e.g. Crawley, 1993; Harr<strong>in</strong>gton et al., 1998). In that way, an <strong>in</strong>troduced mycorrhiza<br />
might have an <strong>in</strong>direct detrimental effect on native biodiversity.<br />
In can be concluded that although quarant<strong>in</strong>e measures are successfully implemented <strong>in</strong><br />
<strong>Switzerland</strong>, non-crop plants need to get more attention, e.g. plants <strong>in</strong> the nursery<br />
<strong>in</strong>dustry. The few examples described here <strong>in</strong>dicate the enormous impacts diseases can<br />
have. Some <strong>in</strong>troduced diseases are completely chang<strong>in</strong>g entire ecosystems by elim<strong>in</strong>at<strong>in</strong>g<br />
important key <strong>species</strong>. Moreover, there are human diseases not mentioned <strong>in</strong><br />
this report with global impact, such as AIDS (a viral disease).<br />
The Fact Sheets are presented after the references, <strong>in</strong> alphabetical order for ease of<br />
location of specific sheets.
9 > Fungi and a selected bacterium 127<br />
References<br />
CABI (2004) Crop Protection Compendium. CD-ROM. CAB International,<br />
Wall<strong>in</strong>gford, UK.<br />
Crawley, M.J. (1993) Succeed<strong>in</strong>g <strong>in</strong> the sand dunes. Nature 362, 17–18.<br />
Harr<strong>in</strong>gton, K.C., Hodder, L.M. and H.A. Outred (1998) Biology and control<br />
of pypgrass. Proceed<strong>in</strong>gs, 51 st New Zealand Plant Protection Conference,<br />
pp. 255–259.<br />
Hasler, T., Schaerer, H.J., Holliger, E., Vogelsanger, J., Vignutelli, A. and<br />
B. Schoch (2002) Fire blight situation <strong>in</strong> <strong>Switzerland</strong>. Acta Horticulturae<br />
590, 73–79.<br />
He<strong>in</strong>iger, U. and Stadler, B. (2003) Gefährliche Quarantänekrankheit<br />
gefunden. Phytophthora ramorum jetzt erstmals auch <strong>in</strong> der Schweiz. Der<br />
Gartenbau 51/52/2003, pp. 10–12.<br />
Kirisits, T., Krumböck, S. Konrad, H., Pennerstorfer, J. and E.<br />
Halmschlager (2001) Untersuchungen über das Auftreten der Erreger der<br />
Holländischen Ulmenwelke <strong>in</strong> Österreich. Forstwissenschaftliches<br />
Zentralblatt 120, 231–241.<br />
Re<strong>in</strong>hardt, F., Herle, M. Bastiansen, F. and B. Streit (2003) Economic<br />
impact of the spread of <strong>alien</strong> <strong>species</strong> <strong>in</strong> Germany. Federal Environmental<br />
Agency of Germany, 190 pp.<br />
Voglmayr, H. and I. Krisai-Greilhuber (2002) Pilze. In: Essl, F. and W.<br />
Rabitsch (eds) Neobiota <strong>in</strong> Österreich. Umweltbundesamt, Wien, pp. 214–<br />
221.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 128<br />
10 > Plants – Planta<br />
Prepared by André Gassmann and Ewald Weber<br />
10.1 Introduction and term<strong>in</strong>ology<br />
The expansion of <strong>alien</strong> plants with<strong>in</strong> Central Europe began with the <strong>in</strong>troduction of<br />
agriculture about 7,000 years ago and the subsequent spread of weeds. In Europe, as <strong>in</strong><br />
most regions of the world, the number of <strong>alien</strong> plant <strong>species</strong> has <strong>in</strong>creased considerably<br />
<strong>in</strong> the past 200 years as a result of <strong>in</strong>creas<strong>in</strong>g trade, tourism and disturbance. The<br />
<strong>in</strong>creas<strong>in</strong>g number of naturalized <strong>alien</strong> plant <strong>species</strong> with negative impacts on plant<br />
communities is viewed as a major component of global change. Successful <strong>in</strong>vaders<br />
can affect the <strong>in</strong>vaded communities <strong>in</strong> various ways, e.g. reduc<strong>in</strong>g the local diversity,<br />
driv<strong>in</strong>g rare native <strong>species</strong> to ext<strong>in</strong>ction (e.g. by competition or hybridization), chang<strong>in</strong>g<br />
habitat structures and ecosystem function<strong>in</strong>g, or <strong>in</strong>creas<strong>in</strong>g erosion. Plants are<br />
particularly notorious <strong>in</strong>vaders, s<strong>in</strong>ce they are capable of chang<strong>in</strong>g the food web at the<br />
base, which can ripple through the entire ecosystem. In <strong>Switzerland</strong>, with the exception<br />
of the Alps, wildlife and areas of conservation value are usually restricted to small<br />
areas, surrounded by heavily disturbed habitats or urban areas. In such small areas,<br />
<strong>in</strong>vasive plant <strong>species</strong> pose additional threats to the native plant and animal diversity.<br />
Moreover, new plant <strong>in</strong>vaders <strong>in</strong> <strong>Switzerland</strong> and <strong>in</strong> Europe can also affect human<br />
health (e.g. Ambrosia artemisiifolia L. and Heracleum mantegazzianum Sommier et<br />
Levier) or are a potential threat to the agro-economy (e.g. Senecio <strong>in</strong>aequidens DC.).<br />
This report gives an overview of <strong>alien</strong> and <strong>in</strong>vasive plant <strong>species</strong> with regard to the<br />
Swiss flora. Some of the ecological and biological characteristics of <strong>alien</strong> plants <strong>in</strong><br />
<strong>Switzerland</strong> are discussed and a list of plant <strong>in</strong>vaders or potential <strong>in</strong>vaders is provided.<br />
In this context, it is important to note that different orig<strong>in</strong>al sources may cite different<br />
numbers of <strong>alien</strong> plant <strong>species</strong>. For example, Moser et al. (2002) list 350 neophytes. A<br />
number of these <strong>species</strong> are given a different status <strong>in</strong> Lauber and Wagner (1998). In<br />
addition, the <strong>species</strong> lists are not identical. For example, Moser et al. (2002) list two<br />
ferns as neophytes (Cyrtomium falcatum K. Presl, C. fortunei J. Sm.) which are not<br />
<strong>in</strong>cluded <strong>in</strong> Lauber and Wagner (1998). The same holds for Crataegus l<strong>in</strong>dmanii<br />
Hrabetova and C. rhipidophylla Gand. In contrast, <strong>species</strong> like Cotoneaster bullata<br />
Bois, C. dammeri Schneider, Helianthus rigidus (Cass<strong>in</strong>i) and Paspalum dilatatum<br />
Poiret are not <strong>in</strong>cluded <strong>in</strong> Moser et al. (2002). Exclud<strong>in</strong>g cultivated <strong>species</strong> that occur<br />
rarely as subspontaneous <strong>species</strong> (‘C’ plants <strong>in</strong> Moser et al. (2002)), we found <strong>in</strong> total<br />
over 70 <strong>alien</strong> <strong>species</strong> which are <strong>in</strong>cluded <strong>in</strong> either one of the two lists but not <strong>in</strong> the<br />
other. Most of the variation orig<strong>in</strong>ates from whether cultivated <strong>species</strong> are considered<br />
to be subspontaneous or not. In addition, another 60 <strong>species</strong> of European orig<strong>in</strong> are<br />
controversial with regard to their <strong>alien</strong> status <strong>in</strong> <strong>Switzerland</strong>. Thus, different authors
10 > Plants – Planta 129<br />
have different approaches to the treatment of <strong>species</strong>, and there is no ultimately correct<br />
source. However, either source can be used to seek general patterns <strong>in</strong> geographic<br />
orig<strong>in</strong>s, life form distribution, or the numbers of neophytes present. Because Moser et<br />
al. (2002) do not provide standardized <strong>in</strong>formation on plant status and plant orig<strong>in</strong>, we<br />
used the <strong>in</strong>formation provided by Lauber and Wagner (1998) to seek these general<br />
patterns.<br />
There is much variation <strong>in</strong> the def<strong>in</strong>ition of some of the terms presented below. In<br />
particular, there is controversy about the def<strong>in</strong>ition of ‘<strong>in</strong>vasive <strong>species</strong>’ <strong>in</strong> the literature.<br />
The term ‘<strong>in</strong>vasive <strong>species</strong>’ often refers exclusively to <strong>species</strong> penetrat<strong>in</strong>g <strong>in</strong>to<br />
natural and semi-natural habitats although it is often difficult to clearly separate sem<strong>in</strong>atural<br />
from human-made habitats. The terms ‘weeds’ and ‘weedy <strong>species</strong>’ are consequently<br />
devoted to plants caus<strong>in</strong>g problems <strong>in</strong> managed areas. Some authors call any<br />
<strong>alien</strong> plant <strong>species</strong> that spreads spontaneously <strong>in</strong> the <strong>in</strong>troduced range an <strong>in</strong>vasive<br />
<strong>species</strong>, irrespective of whether the <strong>species</strong> has harmful effects or not. For practical<br />
reasons, such a broad def<strong>in</strong>ition is not useful.<br />
In this chapter, the term ‘<strong>in</strong>vasive <strong>species</strong>’ always refers to plants of <strong>alien</strong> orig<strong>in</strong>, while<br />
the term ‘weed’ implies an <strong>in</strong>digenous <strong>species</strong>. Alien <strong>species</strong> may become <strong>in</strong>vasive<br />
primarily <strong>in</strong> human-made habitats or agricultural land and thus have economic rather<br />
than ecological effects (e.g. Ambrosia artemisiifolia and Cyperus esculentus L.). It<br />
must be stressed however that most <strong>in</strong>vasive plants are of greater concern <strong>in</strong> natural or<br />
semi-natural areas. It should be po<strong>in</strong>ted out also that the spread of many naturalized<br />
<strong>species</strong> is primarily <strong>in</strong> highly disturbed areas of low ecological (or economic) value but<br />
this situation may change for many plant <strong>species</strong> <strong>in</strong> the future. In contrast, native<br />
<strong>species</strong> can pose problems by becom<strong>in</strong>g abundant <strong>in</strong> conservation areas <strong>in</strong>fluenced by<br />
human activities, e.g. Phragmites australis (Cavanilles) or Rubus spp. This report does<br />
not consider native <strong>species</strong> that are weeds <strong>in</strong> agricultural land or may be of concern <strong>in</strong><br />
natural or semi-natural areas.<br />
The term<strong>in</strong>ology presented below has been adapted from Richardson et al. (2000) and<br />
Weber (1999a).<br />
> Alien (non-native, non-<strong>in</strong>digenous, <strong>in</strong>troduced) plants: plant taxa (<strong>species</strong>, sub<strong>species</strong><br />
or lower taxon) <strong>in</strong> a given area whose presence there is due to <strong>in</strong>tentional or<br />
accidental <strong>in</strong>troduction as a result of human activity. Unless specified, cultivated<br />
plants that have not escaped from cultivation are not treated as <strong>alien</strong> <strong>species</strong>.<br />
> Native (<strong>in</strong>digenous) plants: plant taxa occurr<strong>in</strong>g with<strong>in</strong> their natural range and<br />
dispersal potential (i.e. with<strong>in</strong> the range they occupy naturally or could occupy without<br />
direct or <strong>in</strong>direct <strong>in</strong>troduction or care by humans).<br />
> Neophyte plants (or neophytes): <strong>alien</strong> plants <strong>in</strong>troduced dur<strong>in</strong>g modern times (after<br />
1500 A.D.) and which have become naturalized. Archeophyte plants (or archeophytes)<br />
are those that were <strong>in</strong>troduced before 1500 A.D. This report does not treat<br />
the two groups separately.<br />
> Casual (transient, ephemeral) plants: <strong>alien</strong> plants that may flourish and even<br />
reproduce occasionally <strong>in</strong> an area, but which do not form self-replac<strong>in</strong>g populations,<br />
and which rely on repeated <strong>in</strong>troductions or habitat disturbance for their persistence.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 130<br />
> Adventive plants: casual <strong>alien</strong> plants that have been accidentally <strong>in</strong>troduced as a<br />
result of human activity.<br />
> Subspontaneous plants: casual <strong>alien</strong> plants escaped from cultures.<br />
> Naturalized plants: Alien plants that reproduce consistently and susta<strong>in</strong> populations<br />
over many life cycles without direct <strong>in</strong>tervention by humans, (or <strong>in</strong> spite of human<br />
<strong>in</strong>tervention); they often recruit offspr<strong>in</strong>g freely, usually close to adult plants, and do<br />
not necessarily <strong>in</strong>vade their habitat.<br />
> <strong>Invasive</strong> plants (plant <strong>in</strong>vaders): Naturalized plants that produce reproductive<br />
offspr<strong>in</strong>g, often <strong>in</strong> very large numbers, at considerable distances from parent plants,<br />
and thus have the potential to spread over a considerable area. <strong>Invasive</strong> plants can<br />
affect the <strong>in</strong>vaded natural or semi-natural communities <strong>in</strong> various ways. <strong>Invasive</strong><br />
plants can also affect human-made habitats and have direct economic effects. The<br />
term environmental weeds is sometimes used for those <strong>in</strong>vasive plants hav<strong>in</strong>g an<br />
impact <strong>in</strong> natural areas and semi-natural areas; and the term <strong>alien</strong> weeds is sometimes<br />
used for those <strong>alien</strong> plants that are weedy <strong>in</strong> managed habitats, such as agriculture.<br />
> Weeds: native plants that grow <strong>in</strong> sites where they are not wanted and which have<br />
detectable economic or environmental effects.<br />
10.2 The native and <strong>alien</strong> flora of <strong>Switzerland</strong><br />
The list of <strong>alien</strong> <strong>species</strong> present <strong>in</strong> <strong>Switzerland</strong> (Table 10.6) <strong>in</strong>cludes both archaeophytes<br />
and neophytes. However, <strong>in</strong> this list, the neophytes of Flora Helvetica (Lauber<br />
and Wagner, 1998) are marked as such. A few <strong>species</strong> that have become naturalized<br />
recently (e.g. Ludwigia grandiflora (Michaux) and Lysichiton americanus Hultén & St.<br />
John) have been added to the <strong>alien</strong> <strong>species</strong> amongst the 3000 plant taxa listed <strong>in</strong> Flora<br />
Helvetica. The plant status and life form for each <strong>species</strong> have been extracted from the<br />
<strong>in</strong>formation provided by Lauber and Wagner (1998).<br />
The Swiss flora of vascular plants <strong>in</strong>cludes some 162 families and over 3000 taxa<br />
(<strong>species</strong>, sub<strong>species</strong> and lower taxa). Of these, 20 plant families and 84 plant taxa<br />
belong to the ferns and fern allies (Pteridophyta) while the other families and taxa<br />
belong to the flower<strong>in</strong>g plants (Spermatophyta).<br />
The Swiss flora comprises 2505 native <strong>species</strong> belong<strong>in</strong>g to 136 families (Table 10.1;<br />
exclud<strong>in</strong>g sub<strong>species</strong> and lower taxa). From the 470 taxa of <strong>alien</strong> orig<strong>in</strong> recorded <strong>in</strong><br />
<strong>Switzerland</strong>, over 100 <strong>species</strong> are cultivated <strong>species</strong> that are not, or rarely, found <strong>in</strong> the<br />
environment. The 362 rema<strong>in</strong><strong>in</strong>g <strong>alien</strong> <strong>species</strong> have become subspontaneous, adventive<br />
or naturalized. These <strong>alien</strong> <strong>species</strong>, which represent about 12.6 % of the flora of <strong>Switzerland</strong>,<br />
are discussed below.<br />
This percentage is similar to that observed <strong>in</strong> neighbour<strong>in</strong>g countries, e.g. 9.1 % <strong>in</strong><br />
Austria, 10.2 % <strong>in</strong> France, but much less than <strong>in</strong> countries <strong>in</strong> North America, e.g. 28 %<br />
<strong>in</strong> Canada, or on islands, e.g. 47 % <strong>in</strong> New Zealand (Heywood, 1989). The density of<br />
<strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> (i.e. the number of <strong>alien</strong> <strong>species</strong> per log country size <strong>in</strong>
10 > Plants – Planta 131<br />
10.3<br />
square kilometres) is 78.4 and slightly higher than <strong>in</strong> several other European countries.<br />
This may be due to the topography and diversity of climates encountered <strong>in</strong> <strong>Switzerland</strong><br />
which allow plant <strong>species</strong> as different as Agave americana L. and Reynoutria<br />
japonica Houttuyn to become naturalized (see also Weber, 1999a, b). France, which<br />
offers an even wider range of habitats and climates, has a density of <strong>alien</strong> <strong>species</strong> of<br />
87.1 (Heywood, 1989), whereas <strong>in</strong> Austria, the density of <strong>alien</strong> <strong>species</strong> is 60.9.<br />
Accord<strong>in</strong>g to Lauber and Wagner (1998), the group conta<strong>in</strong><strong>in</strong>g ferns and its allies does<br />
not <strong>in</strong>clude any <strong>alien</strong> <strong>species</strong> (Table 10.1). There is a high proportion of <strong>alien</strong> gymnosperm<br />
<strong>species</strong> (28.6 %), i.e. four <strong>alien</strong> gymnosperm <strong>species</strong> out of 14 recorded. The<br />
frequency of <strong>alien</strong> <strong>species</strong> <strong>in</strong> the dicotyledons and monocotyledons is quite similar, i.e.<br />
13.5 % and 11.0 %, respectively.<br />
Tab. 10.1 > Synopsis of the native and <strong>alien</strong> flora of <strong>Switzerland</strong>: number of <strong>species</strong> (% of total).<br />
Pteridophyta Spermatophyta Total<br />
Gymnosperms Dicotyledons Monocotyledons<br />
Native <strong>species</strong>*<br />
84 (3.4)<br />
10 (0.4)<br />
1879 (75.0)<br />
532 (21.2)<br />
2505 (100)<br />
Families<br />
20 (14.7)<br />
4 (2.9)<br />
93 (68.4)<br />
19 (14.0)<br />
136 (100)<br />
Alien <strong>species</strong>**<br />
0<br />
4 (1.1)<br />
292 (80.7)<br />
66 (18.2)<br />
362 (100)<br />
Families which <strong>in</strong>clude <strong>alien</strong> spp.<br />
0<br />
2 (2.5)<br />
64 (80.0)<br />
14 (17.5)<br />
80 (100)<br />
TOTAL no. <strong>species</strong> 84 (2.9) 14 (0.5) 2171 (75.7) 598 (20.9) 2867 (100)<br />
% <strong>alien</strong> <strong>species</strong> / total no. <strong>species</strong> 0 28.6 13.5 11.0 12.6<br />
* Exclud<strong>in</strong>g sub<strong>species</strong> and varieties<br />
** Exclud<strong>in</strong>g cultivated <strong>species</strong> which are not, or only rarely, found <strong>in</strong> the environment<br />
The geographic orig<strong>in</strong> of <strong>alien</strong> and naturalized <strong>species</strong><br />
About one-fifth of all <strong>alien</strong> plant <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> have been <strong>in</strong>troduced from<br />
each of the follow<strong>in</strong>g areas: North America, Asia and the Mediterranean (Fig. 10.1;<br />
Table 10.2). Some 15 % have been <strong>in</strong>troduced from the rest of Europe and Eurasia/the<br />
Caucasus. The distribution of the so-called Eurasian <strong>species</strong> usually extends from<br />
eastern and south-eastern Europe to Asia M<strong>in</strong>or. Approximately one-fifth of the <strong>species</strong><br />
of Asian orig<strong>in</strong> were restricted to western Asia and the other four-fifths to Central<br />
Asia, Ch<strong>in</strong>a and east Asia. Only three <strong>species</strong> orig<strong>in</strong>ate from Africa, three from Central<br />
America and 14 from South America (Table 10.6). One <strong>species</strong> is native to Australia.<br />
Seven per cent of all <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> are of unknown orig<strong>in</strong>.<br />
One hundred and two <strong>alien</strong> <strong>species</strong> have become naturalized. Although 71 <strong>alien</strong> plant<br />
<strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> have been <strong>in</strong>troduced from North America and 76 from the<br />
Mediterranean region, only 18.4 % from the Mediterranean region have become naturalized<br />
whereas 43.7 % of those of North American orig<strong>in</strong> have done so. Also, 31.1 %<br />
of the <strong>alien</strong> <strong>species</strong> of European orig<strong>in</strong> have become naturalized but only 25.8 % of<br />
those of Asian/Eurasian orig<strong>in</strong>. It should be noted that three out of five <strong>species</strong> <strong>in</strong>troduced<br />
from the Caucasus have become naturalized and two of them are considered to<br />
be <strong>in</strong>vasive (Heracleum mantegazzianum and Rubus armeniacus Focke).
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 132<br />
Tab. 10.2 > The orig<strong>in</strong> of the <strong>alien</strong> flora of <strong>Switzerland</strong>: number of <strong>species</strong> (%).<br />
North America<br />
South America Asia Eurasia/Caucasus Europe Mediterranean. Others Unknown Total<br />
Alien 71 (19.6) 14 (3.9) 68 (18.8) 56 (15.5) 45 (12.4) 76 (21.0) 7 (1.9) 25 (6.9) 362 (100)<br />
Naturalized 31 (30.4) 3 (2.9) 17 (16.7) 15 (14.7) 14 (13.7) 14 (13.7) 3 (2.9) 5 (4.9) 102 (100)<br />
<strong>Invasive</strong> 8 (40.0) 1 (5.0) 8 (40.0) 2 (10.0) 0 0 1 (5.0) 0 20 (100)<br />
Fig. 10.1 > The orig<strong>in</strong> of <strong>alien</strong> plants <strong>in</strong> <strong>Switzerland</strong>.<br />
Mediterranean<br />
21%<br />
Europe<br />
12%<br />
Eurasia/<br />
Caucasus<br />
16%<br />
Asia<br />
19%<br />
Others/<br />
unknown<br />
13%<br />
North America<br />
20%<br />
Fig. 10.2 > The orig<strong>in</strong> of naturalized <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>.<br />
Mediterranean<br />
14%<br />
Europe<br />
14%<br />
Eurasia/<br />
Caucasus<br />
15%<br />
Others/<br />
unknown<br />
11%<br />
In total, 30.4 % of the naturalized flora orig<strong>in</strong>ates from North America and 31.4 % from<br />
Eurasia and the Caucasus (Fig. 10.2; Table 10.2).<br />
Of the 20 <strong>species</strong> that are considered to be <strong>in</strong>vasive <strong>in</strong> <strong>Switzerland</strong> (http://www.cpsskew.ch/),<br />
40 % orig<strong>in</strong>ate each from North America and Asia (Table 10.2). With the<br />
exception of the Caucasian <strong>species</strong>, no Eurasian or Mediterranean <strong>species</strong> have become<br />
<strong>in</strong>vasive. One might argue that Eurasian and Mediterranean <strong>species</strong> capable of be<strong>in</strong>g<br />
<strong>in</strong>vasive <strong>in</strong> <strong>Switzerland</strong> will have arrived already. Thus, not surpris<strong>in</strong>gly, the majority<br />
of naturalized and <strong>in</strong>vasive <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> orig<strong>in</strong>ate from temperate North<br />
America and Asia. Europe and the Mediterranean are a major source of <strong>alien</strong> <strong>species</strong>,<br />
but <strong>in</strong>terest<strong>in</strong>gly not of naturalized or <strong>in</strong>vasive plants.<br />
Pathways of <strong>in</strong>troduction<br />
From the 20 <strong>species</strong> on the Black List of <strong>in</strong>vasive <strong>species</strong> (http://www.cps-skew.ch/),<br />
15 (75 %) have been deliberately <strong>in</strong>troduced, usually as ornamentals. The pathway of<br />
<strong>in</strong>troduction of the two aquatic <strong>species</strong>, Elodea canadensis Michaux and E. nuttallii<br />
(Planchon), is unknown but they might well have escaped from garden ponds or been<br />
released from aquariums and thus are likely to be deliberate <strong>in</strong>troductions as well.<br />
Kowarik (2003) separates the pathways of the 25 problematic <strong>species</strong> <strong>in</strong> Germany <strong>in</strong>to<br />
21 deliberate (84 %) and four accidental (16 %) <strong>in</strong>troductions. These figures are summarized<br />
<strong>in</strong> Table 10.3 together with the correspond<strong>in</strong>g numbers of the total neophytes<br />
<strong>in</strong> Austria and the Czech Republic.<br />
Asia<br />
17%<br />
North America<br />
30%
10 > Plants – Planta 133<br />
10.4<br />
These two sets of data each give a similar picture for the Central European flora. The<br />
total numbers of neophytes <strong>in</strong> Austria and the Czech Republic are around 1,000 <strong>species</strong><br />
and the numbers of problematic <strong>species</strong> <strong>in</strong> Germany and <strong>Switzerland</strong> are 25 and 27,<br />
respectively. It should be noted that the figure of 1,000 neophytes given by Essl and<br />
Rabitsch (2002) for Austria is much higher than the 300 <strong>in</strong>troduced <strong>species</strong> given by<br />
Heywood (1989) for the same country.<br />
The percentage of deliberate <strong>in</strong>troductions that have become problematic is between<br />
75 % and 84 %, while it is lower for all neophytes, i.e. between 55 % and 59 %. The<br />
comparison <strong>in</strong>dicates that <strong>species</strong> selected for <strong>in</strong>troduction are more likely to become<br />
problematic <strong>species</strong> than those arriv<strong>in</strong>g accidentally. These differences may be attributed<br />
to human dimensions <strong>in</strong> the success of <strong>in</strong>vasions, <strong>in</strong> particular propagule pressure.<br />
Tab. 10.3 > Pathways of <strong>in</strong>troductions <strong>in</strong>to four European countries.<br />
The numbers for <strong>Switzerland</strong> and Germany are based on problematic plants (authors’ data and<br />
Kowarik, 2003), while the numbers for Austria (Essl and Rabitsch, 2002) and Czech Republic<br />
(Pysek et al., 2002) are neophytes.<br />
<strong>Switzerland</strong><br />
Germany Austria Czech Republic<br />
Total number 20 25 1110 924<br />
Deliberate 15 21 652 504<br />
Accidental 4 4 372 420<br />
Unknown 1 - 86 -<br />
Deliberate/Accidental as percentage of total 75 / 25 84 / 16 59 / 33 55 / 45<br />
Status of the <strong>alien</strong> <strong>species</strong> of <strong>Switzerland</strong><br />
Of the 362 <strong>alien</strong> <strong>species</strong> recorded <strong>in</strong> <strong>Switzerland</strong>, 102 <strong>species</strong> (28.2 %) have become<br />
naturalized and 20 <strong>species</strong> have become <strong>in</strong>vasive (5.5 %) (Fig. 10.3), which also means<br />
that a quarter of all naturalized <strong>species</strong> have become <strong>in</strong>vasive. The rema<strong>in</strong><strong>in</strong>g <strong>species</strong><br />
are either adventive or subspontaneous, or of unknown status. Thus, 3.6 % of the Swiss<br />
flora consists of naturalized <strong>species</strong>.<br />
The 362 <strong>alien</strong> <strong>species</strong> belong to 80 plant families, i.e. about half of all plant families<br />
present <strong>in</strong> <strong>Switzerland</strong> (Table 10.1). The percentage of <strong>alien</strong> <strong>species</strong> per family ranges<br />
from 2.8 % (Juncaceae) to 100 % for 20 plant families which are represented by <strong>alien</strong><br />
<strong>species</strong> only, usually by one or two <strong>species</strong> (Table 10.7). The percentage of <strong>alien</strong><br />
<strong>species</strong> <strong>in</strong> the large families (>30 <strong>species</strong>) ranges from 2.8 (Juncaceae) to 25.6 %<br />
(Polygonaceae). The largest family, Asteraceae, comprises 340 <strong>species</strong> of which some<br />
12.4 % are <strong>alien</strong> <strong>species</strong>. Thirty percent of all <strong>alien</strong> <strong>species</strong> belong to the Asteraceae,<br />
Brassicaceae and Poaceae.<br />
Some 50 plant families <strong>in</strong> <strong>Switzerland</strong> <strong>in</strong>clude no <strong>alien</strong> <strong>species</strong>. Most of these are small<br />
or very small families. Exceptions are the Orchidaceae (62 <strong>species</strong>), Gentianaceae (34<br />
<strong>species</strong>), Potamogetonaceae (21 <strong>species</strong>) and Orobanchaceae (20 <strong>species</strong>).
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 134<br />
Fig. 10.2 > Status of <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>.<br />
Unknown<br />
22%<br />
<strong>Invasive</strong><br />
5.5%<br />
Naturalized<br />
28%<br />
Adventive<br />
22%<br />
Subspont.<br />
28%<br />
10.5 Naturalized <strong>species</strong> of <strong>Switzerland</strong><br />
Naturalized <strong>species</strong> are represented by 102 <strong>species</strong> belong<strong>in</strong>g to 49 plant families, i.e.<br />
less than one-third of the total number of families <strong>in</strong> <strong>Switzerland</strong> (Table 10.7). The<br />
majority (30 families) <strong>in</strong>clude only one naturalized <strong>species</strong>. Eleven families <strong>in</strong>clude<br />
two naturalized <strong>species</strong>, and only eight families have more than two naturalized <strong>species</strong>.<br />
The Asteraceae alone <strong>in</strong>cludes ten naturalized <strong>species</strong> followed by the Rosaceae<br />
(n<strong>in</strong>e), the Brassicaceae (seven) and the Fabaceae (six). Almost one-third of all naturalized<br />
<strong>species</strong> belong to one of these four families.<br />
The percentage of naturalized <strong>species</strong> per family ranges from 0.8 % (Caryophyllaceae)<br />
to 100 % for eight families that are represented exclusively by their naturalized <strong>species</strong><br />
(usually one <strong>species</strong> – with the exception of the Phytolaccaceae which is represented<br />
by two <strong>species</strong>) (Table 10.7). The percentage of naturalized <strong>species</strong> <strong>in</strong> the large families<br />
(>30 <strong>species</strong>) ranges from 0.8 % (Caryophyllaceae) to 7.7 % (Polygonaceae). The<br />
largest family, Asteraceae, comprises 2.9 % naturalized <strong>species</strong>. The aquatic plant<br />
family Hydrocharitaceae comprises 71.4 % naturalized <strong>species</strong>.<br />
Fig. 10.3 > Number of established <strong>species</strong> as a function of the number of <strong>alien</strong> <strong>species</strong> (r 2 =0.8).<br />
No. of naturalized <strong>species</strong> /<br />
family<br />
12<br />
10<br />
8<br />
6<br />
4<br />
2<br />
0<br />
0 5 10 15 20 25 30 35 40<br />
No. of exotic <strong>species</strong> / family
10 > Plants – Planta 135<br />
In total, 28.2 % of all <strong>alien</strong> <strong>species</strong> have become naturalized. No naturalization at all is<br />
observed <strong>in</strong> 31 families (compris<strong>in</strong>g 55 <strong>alien</strong> <strong>species</strong>). The highest rate of failure is <strong>in</strong><br />
the Amaranthaceae with n<strong>in</strong>e <strong>alien</strong> but no naturalized <strong>species</strong>. In contrast, 100 %<br />
naturalization is observed <strong>in</strong> 17 families account<strong>in</strong>g for 20 <strong>alien</strong> <strong>species</strong>. The number<br />
of naturalized <strong>species</strong> is correlated to the number of <strong>species</strong> <strong>in</strong>troduced (Fig. 10.4).<br />
10.6 Life form<br />
The 11 life forms that are def<strong>in</strong>ed <strong>in</strong> Flora Helvetica (see also Table 10.6) have been<br />
pooled <strong>in</strong>to eight different ones. The categories ‘trees’ and ‘shrubs’ <strong>in</strong>clude both<br />
deciduous and evergreen plants. The category ‘small shrubs’ <strong>in</strong>cludes both woody and<br />
herbaceous chamaephytic plants. Chamaephytic plants are perennial <strong>species</strong> with<br />
persistent stems and buds overw<strong>in</strong>ter<strong>in</strong>g above ground level. Geophytic plants are<br />
perennial herbs with tubers, bulbs or rhizomes.<br />
In all, 46.4 % of the <strong>alien</strong> flora of <strong>Switzerland</strong> consists of annual and biennial <strong>species</strong><br />
(n=168) but only 21.6 % of the naturalized flora is <strong>in</strong> these groups (n=22) (Table 10.4).<br />
In contrast, perennial <strong>species</strong> form 53.6 % of the <strong>alien</strong> flora but 78.4 % of the naturalized<br />
flora. Thus, of the <strong>alien</strong> <strong>species</strong> present <strong>in</strong> <strong>Switzerland</strong>, relatively more aquatic<br />
plants (78 %), small shrubs (55 %) and trees (47 %), and <strong>in</strong>deed perennial plants as a<br />
whole (41 %) have become naturalised, compared to biennial plants (13 %) and annual<br />
herbs (13 %).<br />
Tab. 10.4 > The life form of vascular plants <strong>in</strong> <strong>Switzerland</strong>: number of <strong>species</strong> (% of total).<br />
Life forms Alien <strong>species</strong> Naturalized <strong>species</strong> <strong>Invasive</strong> <strong>species</strong> Native <strong>species</strong><br />
Perennials Trees 30 (8.3) 14 (13.7) 5 (25.0) 68 (2.7)<br />
Shrubs 25 (6.9) 8 (7.8) 2 (10.0) 103 (4.1)<br />
Small shrubs 22 (6.1) 12 (11.8) 0 238 (9.6)<br />
Geophytic plants 48 (13.3) 18 (17.6) 7 (35.0) 383 (15.4)<br />
Herbs 60 (16.6) 21 (20.6) 1 (5.0) 1120 (44.9)<br />
Aquatic plants 9 (2.5) 7 (6.9) 2 (10.0) 62 (2.5)<br />
Perennials (total) 194 (53.6) 80 (78.4) 17 (85) 1974 (79.1)<br />
Biennial herbs 54 (14.9) 7 (6.9) 1 (5.0) 239 (9.6)<br />
Annual herbs 114 (31.5) 15 (14.7) 2 (10.0) 279 (11.2)<br />
Total 362 102 20 2492<br />
Woody and geophytic plants account for 70 % of <strong>in</strong>vasive <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong><br />
(Table 10.4). Perhaps surpris<strong>in</strong>gly, no chamaephytic and only one herbaceous perennial<br />
have become <strong>in</strong>vasive. The proportions of shrubs and aquatic plants <strong>in</strong> the <strong>in</strong>vasive<br />
flora are similar, as are the proportions of these two life forms <strong>in</strong> the naturalized flora,<br />
i.e. 7–10 %. Thus, there are aga<strong>in</strong> a disproportionate number of large woody perennials
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 136<br />
and geophytic <strong>species</strong> that have become <strong>in</strong>vasive compared to those that have become<br />
naturalized. Only few short-lived <strong>species</strong> have become <strong>in</strong>vasive.<br />
In total, 85 % of the <strong>in</strong>vasive flora <strong>in</strong> <strong>Switzerland</strong> consists of perennial <strong>species</strong> of which<br />
more than two-thirds are trees and geophytic plants. The proportion of <strong>alien</strong> <strong>species</strong><br />
that are trees is three times the proportion of native <strong>species</strong> that are trees (Table 10.4),<br />
but the proportion of <strong>in</strong>vasive <strong>species</strong> that are trees is about ten times the figure for<br />
native <strong>species</strong>. For comparison, the proportion of short-lived plants that are <strong>alien</strong> is<br />
about twice the proportion for native <strong>species</strong>, but the proportions of short-lived <strong>species</strong><br />
that are <strong>in</strong>vasive and native are similar. In contrast, the proportion of herbaceous<br />
perennials that are native is almost three times higher than the figure for <strong>alien</strong> <strong>species</strong><br />
and nearly ten times higher than for <strong>in</strong>vasive <strong>species</strong> (Table 10.4). In summary, the life<br />
form composition of the <strong>alien</strong> flora is different from the native one and it changes<br />
dur<strong>in</strong>g the process of naturalization and <strong>in</strong>vasion. While the highest number of <strong>in</strong>troduced<br />
<strong>species</strong> are annual plants, they do not naturalize well. The relatively small<br />
number of <strong>in</strong>troduced trees and geophytic plants are successful <strong>in</strong> the naturalization<br />
process and are aggressive <strong>in</strong>vaders. The <strong>in</strong>troduction, naturalization and <strong>in</strong>vasion of<br />
<strong>alien</strong> plant <strong>species</strong> results not only <strong>in</strong> a change <strong>in</strong> the floristic composition, but also <strong>in</strong><br />
plant life form changes with probable consequences on habitat structure and ecosystem<br />
function<strong>in</strong>g.<br />
10.7 The habitats of <strong>alien</strong> plants <strong>in</strong> <strong>Switzerland</strong><br />
The ecological plant groups used here have been taken from Lauber and Wagner<br />
(1998) to def<strong>in</strong>e plant habitats. N<strong>in</strong>e ecological groups have been def<strong>in</strong>ed:<br />
> F = forest plant<br />
> M = mounta<strong>in</strong> plant<br />
> P = (lowland) pioneer plant<br />
> E = aquatic plant<br />
> H = marsh plant<br />
> S = dry grassland plant<br />
> G = grassland plant<br />
> R = ruderal plant<br />
Tab. 10.5 > The ecological groups of <strong>alien</strong> plants <strong>in</strong> <strong>Switzerland</strong>, exclud<strong>in</strong>g cultivated plants:<br />
number of <strong>species</strong> (%).<br />
Forest<br />
Mounta<strong>in</strong> Pioneer Aquatic Marsh Dry grassland Grassland Ruderal Total<br />
Alien 34 (11.3) 4 (1.3) 19 (6.3) 12 (4.1) 26 (8.6) 14 (4.7) 4 (1.3) 189 (62.6) 302<br />
Naturalized 23 (23.5) 3 (3.1) 11 (11.2) 8 (8.2) 11 (11.2) 4 (4.1) 0 38 (38.8) 98<br />
<strong>Invasive</strong> 3 (15.0) 0 7 (35.0) 2 (10.0) 3 (15.5) 0 0 5 (25.0) 20<br />
Native 443 (17.9) 644 (26.0) 126 (5.1) 96 (3.9) 308 (12.4) 347 (14.0) 74 (3.0) 438 (17.7) 2476
10 > Plants – Planta 137<br />
The majority of <strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> are ruderal plants (62.6 %), followed by<br />
forest plants (11.3 %) and marsh plants (8.6 %) (Table 10.5). However, ruderal plants<br />
represent only 38.8 % of all naturalized <strong>species</strong>. Forest <strong>species</strong> <strong>in</strong>crease to 23.5 % of all<br />
naturalized <strong>species</strong>, followed by pioneer and marsh <strong>species</strong> (11.2 %), and aquatic<br />
<strong>species</strong> (8.2 %). There are very few mounta<strong>in</strong> and grassland <strong>species</strong> naturalized <strong>in</strong><br />
<strong>Switzerland</strong>. Thus, it appears that forest and wetland habitats are more suited to plant<br />
naturalization s<strong>in</strong>ce 42.9 % of all naturalized <strong>species</strong> belong to these ecological groups<br />
(F, E, H). It should be noted, however, that three out of four mounta<strong>in</strong> <strong>species</strong> have<br />
become naturalized. Only 20.2 % of ruderal <strong>species</strong> have become naturalized.<br />
Ruderal and pioneer <strong>species</strong> represent 60 % of all <strong>in</strong>vasive <strong>species</strong>, <strong>in</strong>dicat<strong>in</strong>g perhaps<br />
less resistance of these habitats to <strong>in</strong>vasion. Wetlands and forests seem to be more<br />
resistant to <strong>in</strong>vasion. However, 40 % of <strong>in</strong>vasive <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong> belong to either<br />
the forest, aquatic or marsh ecological plant groups (F, E, H) and thus are a potential<br />
threat to the most valuable ecosystems of <strong>Switzerland</strong>, i.e. forests and wetland habitats.<br />
Mounta<strong>in</strong> and grassland habitats seem to be at less risk based on <strong>in</strong>vasions to date.<br />
About 50 % of all naturalized and <strong>in</strong>vasive <strong>species</strong> are ruderal and lowland pioneer<br />
<strong>species</strong>. These <strong>species</strong> are often not restricted to waste ground or urban areas, but are<br />
known to <strong>in</strong>vade (sometimes after a long period) semi-natural habitats such as meadows,<br />
riverbanks, gravel shores, forest marg<strong>in</strong>s or clear<strong>in</strong>gs. Several of the worst <strong>in</strong>vasive<br />
plants <strong>in</strong> <strong>Switzerland</strong> such as Reynoutria japonica, Heracleum manteggazianum,<br />
Impatiens glandulifera Royle, Buddleja davidii Franchet and Solidago canadensis L.<br />
are treated as ruderal or pioneer plants but are known to <strong>in</strong>vade less disturbed places as<br />
well. Solidago spp. <strong>in</strong> particular have been recognized for over a decade as a threat to<br />
protected areas (Voser-Huber, 1992). Impatiens parviflora DC. was <strong>in</strong>troduced from<br />
Asia <strong>in</strong>to Central Europe <strong>in</strong> 1837 and was for many years a typical ruderal plant occurr<strong>in</strong>g<br />
only <strong>in</strong> towns, gardens, parks or cemeteries. A few decades ago, however, it<br />
started to penetrate <strong>in</strong>to woods, at first only <strong>in</strong> badly degraded areas, but then becom<strong>in</strong>g<br />
firmly established <strong>in</strong> relatively natural stands of deciduous forests <strong>in</strong> Central Europe<br />
(Kornas, 1990).<br />
Weber (1999a) gives the number of <strong>alien</strong> plant <strong>species</strong> recorded <strong>in</strong> various habitats<br />
(s.str.) <strong>in</strong> <strong>Switzerland</strong>: 16 % are known from forests and related habitats, 15 % from<br />
lakes and wetland areas, and 8 % from grassland habitats. Interest<strong>in</strong>gly, 16 % of <strong>alien</strong><br />
plants are found <strong>in</strong> rocky sites and on walls. As compared to the 61 % of ruderal <strong>species</strong>,<br />
some 35 % of <strong>alien</strong> plants grow only <strong>in</strong> ruderal sites. The number of <strong>alien</strong> plants<br />
recorded <strong>in</strong> various habitats is relatively similar to the numbers that can be extrapolated<br />
from the ecological groups. The data suggest that many ruderal <strong>alien</strong> plants are not<br />
restricted to ruderal sites but grow ma<strong>in</strong>ly <strong>in</strong> open habitats or less disturbed sites such<br />
as rocky areas, walls, or forests subjected to some disturbance.<br />
As shown above, the great majority of naturalized and <strong>in</strong>vasive <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong><br />
are perennial <strong>species</strong> and most of them are forest, wetland and ruderal <strong>species</strong>. With<br />
very few exceptions (e.g. Impatiens glandulifera), naturalized and <strong>in</strong>vasive short-lived<br />
<strong>species</strong> are uncommon <strong>in</strong> semi-natural or natural habitats.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 138<br />
With respect to plant life form, the composition of the ecological plant groups of the<br />
<strong>alien</strong> flora differs from the native one, and it changes dur<strong>in</strong>g the process of naturalization<br />
and <strong>in</strong>vasion. The proportion of ruderal <strong>species</strong> <strong>in</strong> the <strong>alien</strong> flora is much higher<br />
than <strong>in</strong> the native one but this discrepancy is reduced dur<strong>in</strong>g the naturalization process.<br />
In contrast, the proportion of <strong>alien</strong> forest plant <strong>species</strong> is lower than that of native<br />
forest plant <strong>species</strong> but proportions of <strong>in</strong>vasive and native forest plant <strong>species</strong> are fairly<br />
similar.<br />
10.8 <strong>Invasive</strong> plant <strong>species</strong> <strong>in</strong> Europe<br />
Up to 2003, only a few European countries had compiled data on their <strong>alien</strong> and <strong>in</strong>vasive<br />
floras at a national level. However, <strong>in</strong>formation from several countries is quite<br />
<strong>in</strong>formative about <strong>in</strong>vasive <strong>species</strong> elsewhere <strong>in</strong> Europe that could be a threat to <strong>Switzerland</strong>.<br />
Countries usually make explicit dist<strong>in</strong>ctions between <strong>in</strong>vasive <strong>species</strong>, moderately<br />
or potentially <strong>in</strong>vasive <strong>species</strong>, and <strong>species</strong> for which special attention is needed<br />
(usually the so-called ‘Watch List’). We had to <strong>in</strong>terpret some data to a certa<strong>in</strong> extent<br />
to obta<strong>in</strong> a more consistent result. Austria <strong>in</strong>cludes an assessment of economic impact,<br />
which we have reproduced <strong>in</strong> Table 10.8 (Essl and Rabitsch, 2002). Also, the European<br />
and Mediterranean Plant Protection Organization (EPPO) is prepar<strong>in</strong>g a list of <strong>in</strong>vasive<br />
and potentially <strong>in</strong>vasive <strong>alien</strong> plants for the EPPO region. This <strong>in</strong>formation is still<br />
partial and it has not been used for the present review.<br />
Plant <strong>species</strong> declared <strong>in</strong>vasive or potentially <strong>in</strong>vasive <strong>in</strong> n<strong>in</strong>e European countries and<br />
occurr<strong>in</strong>g <strong>in</strong> <strong>Switzerland</strong> are presented <strong>in</strong> Table 10.8. The list <strong>in</strong>cludes over 130 <strong>alien</strong><br />
plant <strong>species</strong> of concern <strong>in</strong> Europe. Quite obviously, not all of them are sufficiently<br />
pre-adapted to the eco-climatic conditions of <strong>Switzerland</strong> to present a threat <strong>in</strong> this part<br />
of Europe. On the other hand, some <strong>in</strong>vasive <strong>species</strong> <strong>in</strong> northern European countries,<br />
for example, which are not listed <strong>in</strong> Table 10.8, could become a problem <strong>in</strong> <strong>Switzerland</strong>.<br />
Unfortunately, no detailed list of <strong>alien</strong> plants <strong>in</strong> Germany for each of the three<br />
categories def<strong>in</strong>ed was available at the time of this review. In contrast, the detailed<br />
<strong>in</strong>formation on the status of <strong>alien</strong> plants <strong>in</strong> France is highly relevant to <strong>Switzerland</strong><br />
(Muller, 2004).<br />
The list of <strong>in</strong>vasive <strong>species</strong> (‘Black List’) and ‘Watch Species’ <strong>in</strong> <strong>Switzerland</strong> is taken<br />
from the CPS-SKEW work<strong>in</strong>g group (http://www.cps-skew.ch/<strong>in</strong>dex.htm). A few<br />
<strong>species</strong> <strong>in</strong> these two categories do not have this status <strong>in</strong> the other European countries<br />
from which <strong>in</strong>formation could be collected (Table 10.8). However, they are <strong>in</strong>vasive on<br />
other cont<strong>in</strong>ents, e.g. Pueraria lobata (Willd.), and Lonicera japonica Thunberg<br />
(Cronk and Fuller, 1995; Weber, 2003) or their taxonomy is complex and their <strong>in</strong>vasive<br />
status may not yet have been recognized (e.g. Rubus armeniacus).<br />
Fact Sheets have been prepared for 48 plant <strong>species</strong>. They <strong>in</strong>clude 19 <strong>in</strong>vasive <strong>species</strong><br />
and 11 ‘Watch Species’ of the CPS-SKEW work<strong>in</strong>g group. In this report, another 18<br />
<strong>species</strong> to which special attention should be given <strong>in</strong> the future are also presented <strong>in</strong><br />
Fact Sheets. These <strong>species</strong> have been selected accord<strong>in</strong>g to their status <strong>in</strong> neighbour<strong>in</strong>g<br />
countries and a previous list of potentially <strong>in</strong>vasive <strong>species</strong> prepared by the CPS-<br />
SKEW work<strong>in</strong>g group (Table 10.8).
10 > Plants – Planta 139<br />
10.9 Discussion<br />
About 362 <strong>alien</strong> <strong>species</strong> are established <strong>in</strong> <strong>Switzerland</strong> and almost one third of these<br />
have become naturalized with certa<strong>in</strong>ty. With the exception of aquatic plants <strong>in</strong> the<br />
Hydrocharitaceae, no one plant family appears to be particularly successful <strong>in</strong> naturalization.<br />
This review stresses the importance of establish<strong>in</strong>g the biological and ecological<br />
characteristics of the naturalized flora for determ<strong>in</strong><strong>in</strong>g the potential <strong>in</strong>vasiveness of<br />
<strong>alien</strong> <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>. The life form composition of the <strong>alien</strong> flora is different<br />
from that of the native one and changes dur<strong>in</strong>g the process of naturalization from<br />
dom<strong>in</strong>ance by short-lived to perennial <strong>species</strong>. The flora of naturalized plants consists<br />
of almost 80 % perennial <strong>species</strong>. There is a further shift dur<strong>in</strong>g the process of <strong>in</strong>vasion<br />
towards large woody perennial and geophytic plants. The <strong>in</strong>vasion by herbaceous<br />
perennials and short-lived plants has been negligible so far. <strong>Invasive</strong> <strong>species</strong> come<br />
from geographically distant areas and no European or Mediterranean <strong>species</strong> is considered<br />
to be <strong>in</strong>vasive or potentially <strong>in</strong>vasive <strong>in</strong> <strong>Switzerland</strong>.<br />
More than 40 % of all naturalized and <strong>in</strong>vasive <strong>species</strong> consist of either forest or wetland<br />
<strong>species</strong>, thus it appears that these habitats are at a higher risk of <strong>in</strong>vasion. In<br />
Europe, over 50 % of the naturalized flora occurs <strong>in</strong> river border communities (Sykora,<br />
1990). This is because rivers are an effective means of transportation for many <strong>species</strong>,<br />
natural riverbank communities have been largely destroyed by human activities, and<br />
riverbanks are regularly disturbed by water movement. Alluvial zones and mires are<br />
therefore of primary concern with regard to <strong>in</strong>vasive <strong>species</strong>. The typical vegetation of<br />
lowland alluvial zones such as river<strong>in</strong>e floodpla<strong>in</strong>s consists of a mosaic of pioneer<br />
communities, shrubs and alluvial forests. Invasion by pioneer or ruderal <strong>alien</strong> <strong>species</strong> is<br />
likely to occur and expand <strong>in</strong> such sites due to human activities or colonization from<br />
upstream or adjacent fields. Fenlands also are at risk because they are often used for<br />
agricultural purposes. In contrast, raised bogs are at a lower risk s<strong>in</strong>ce only highly<br />
specialized plant <strong>species</strong> can flourish <strong>in</strong> such habitats, and there is less permanent<br />
natural disturbance.<br />
To date, the <strong>in</strong>vasibility of mounta<strong>in</strong> and grassland habitats has been low and few <strong>alien</strong><br />
<strong>species</strong> have become naturalized <strong>in</strong> these habitats. Because meadows and grasslands<br />
owe their existence almost exclusively to human management, the threat from <strong>alien</strong><br />
<strong>species</strong> will <strong>in</strong>crease with land-use changes and reduced grassland ma<strong>in</strong>tenance. In the<br />
absence of any management, most meadows and grasslands below the timberl<strong>in</strong>e<br />
degenerate to scrub and revert to woodland. Alien shrubs and trees may profit from<br />
such a situation.<br />
The restoration of biological diversity <strong>in</strong> <strong>in</strong>tensively used agricultural land has been<br />
supported for several years through various agri-environmental schemes. In many areas<br />
this <strong>in</strong>volves the conversion of <strong>in</strong>tensively managed arable land to extensive pastures<br />
or so-called areas of ecological compensation. The transition period from <strong>in</strong>tensively to<br />
extensively managed land or semi-natural habitats is highly favourable to <strong>alien</strong> <strong>species</strong>.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 140<br />
Alien plants are a threat to the areas of ecological compensation, the restoration of<br />
riverbanks and mires, abandoned grassland and forests, and all previously or currently<br />
disturbed natural areas <strong>in</strong> <strong>Switzerland</strong>. For example, the rehabilitation of riverbanks<br />
can be seriously handicapped by <strong>alien</strong> <strong>species</strong>. Many <strong>in</strong>vasive <strong>species</strong> <strong>in</strong>crease erosion<br />
(e.g. Reynoutria japonica, Buddleja davidii), which <strong>in</strong> turn accelerates the establishment<br />
of <strong>in</strong>vasive <strong>species</strong>, thus prevent<strong>in</strong>g the restoration of native plant communities.<br />
The problems caused by <strong>alien</strong> <strong>species</strong> <strong>in</strong> cultivated land are still relatively m<strong>in</strong>or but<br />
may <strong>in</strong>crease <strong>in</strong> the future because of newly naturalized <strong>species</strong> and because changes <strong>in</strong><br />
land use may favour establishment of <strong>alien</strong> <strong>species</strong> <strong>in</strong> extensive agro-ecosystems.<br />
Weber (1999a) recorded 38 <strong>alien</strong> plants that are known to occur <strong>in</strong> arable land. For<br />
example, Conyza canadensis (L.) and Epilobium ciliatum Raf<strong>in</strong>esque are of <strong>in</strong>creas<strong>in</strong>g<br />
importance <strong>in</strong> arable land, gardens, orchards and tree nurseries. Gal<strong>in</strong>soga parviflora<br />
Cavanilles is known to be a serious problem <strong>in</strong> vegetable crops <strong>in</strong> other countries and<br />
Cyperus esculentus is considered to be one of the world’s worst weeds. Ambrosia<br />
artemisiifolia is most abundant <strong>in</strong> sunflower crops ow<strong>in</strong>g to the botanical similarity<br />
between the weed and the crop itself, and therefore only a limited number of herbicides<br />
are available that are effective aga<strong>in</strong>st A. artemisiifolia alone. Senecio <strong>in</strong>aequidens is a<br />
potential problem <strong>in</strong> pastures and meadows. Unlike those <strong>in</strong> a natural environment,<br />
most <strong>alien</strong> plants <strong>in</strong> arable land are ruderal <strong>species</strong> and the majority of them are short<br />
lived.<br />
Alien <strong>species</strong> can also become a public health problem. Ambrosia artemisiifolia is a<br />
strongly allergenic plant. Exposure to the sap of Heracleum manteggazianum sensitizes<br />
the sk<strong>in</strong> to sunlight and causes severe irritation and pa<strong>in</strong>ful blisters. Contact with the<br />
sap of Ailanthus altissima (Miller) may also cause sk<strong>in</strong> eruptions.<br />
Most of the 20 declared <strong>in</strong>vasive plants <strong>in</strong> <strong>Switzerland</strong> are also <strong>in</strong>vasive <strong>in</strong> neighbour<strong>in</strong>g<br />
countries. An analysis of the <strong>in</strong>vasive flora of several European countries shows<br />
that over 130 <strong>alien</strong> plants are of concern <strong>in</strong> Europe. The status of <strong>alien</strong> plants <strong>in</strong> Europe<br />
should be one aspect considered <strong>in</strong> develop<strong>in</strong>g a dynamic ‘Watch List’ of <strong>alien</strong> plants<br />
<strong>in</strong> <strong>Switzerland</strong>. In addition to the known <strong>in</strong>vasive <strong>species</strong> of <strong>Switzerland</strong>, over 30 <strong>alien</strong><br />
plants should be monitored over time to predict their <strong>in</strong>vasive potential.<br />
In conclusion, a careful review of <strong>in</strong>vasive and potentially <strong>in</strong>vasive plants <strong>in</strong> Europe<br />
comb<strong>in</strong>ed with some key biological and ecological traits associated with naturalized<br />
<strong>species</strong> rather than <strong>alien</strong> plants, as well as field observations, should contribute to<br />
assess<strong>in</strong>g the <strong>in</strong>vasiveness of <strong>alien</strong> plant <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>. More research is<br />
needed to understand the naturalization and <strong>in</strong>vasion process and to evaluate the impact<br />
of <strong>in</strong>vasive <strong>species</strong> on the environment and the agro-ecosystem. More research is also<br />
needed to establish long-term and environmentally friendly management tools for<br />
<strong>in</strong>vasive plants <strong>in</strong> <strong>Switzerland</strong>.
10 > Plants – Planta 141<br />
References<br />
Cronk, Q.C.B. and J.L. Fuller (1995) Plant <strong>in</strong>vaders. Chapman & Hall, 241<br />
pp.<br />
Dana, E.D., Sanz-Elorza, M. and E. Sobr<strong>in</strong>o (2001) Plant <strong>in</strong>vaders <strong>in</strong><br />
Spa<strong>in</strong>. http://www.ual.es/personal/edana/<strong>alien</strong>plants/checklist.pdf<br />
De Almeida, J.D. (1999) Flora exotica subespontanea de Portugal<br />
cont<strong>in</strong>ental. Universidade de Coimbra, 151 pp.<br />
Essl, F. and W. Rabitsch (eds) (2002) Neobiota <strong>in</strong> Österreich. Federal<br />
Environment Agency, 432 pp.<br />
Heywood, V.H. (1989) Patterns, extends and modes of <strong>in</strong>vasions by<br />
terrestrial plants. In: Drake, J.A. et al. (eds) Biological <strong>in</strong>vasions: a global<br />
perspective. SCOPE 37, John Wiley & Sons, pp. 31–55.<br />
Kornas, J. (1990) Plant <strong>in</strong>vasions <strong>in</strong> Central Europe: historical and<br />
ecological aspects. In: di Castri, F., Hansen, A.J. and M. Debussche (eds)<br />
Biological <strong>in</strong>vasions <strong>in</strong> Europe and the Mediterranean Bas<strong>in</strong>. Kluwer<br />
Academic Publishers, pp. 19–36.<br />
Kowarik, I. (2003) Biologische Invasionen: Neophyten und Neozoen <strong>in</strong><br />
Mitteleuropa. Verlag Eugen Ulmer GmbH, Stuttgart, 380 pp.<br />
Lauber, K. and G. Wagner (1998) Flora Helvetica. Paul Haupt Publishers,<br />
1616 pp.<br />
Moser, D.M., Gygax, A., Bäumler. B., Wyler, N. and R. Palese (2002) Rote<br />
Liste der gefährdeten Farn- und Blütenpflanzen der Schweiz. BUWAL-<br />
Reihe ”Vollzug Umwelt“.<br />
Muller, S. (2004) Plantes <strong>in</strong>vasives en France. Paris, Muséum National<br />
d’Histoire Naturelle. Patrimo<strong>in</strong>es naturels, 62, 176 pp.<br />
Pyšek, P., Sádlo, J. and B. Mandák (2002) Catalogue of <strong>alien</strong> plants of the<br />
Czech Republic. Preslia, Praha, 97–186.<br />
Richardson, D.M., Pyšek, P., Rejmanek, M., Barbour, M.G., Panetta, F.D.<br />
and C.J. West (2000) Naturalization and <strong>in</strong>vasion of <strong>alien</strong> plants: concepts<br />
and def<strong>in</strong>itions. Diversity and Distributions 6, 93–107.<br />
Sykora, K.V. (1990) History of the impact of man on the distribution of<br />
plant <strong>species</strong>. In: di Castri, F., Hansen, A.J. and M. Debussche (eds)<br />
Biological <strong>in</strong>vasions <strong>in</strong> Europe and the Mediterranean Bas<strong>in</strong>. Kluwer<br />
Academic Publishers, pp. 37–50.<br />
Voser-Huber, M.-L. (1992) Verges d’or – problèmes dans les réserves<br />
naturelles. OFEFP, Cahier de l’environnement N° 167, 22 pp.<br />
Weber, E. (1999a) <strong>Switzerland</strong> and the <strong>in</strong>vasive plant issue. Bot Helv.<br />
110, 11–24.<br />
Weber, E (1999b) Gebietsfremde Arten der <strong>Schweizer</strong> Flora – Ausmass<br />
und Bedeutung. Bauh<strong>in</strong>ia 13, 1–10.<br />
Weber, E. (2003) <strong>Invasive</strong> plant <strong>species</strong> of the world. A reference guide to<br />
environmental weeds. CABI Publish<strong>in</strong>g, Wall<strong>in</strong>gford, 548 pp.<br />
Welch, D. et al. (2001) An audit of <strong>alien</strong> <strong>species</strong> <strong>in</strong> Scotland. Scottish<br />
Natural Heritage Review No 139, 225 pp.
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 142<br />
Tab. 8.6 > Alien <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>.<br />
Species<br />
Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Abutilon theophrasti Medic. Malvaceae Unknown t adventive N R<br />
Acalypha virg<strong>in</strong>ica L. Euphorbiaceae North America t naturalized N R<br />
Acer negundo L. Aceraceae North America t subspontaneous F<br />
Acorus calamus L. Araceae Unknown g unknown N E<br />
Aegilops cyl<strong>in</strong>drica Host Poaceae Mediterranean t adventive R<br />
Aegilops ovata L. Poaceae Mediterranean t adventive R<br />
Agave americana L. Agavaceae Central America j naturalized N R<br />
Ailanthus altissima (Miller) Simaroubaceae Asia p naturalized N P<br />
Aldrovanda vesiculosa L. Droseraceae Unknown a unknown N E<br />
Allium scorodoprasum L. Liliaceae Europa g unknown N R<br />
Alopecurus rendlei Eig Poaceae Mediterranean t adventive H<br />
Althaea offic<strong>in</strong>alis L. Malvaceae Asia h subspontaneous C<br />
Amaranthus albus L. Amaranthaceae North America t unknown N R<br />
Amaranthus blitum L. Amaranthaceae Unknown t unknown N R<br />
Amaranthus cruentus L. Amaranthaceae North America t subspontaneous N R<br />
Amaranthus deflexus L. Amaranthaceae South America u unknown N R<br />
Amaranthus graecizans L. Amaranthaceae Unknown t unknown N R<br />
Amaranthus hypochondriacus L. Amaranthaceae Unknown t unknown N R<br />
Amaranthus retroflexus L. Amaranthaceae Unknown t unknown N R<br />
Ambrosia artemisiifolia L. Asteraceae North America t adventive N R<br />
Amorpha fructicosa L. Fabaceae North America n subspontaneous N H<br />
Arabis rosea DC. Brassicaceae Europa h naturalized N P<br />
Arabis caucasica (Willdenow) Brassicaceae Eurasia c naturalized N M<br />
Aremonia agrimonioides (L.) Rosaceae Europa h naturalized N P<br />
Armoracia rusticana P.Gaertn., B. Mey. & Scherb. Brassicaceae Eurasia g subspontaneous C<br />
Artemisia annua L. Asteraceae Eurasia t adventive R<br />
Artemisia biennis Willdenow Asteraceae Eurasia u adventive R<br />
Artemisia verlotiorum Lamotte Asteraceae Asia g unknown N R<br />
Arum italicum Miller Araceae Mediterranean g naturalized R<br />
Asar<strong>in</strong>a procumbens Mill. Scrophulariaceae Mediterranean c naturalized N P<br />
Asclepias syriaca L. Asclepiadaceae North America g subspontaneous N CR<br />
Aster novae-angliae L. Asteraceae North America g subspontaneous N H<br />
Aster novi-belgii L. Asteraceae North America c subspontaneous N H<br />
Aster tradescantii L. Asteraceae North America g subspontaneous C<br />
Aubrieta deltoidea (L.) Brassicaceae Mediterranean c naturalized N S<br />
Avena barbata Pott Poaceae Mediterranean t adventive R<br />
Avena sativa L. Poaceae Eurasia t subspontaneous C<br />
Bidens bip<strong>in</strong>nata L. Asteraceae North America t naturalized N R<br />
Bidens connata Willdenow Asteraceae North America t naturalized N H<br />
Bidens frondosa L. Asteraceae North America t naturalized N R
10 > Plants – Planta 143<br />
Species<br />
Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Bidens subalternans DC. Asteraceae South America t adventive R<br />
Brassica juncea (L.) Brassicaceae Asia t adventive R<br />
Brassica nigra (L.) Brassicaceae Unknown t naturalized R<br />
Brassica rapa L. Brassicaceae Europa u subspontaneous C<br />
Bromus diandrus Roth Poaceae Europa t adventive R<br />
Bromus <strong>in</strong>ermis Leysser Poaceae Eurasia h unknown N R<br />
Bromus madritensis L. Poaceae Mediterranean t adventive R<br />
Bromus rigidus Roth Poaceae Mediterranean t adventive R<br />
Buddleja davidii Franchet Buddlejaceae Asia n naturalized N P<br />
Bunias orientalis L. Brassicaceae Eurasia h unknown N R<br />
Calla palustris L. Araceae Unknown g naturalized N E<br />
Cannabis sativa L. Cannabaceae Asia t subspontaneous C<br />
Carex vulp<strong>in</strong>oidea Michaux Cyperaceae North America h naturalized N H<br />
Centaurea diffusa Lamarck Asteraceae Mediterranean t adventive R<br />
Centranthus ruber (L.) Valerianaceae Mediterranean h naturalized N P<br />
Cerastium dubium (Bastard) Caryophyllaceae Mediterranean t adventive H<br />
Cerastium ligusticum Viviani Caryophyllaceae Mediterranean t adventive R<br />
Cerastium tomentosum L. Caryophyllaceae Europa c subspontaneous C<br />
Cer<strong>in</strong>the major L. Borag<strong>in</strong>aceae Eurasia t adventive R<br />
Cer<strong>in</strong>the m<strong>in</strong>or L. Borag<strong>in</strong>aceae Eurasia u adventive R<br />
Chenopodium ambrosioides L. Chenopodiaceae South America t adventive R<br />
Chenopodium pratericola Rydberg Chenopodiaceae North America t adventive R<br />
Chrysanthemum segetum L. Asteraceae Mediterranean t unknown N R<br />
Commel<strong>in</strong>a communis L. Commel<strong>in</strong>aceae Asia c naturalized N R<br />
Consolida ajacis (L.) Ranunculaceae Eurasia t subspontaneous C<br />
Conyza canadensis (L.) Asteraceae North America u unknown N P<br />
Cornus sericea L. Cornaceae North America n naturalized N F<br />
Coronopus didymus (L.) Brassicaceae South America u adventive N R<br />
Crepis nemauensis Gouan Asteraceae Mediterranean t naturalized N R<br />
Crepis nicaeensis Persoon Asteraceae Mediterranean u adventive R<br />
Crepis pulchra L. Asteraceae Mediterranean t unknown N R<br />
Cuscuta campestris Yuncker Cuscutaceae North America t adventive N R<br />
Cuscuta cesatiana Bertoloni Cuscutaceae Eurasia t unknown N R<br />
Cymbalaria muralis P. Gaertn., B. Mey. & Scherb. Scrophulariaceae Europa c unknown N P<br />
Cynodon dactylon (L.) Poaceae Mediterranean g naturalized N R<br />
Cyperus eragrostis Lamarck Cyperaceae South America h unknown H<br />
Cyperus esculentus L. Cyperaceae Unknown g unknown N H<br />
Cyperus rotundus L. Cyperaceae Eurasia g naturalized H<br />
Datura stramonium L. Solanaceae Central America t unknown N R<br />
Dianthus barbatus L. Caryophyllaceae Europa h subspontaneous C<br />
Diplotaxis erucoides (Torner) Brassicaceae Mediterranean u adventive R<br />
Dipsacus lac<strong>in</strong>iatus L. Dipsacaceae Mediterranean u unknown N R
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 144<br />
Species<br />
Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Duchesnea <strong>in</strong>dica (Andrews) Rosaceae Asia h naturalized N F<br />
Eleus<strong>in</strong>e <strong>in</strong>dica (L.) Poaceae Unknown t naturalized N R<br />
Elodea canadensis Michaux Hydrocharitaceae North America a unknown N E<br />
Elodea densa (Planchon) Hydrocharitaceae South America a naturalized N E<br />
Elodea nuttallii (Planchon) Hydrocharitaceae North America a naturalized N E<br />
Epilobium ciliatum Raf<strong>in</strong>esque Onagraceae North America h unknown N R<br />
Epimedium alp<strong>in</strong>um L. Berberidaceae Europa g naturalized N F<br />
Eragrostis cilianensis (Allioni) Poaceae Mediterranean t unknown N R<br />
Eragrostis multicaulis Steudel Poaceae Asia t adventive R<br />
Eranthis hyemalis (L.) Ranunculaceae Europa g subspontaneous N F<br />
Erica tetralix L. Ericaceae Europa z naturalized N H<br />
Erigeron annuus (L.) Asteraceae North America u naturalized N R<br />
Erigeron karv<strong>in</strong>skianus DC. Asteraceae Central America h unknown N P<br />
Erodium ciconium (L.) Geraniaceae Mediterranean t adventive R<br />
Erodium moschatum (L.) Geraniaceae Mediterranean u unknown N R<br />
Erysimum cheiri (L.) Brassicaceae Mediterranean c naturalized C<br />
Erysimum hieraciifolium L. Brassicaceae Eurasia u unknown N R<br />
Erysimum repandum L. Brassicaceae Europa t adventive R<br />
Euclidium syriacum (L.) Brassicaceae Eurasia t adventive R<br />
Euphorbia chamaesyce L. Euphorbiaceae Eurasia t adventive R<br />
Euphorbia humifusa Willdenow Euphorbiaceae Asia t unknown N R<br />
Euphorbia lathyris L. Euphorbiaceae Eurasia u subspontaneous N R<br />
Euphorbia maculata L. Euphorbiaceae North America t unknown N R<br />
Euphorbia nutans Lagasca Euphorbiaceae North America t unknown N R<br />
Euphorbia prostrata Aiton Euphorbiaceae North America t adventive R<br />
Euphorbia virgata Waldste<strong>in</strong> et Kitaibel Euphorbiaceae Eurasia h unknown N R<br />
Fagopyrum esculentum Moench Polygonaceae Asia t subspontaneous RC<br />
Fagopyrum tataricum (L.) Polygonaceae Asia t unknown R<br />
Ficus carica L. Moraceae Mediterranean p subspontaneous C<br />
Foeniculum vulgare Miller Apiaceae Mediterranean u naturalized N R<br />
Galega offic<strong>in</strong>alis L. Fabaceae Mediterranean h subspontaneous N G<br />
Gal<strong>in</strong>soga ciliata (Raf<strong>in</strong>esque) Asteraceae South America t unknown N R<br />
Gal<strong>in</strong>soga parviflora Cavanilles Asteraceae South America t unknown N R<br />
Galium saxatile L. Rubiaceae Europa c unknown N H<br />
Galium verrucosum Hudson Rubiaceae Mediterranean t adventive S<br />
Geranium sibiricum L. Geraniaceae Asia u unknown N F<br />
Glaucium corniculatum (L.) Papaveraceae Mediterranean t adventive P<br />
Glaucium flavum Crantz Papaveraceae Europa u adventive P<br />
Glyceria striata (Lamarck) Poaceae North America g unknown N H<br />
Gypsophila paniculata L. Caryophyllaceae Eurasia c naturalized N P<br />
Helianthus annuus L. Asteraceae North America t subspontaneous C<br />
Helianthus rigidus (Cass<strong>in</strong>i) Asteraceae North America g subspontaneous C
10 > Plants – Planta 145<br />
Species<br />
Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Helianthus tuberosus L. Asteraceae North America g subspontaneous N R<br />
Hemerocallis fulva (L.) Liliaceae Asia g naturalized N R<br />
Heracleum mantegazzianum Sommier & Levier Apiaceae Caucasus h naturalized N R<br />
Hibiscus trionum L. Malvaceae Asia t subspontaneous C<br />
Hordeum distichon L. Poaceae Western Asia t subspontaneous C<br />
Hordeum vulgare L. Poaceae Africa u subspontaneous C<br />
Hypericum calyc<strong>in</strong>um L. Hypericaceae Eurasia z subspontaneous C<br />
Iberis umbellata L. Brassicaceae Mediterranean u subspontaneous PC<br />
Impatiens balfourii Hooker F. Balsam<strong>in</strong>aceae Asia t unknown N R<br />
Impatiens glandulifera Royle Balsam<strong>in</strong>aceae Asia t naturalized N R<br />
Impatiens parviflora DC. Balsam<strong>in</strong>aceae Asia t naturalized N F<br />
Inula helenium L. Asteraceae Europa h subspontaneous C<br />
Iris foetidissima L. Iridaceae Europa g subspontaneous N S<br />
Iris germanica L. Iridaceae Mediterranean g naturalized C<br />
Iris lutescens Lamarck Iridaceae Eurasia g subspontaneous N S<br />
Iris sambuc<strong>in</strong>a L. Iridaceae Unknown g unknown N S<br />
Iris squalens L. Iridaceae Unknown g subspontaneous N S<br />
Juncus tenuis Willdenow Juncaceae North America h unknown N R<br />
Lagarosiphon major (Ridley) Hydrocharitaceae South Africa a naturalized N E<br />
Laurus nobilis L. Lauraceae Mediterranean i naturalized N F<br />
Legousia hybrida (L.) Campanulaceae Mediterranean t adventive R<br />
Lemna m<strong>in</strong>uta Humboldt et al. Lemnaceae North America a naturalized N E<br />
Lepidium densiflorum Schrader Brassicaceae North America u unknown N R<br />
Lepidium neglectum Thellung Brassicaceae North America u unknown N R<br />
Lepidium sativum L. Brassicaceae Western Asia t subspontaneous RC<br />
Lepidium virg<strong>in</strong>icum L. Brassicaceae North America u unknown N R<br />
Lepidium latifolium L. Brassicaceae Europa h subspontaneous RC<br />
Leucanthemum gaud<strong>in</strong>ii Della Torre Asteraceae Unknown h unknown N R<br />
Ligustrum lucidum Aiton Oleaceae Asia i subspontaneous C<br />
L<strong>in</strong>aria arvensis (L.) Scrophulariaceae Europa t adventive R<br />
L<strong>in</strong>aria repens (L.) Scrophulariaceae Europa h naturalized N R<br />
L<strong>in</strong>aria simplex (Willdenow) Scrophulariaceae Europa t adventive R<br />
L<strong>in</strong>aria dalmatica (L.) Scrophulariaceae Europa h naturalized N R<br />
L<strong>in</strong>um bienne Miller L<strong>in</strong>aceae Europa u adventive S<br />
L<strong>in</strong>um narbonense L. L<strong>in</strong>aceae Mediterranean h naturalized N S<br />
Lonicera japonica Thunberg Caprifoliaceae Asia i naturalized N F<br />
Lonicera henryi Hemsley Caprifoliaceae Asia i subspontaneous N F<br />
Ludwigia grandiflora (Michaux) Onagraceae South America a naturalized N E<br />
Lunaria annua L. Brassicaceae Europa u naturalized N R<br />
Lup<strong>in</strong>us polyphyllus L<strong>in</strong>dley Fabaceae North America h subspontaneous N F<br />
Mahonia aquifolium (Pursh) Berberidaceae North America j subspontaneous N FR<br />
Malus domestica Borkhausen Rosaceae Western Asia p subspontaneous C
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 146<br />
Species<br />
Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Matricaria discoidea DC. Asteraceae Asia t unknown N R<br />
Meconopsis cambrica (L.) Papaveraceae Europa h unknown N R<br />
Medicago polymorpha L. Fabaceae Mediterranean u unknown N R<br />
Medicago sativa L. Fabaceae Mediterranean h subspontaneous C<br />
Melilotus <strong>in</strong>dicus (L.) Fabaceae Eurasia t unknown N R<br />
Melilotus sulcatus Desfonta<strong>in</strong>es Fabaceae Mediterranean t unknown N R<br />
Mespilus germanica L. Rosaceae Eurasia p naturalized C<br />
Mimulus guttatus DC. Scrophulariaceae North America g naturalized N H<br />
Muhlenbergia schreberi Gmel<strong>in</strong> Poaceae North America h unknown N R<br />
Narcissus <strong>in</strong>comparabilis Miller Amaryllidaceae Europa g subspontaneous C<br />
Narcissus medioluteus Miller Amaryllidaceae Unknown g naturalized N R<br />
Nigella damascena L. Ranunculaceae Mediterranean t subspontaneous C<br />
Nonea erecta Bernhardi Borag<strong>in</strong>aceae Asia h unknown N R<br />
Nonea lutea (Desrousseaux) Borag<strong>in</strong>aceae Eurasia u unknown N R<br />
Nymphoides peltata (Gmel<strong>in</strong>) Menyanthaceae Eurasia a naturalized N E<br />
Oenothera biennis L. Onagraceae North America u unknown N R<br />
Oenothera glazioviana Micheli Onagraceae Unknown u unknown N R<br />
Oenothera parviflora L. Onagraceae North America u unknown N R<br />
Oplismenus undulatifolius (Ardu<strong>in</strong>o) Poaceae Eurasia c naturalized F<br />
Opuntia humifusa (Raf<strong>in</strong>esque) Cactaceae North America c unknown N S<br />
Opuntia imbricata (Haworth) Cactaceae North America c unknown N S<br />
Ornithogalum nutans L. Liliaceae Eurasia g unknown N R<br />
Oxalis fontana Bunge Oxalidaceae Unknown u unknown N R<br />
Panicum capillare L. Poaceae North America t naturalized N R<br />
Panicum dichotomiflorum Michaux Poaceae North America t naturalized N R<br />
Papaver apulum Tenore Papaveraceae Mediterranean u adventive R<br />
Papaver croceum Ledebour Papaveraceae Asia h naturalized N M<br />
Papaver somniferum L. Papaveraceae Unknown t subspontaneous N R<br />
Parthenocissus qu<strong>in</strong>quefolia (L.) Vitaceae North America p naturalized N F<br />
Parthenocissus tricuspidata (Siebold et Zuccar<strong>in</strong>i) Vitaceae Western Asia p subspontaneous N C<br />
Paspalum dilatatum Poiret Poaceae South America g adventive H<br />
Paulownia tomentosa (Thunberg) Bignoniaceae Asia p subspontaneous N F<br />
Phacelia tanacetifolia Bentham Hydrophyllaceae North America t subspontaneous N R<br />
Phalaris canariensis L. Poaceae Mediterranean t adventive R<br />
Philadelphus coronarius L. Philadelphaceae Europa n naturalized N F<br />
Physalis alkekengi L. Solanaceae Eurasia g naturalized N R<br />
Physocarpus opulifolius (L.) Rosaceae North America n naturalized N F<br />
Phyteuma nigrum F.W. Schmidt Campanulaceae Europa h adventive F<br />
Phytolacca americana L. Phytolaccaceae North America h naturalized N R<br />
Phytolacca esculenta Van Houtte Phytolaccaceae Western Asia h naturalized N R<br />
Pimp<strong>in</strong>ella peregr<strong>in</strong>a L. Apiaceae Mediterranean t adventive R<br />
Pisum sativum L. Fabaceae Eurasia t naturalized N R
10 > Plants – Planta 147<br />
Species<br />
Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Plantago arenaria Waldste<strong>in</strong> et Kitaibel Plantag<strong>in</strong>aceae Eurasia t unknown N R<br />
Polygonum orientale L. Polygonaceae Asia t subspontaneous R<br />
Polygonum polystachyum Meissner Polygonaceae Asia g naturalized N R<br />
Polypogon monspeliensis (L.) Poaceae Eurasia t adventive R<br />
Pontederia cordata L. Pontederiaceae North America g adventive E<br />
Potentilla <strong>in</strong>termedia L. Rosaceae Europa h adventive R<br />
Potentilla recta L. Rosaceae Eurasia h unknown N R<br />
Prunus cerasus L. Rosaceae Western Asia p naturalized N F<br />
Prunus dulcis (Miller) Rosaceae Western Asia p naturalized N F<br />
Prunus laurocerasus L. Rosaceae Eurasia i naturalized N F<br />
Prunus serot<strong>in</strong>a Ehrhart Rosaceae North America p naturalized N F<br />
Pseudotsuga menziesii (Mirbel) P<strong>in</strong>aceae North America i naturalized N F<br />
Pueraria hirsuta (Thunberg) Fabaceae Western Asia p subspontaneous N FR<br />
Punica granatum L. Punicaceae Western Asia n subspontaneous C<br />
Pyrus pyraster (L.) Rosaceae Eurasia p subspontaneous FC<br />
Quercus rubra L. Fagaceae North America p naturalized N F<br />
Ranunculus muricatus L. Ranunculaceae Mediterranean t adventive H<br />
Raphanus sativus L. Brassicaceae Mediterranean u subspontaneous C<br />
Rapistrum perenne (L.) Brassicaceae Europa h adventive R<br />
Reynoutria japonica Houttuyn Polygonaceae Asia g naturalized N R<br />
Reynoutria sachal<strong>in</strong>ensis (F. Schmidt) Polygonaceae Asia g subspontaneous N R<br />
Rhus typh<strong>in</strong>a L. Anacardiaceae North America p naturalized N P<br />
Rob<strong>in</strong>ia pseudoacacia L. Fabaceae North America p naturalized N F<br />
Rorippa austriaca (Crantz) Brassicaceae Europa h adventive H<br />
Rosa rugosa Thunberg Rosaceae Western Asia n subspontaneous N R<br />
Rostraria cristata (L.) Poaceae Mediterranean h adventive R<br />
Rubia t<strong>in</strong>ctorum L. Rubiaceae Eurasia h naturalized N R<br />
Rubus armeniacus Focke Rosaceae Caucasus n naturalized N F<br />
Rudbeckia hirta L. Asteraceae North America u subspontaneous C<br />
Rudbeckia lac<strong>in</strong>iata L. Asteraceae North America g subspontaneous N R<br />
Rumex longifolius DC. Polygonaceae Eurasia h unknown N P<br />
Rumex palustris J.E. Smith Polygonaceae Eurasia u adventive R<br />
Rumex thyrsiflorus F<strong>in</strong>gerhuth Polygonaceae Eurasia h unknown N R<br />
Rumex confertus Willdenow Polygonaceae Asia h naturalized N P<br />
Salvia sylvestris L. Lamiaceae Eurasia h naturalized N R<br />
Salvia verbenaca L. Lamiaceae Mediterranean h adventive S<br />
Salvia verticillata L. Lamiaceae Mediterranean h unknown N R<br />
Sarracenia purpurea L. Sarraceniaceae North America h naturalized H<br />
Saxifraga hirsuta L. Saxifragaceae Europa h subspontaneous C<br />
Saxifraga stolonifera Meerburgh Saxifragaceae Western Asia h naturalized N P<br />
Saxifraga umbrosa L. Saxifragaceae Europa c subspontaneous C<br />
Scabiosa ochroleuca L. Dipsacaceae Europa h naturalized N S
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 148<br />
Species<br />
Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Scilla non-scripta (L.) Liliaceae Europa g naturalized N F<br />
Scrophularia vernalis L. Scrophulariaceae Mediterranean u adventive N R<br />
Sedum sarmentosum Bunge Crassulaceae Asia c naturalized C<br />
Sedum sediforme (Jacqu<strong>in</strong>) Crassulaceae Mediterranean c adventive P<br />
Sedum spurium M. Bieberste<strong>in</strong> Crassulaceae Western Asia c subspontaneous N R<br />
Sedum hispanicum L. Crassulaceae Europa u unknown P<br />
Senecio <strong>in</strong>aequidens DC. Asteraceae South Africa u naturalized N R<br />
Senecio rupestris Waldste<strong>in</strong> et Kitaibel Asteraceae Europa u unknown N R<br />
Setaria italica (L.) Poaceae Unknown t subspontaneous RC<br />
Silene conica L. Caryophyllaceae Mediterranean t adventive R<br />
Silene dichotoma Ehrhart Caryophyllaceae Europa u adventive R<br />
S<strong>in</strong>apis alba L. Brassicaceae Mediterranean t subspontaneous RC<br />
Sisymbrium altissimum L. Brassicaceae Eurasia u unknown N R<br />
Sisymbrium irio L. Brassicaceae Mediterranean u unknown N R<br />
Sisymbrium loeselii L. Brassicaceae Eurasia u unknown N R<br />
Sisyr<strong>in</strong>chium montanum Greene Iridaceae North America h unknown N H<br />
Solanum sublobatum Roemer et Schultes Solanaceae South America t naturalized N R<br />
Solidago canadensis L. Asteraceae North America g naturalized N R<br />
Solidago gigantea Aiton Asteraceae North America g naturalized N H<br />
Solidago gram<strong>in</strong>ifolia (L.) Asteraceae North America g naturalized N R<br />
Sorghum halepense (L.) Poaceae Unknown h unknown N R<br />
Sorghum vulgare Persoon Poaceae Western Asia t adventive C<br />
Spiraea salicifolia L. Rosaceae Eurasia n subspontaneous N R<br />
Spiraea ulmifolia Scopoli Rosaceae Eurasia n subspontaneous N R<br />
Staphylea p<strong>in</strong>nata L. Staphyleaceae Eurasia n subspontaneous FC<br />
Stratiotes aloides L. Hydrocharitaceae Eurasia a naturalized N H<br />
Symphoricarpos albus (L.) Caprifoliaceae North America n naturalized N F<br />
Symphytum asperum Lepech<strong>in</strong> Borag<strong>in</strong>aceae Caucasus h naturalized N R<br />
Tanacetum c<strong>in</strong>erariifolium (Treviranus) Asteraceae Europa h naturalized N P<br />
Thlaspi alliaceum L. Brassicaceae Europa u naturalized N R<br />
Tolpis barbata (L.) Asteraceae Mediterranean t adventive R<br />
Tordylium maximum L. Apiaceae Mediterranean u adventive R<br />
Torilis leptophylla (L.) Apiaceae Mediterranean t adventive R<br />
Torilis nodosa (L.) Apiaceae Europa t unknown N R<br />
Trachycarpus fortunei (Hooker) Palmae Asia i subspontaneous N F<br />
Tragopogon crocifolius L. Asteraceae Mediterranean u adventive S<br />
Tribulus terrestris L. Zygophyllaceae Mediterranean t adventive R<br />
Trifolium alexandr<strong>in</strong>um L. Fabaceae Mediterranean t subspontaneous N R<br />
Trifolium hybridum L. Fabaceae Europa u naturalized N R<br />
Trifolium <strong>in</strong>carnatum L. Fabaceae Europa u subspontaneous N G<br />
Trifolium resup<strong>in</strong>atum L. Fabaceae Mediterranean u naturalized N R<br />
Trifolium suaveolens Willdenow Fabaceae Mediterranean t naturalized N R
10 > Plants – Planta 149<br />
Species Family Orig<strong>in</strong> Life form Status Ecol. group<br />
Tulipa didieri Jordan Liliaceae Western Asia g unknown N R<br />
Tulipa grengiolensis Thommen Liliaceae Unknown g unknown N R<br />
Typha laxmannii Lepech<strong>in</strong> Typhaceae Eurasia g adventive H<br />
Ulex europaeus L. Fabaceae Europa n naturalized N F<br />
Ulmus laevis Pallas Ulmaceae Europa p subspontaneous C<br />
Vacc<strong>in</strong>ium macrocarpon Aiton Ericaceae North America z naturalized N H<br />
Valerianella eriocarpa Desvaux Valerianaceae Mediterranean t adventive R<br />
Vallisneria spiralis L. Hydrocharitaceae Unknown a naturalized N E<br />
Veronica filiformis Smith Scrophulariaceae Eurasia h unknown N G<br />
Veronica peregr<strong>in</strong>a L. Scrophulariaceae North America t unknown N R<br />
Veronica persica Poiret Scrophulariaceae Western Asia u unknown N R<br />
Vicia hybrida L. Fabaceae Europa u unknown N R<br />
Vicia lutea L. Fabaceae Mediterranean u adventive N R<br />
Vicia pannonica Crantz Fabaceae Europa u adventive N R<br />
Vicia peregr<strong>in</strong>a L. Fabaceae Mediterranean t adventive R<br />
Vicia sativa L. Fabaceae Mediterranean u subspontaneous N R<br />
V<strong>in</strong>ca major L. Apocynaceae Eurasia z naturalized N M<br />
Viola obliqua Hill Violaceae North America g naturalized N F<br />
Vitis v<strong>in</strong>ifera L. Vitaceae Europa p subspontaneous C<br />
Vulpia ciliata Dumortier Poaceae Mediterranean t adventive R<br />
Xanthium italicum Moretti Asteraceae North America t subspontaneous N G<br />
Xanthium sp<strong>in</strong>osum L. Asteraceae South America t unknown N R<br />
Xeranthemum annuum L. Asteraceae Mediterranean t adventive R<br />
* Exclud<strong>in</strong>g cultivated plants which have not escaped <strong>in</strong> the environment<br />
Life form: p, deciduous tree; i, evergreen tree; n, deciduous shrub; j, evergreen shrub; z, woody chamaephytic plant (small shrub); c, herbaceous chamaephytic plant; hhemicryptophytic plant<br />
(perennial herb); g, geophytic plant (perennial herb with tubers, bulbs or rhizomes); t, annual plant; u, biennal plant; a, aquatic plant.<br />
Ecological group: R, ruderal plant; P, pioneer plant; F, forest plant; M, mounta<strong>in</strong> plant; E, aquatic plant; H, marsh plant; S, dry meadow plant; G, meadow plant; C, cultivated plant; N, neophyte<br />
(accord<strong>in</strong>g to Lauber and Wagner, 1998).<br />
Tab. 10.7 > Plant families with <strong>alien</strong> and naturalized <strong>species</strong> <strong>in</strong> <strong>Switzerland</strong>.*<br />
Family Taxonomy No. native<br />
<strong>species</strong><br />
No. <strong>alien</strong><br />
<strong>species</strong><br />
No. naturalized<br />
<strong>species</strong><br />
Total no.<br />
<strong>species</strong><br />
% <strong>alien</strong><br />
<strong>species</strong><br />
% naturalized<br />
<strong>species</strong><br />
% naturalized/<br />
<strong>alien</strong><br />
Aceraceae Dicot 4 1 0 5 20.0 0.0 0.0<br />
Agavaceae Monoc 0 1 1 1 100.0 100.0 100.0<br />
Aizoaceae Dicot 0 1 0 1 100.0 0.0 0.0<br />
Alismataceae Monoc 5 1 1 6 16.7 16.7 100.0<br />
Amarantaceae Dicot 0 9 0 9 100.0 0.0 0.0<br />
Amaryllidaceae Monoc 7 2 1 9 22.2 11.1 50.0<br />
Anacardiaceae Dicot 1 1 1 2 50.0 50.0 100.0<br />
Apiaceae Dicot 85 7 2 92 7.6 2.2 28.6<br />
Apocynaceae Dicot 1 1 1 2 50.0 50.0 100.0<br />
Araceae Monoc 3 3 2 6 50.0 33.3 66.7
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 150<br />
Family Taxonomy No. native<br />
<strong>species</strong><br />
No. <strong>alien</strong><br />
<strong>species</strong><br />
No. naturalized<br />
<strong>species</strong><br />
Total no.<br />
<strong>species</strong><br />
% <strong>alien</strong><br />
<strong>species</strong><br />
% naturalized<br />
<strong>species</strong><br />
% naturalized/<br />
<strong>alien</strong><br />
Asclepiadaceae Dicot 1 1 0 2 50.0 0.0 0.0<br />
Asteraceae Dicot 298 42 10 340 12.4 2.9 23.8<br />
Balsam<strong>in</strong>aceae Dicot 1 3 2 4 75.0 50.0 66.7<br />
Berberidaceae Dicot 1 2 1 3 66.7 33.3 50.0<br />
Bignoniaceae Dicot 0 1 0 1 100.0 0.0 0.0<br />
Borag<strong>in</strong>aceae Dicot 35 6 1 41 14.6 2.4 16.7<br />
Brassicaceae Dicot 129 35 7 164 21.3 4.3 20.0<br />
Buddlejaceae Dicot 0 1 1 1 100.0 100.0 100.0<br />
Cactaceae Dicot 0 2 0 2 100.0 0.0 0.0<br />
Campanulaceae Dicot 34 2 0 36 5.6 0.0 0.0<br />
Cannabaceae Dicot 1 1 0 2 50.0 0.0 0.0<br />
Caprifoliaceae Dicot 12 6 2 18 33.3 11.1 33.3<br />
Caryophyllaceae Dicot 117 6 1 123 4.9 0.8 16.7<br />
Chenopodiaceae Dicot 21 6 1 27 22.2 3.7 16.7<br />
Commel<strong>in</strong>aceae Monoc 0 1 1 1 100.0 100.0 100.0<br />
Cornaceae Dicot 2 1 1 3 33.3 33.3 100.0<br />
Crassulaceae Dicot 26 3 1 29 10.3 3.4 33.3<br />
Cupressaceae Gymnosperm 2 2 0 4 50.0 0.0 0.0<br />
Cuscutaceae Dicot 2 2 0 4 50.0 0.0 0.0<br />
Cyperaceae Monoc 131 4 2 135 3.0 1.5 50.0<br />
Dipsacaceae Dicot 15 2 1 17 11.8 5.9 50.0<br />
Droseraceae Dicot 4 1 0 5 20.0 0.0 0.0<br />
Elaeagnaceae Dicot 1 1 0 2 50.0 0.0 0.0<br />
Ericaceae Dicot 15 2 2 17 11.8 11.8 100.0<br />
Euphorbiaceae Dicot 17 8 1 25 32.0 4.0 12.5<br />
Fabaceae Dicot 135 25 6 160 15.6 3.8 24.0<br />
Fagaceae Dicot 7 1 1 8 12.5 12.5 100.0<br />
Fumariaceae Dicot 8 1 0 9 11.1 0.0 0.0<br />
Geraniaceae Dicot 20 4 0 24 16.7 0.0 0.0<br />
Hydrocharitaceae Monoc 1 6 5 7 85.7 71.4 83.3<br />
Hydrophyllaceae Dicot 0 1 0 1 100.0 0.0 0.0<br />
Hypericaceae Dicot 12 1 0 13 7.7 0.0 0.0<br />
Iridaceae Monoc 8 6 1 14 42.9 7.1 16.7<br />
Juncaceae Monoc 35 1 0 36 2.8 0.0 0.0<br />
Lamiaceae Dicot 82 4 2 86 4.7 2.3 50.0<br />
Lauraceae Dicot 0 1 1 1 100.0 100.0 100.0<br />
Lemnaceae Monoc 4 1 1 5 20.0 20.0 100.0<br />
Liliaceae Monoc 54 9 2 63 14.3 3.2 22.2<br />
L<strong>in</strong>aceae Dicot 4 2 1 6 33.3 16.7 50.0<br />
Malvaceae Dicot 4 4 0 8 50.0 0.0 0.0<br />
Mimosaceae Dicot 0 1 0 1 100.0 0.0 0.0<br />
Moraceae Dicot 1 1 0 2 50.0 0.0 0.0
10 > Plants – Planta 151<br />
Family Taxonomy No. native<br />
<strong>species</strong><br />
No. <strong>alien</strong><br />
<strong>species</strong><br />
No. naturalized<br />
<strong>species</strong><br />
Total no.<br />
<strong>species</strong><br />
% <strong>alien</strong><br />
<strong>species</strong><br />
% naturalized<br />
<strong>species</strong><br />
% naturalized/<br />
<strong>alien</strong><br />
Oleaceae Dicot 3 2 0 5 40.0 0.0 0.0<br />
Onagraceae Dicot 21 5 1 26 19.2 3.8 20.0<br />
Oxalidaceae Dicot 2 1 0 3 33.3 0.0 0.0<br />
Palmae Monoc 0 1 0 1 100.0 0.0 0.0<br />
Papaveraceae Dicot 10 5 1 15 33.3 6.7 20.0<br />
Philadelphaceae Dicot 0 1 1 1 100.0 100.0 100.0<br />
Phytolaccaceae Dicot 0 2 2 2 100.0 100.0 100.0<br />
P<strong>in</strong>aceae Gymnosperm 7 2 2 9 22.2 22.2 100.0<br />
Plantag<strong>in</strong>aceae Dicot 8 2 0 10 20.0 0.0 0.0<br />
Poaceae Monoc 189 31 5 220 14.1 2.3 16.1<br />
Polygonaceae Dicot 29 10 3 39 25.6 7.7 30.0<br />
Pontederiaceae Monoc 0 1 0 1 100.0 0.0 0.0<br />
Punicaceae Dicot 0 1 0 1 100.0 0.0 0.0<br />
Ranunculaceae Dicot 93 5 0 98 5.1 0.0 0.0<br />
Rosaceae Dicot 126 20 9 146 13.7 6.2 45.0<br />
Rubiaceae Dicot 34 3 1 37 8.1 2.7 33.3<br />
Salicaceae Dicot 33 1 0 34 2.9 0.0 0.0<br />
Sarraceniaceae Dicot 0 1 1 1 100.0 100.0 100.0<br />
Saxifragaceae Dicot 30 4 1 34 11.8 2.9 25.0<br />
Scrophulariaceae Dicot 106 12 5 118 10.2 4.2 41.7<br />
Simourabaceae Dicot 0 1 1 1 100.0 100.0 100.0<br />
Solanaceae Dicot 5 5 2 10 50.0 20.0 40.0<br />
Staphyleaceae Dicot 0 1 0 1 100.0 0.0 0.0<br />
Ulmaceae Dicot 3 1 0 4 25.0 0.0 0.0<br />
Valerianaceae Dicot 18 2 1 20 10.0 5.0 50.0<br />
Violaceae Dicot 25 1 1 26 3.8 3.8 100.0<br />
Vitaceae Dicot 0 3 1 3 100.0 33.3 33.3<br />
Zygopphyllaceae Dicot 0 1 0 1 100.0 0.0 0.0<br />
*Exclud<strong>in</strong>g cultivated plants which have not escaped <strong>in</strong>to the environment<br />
Tab. 10.8 > <strong>Invasive</strong> plant <strong>species</strong> <strong>in</strong> Europe.<br />
Fact<br />
Sheet<br />
Species Ecol. Life<br />
group form<br />
Status CH<br />
1<br />
Austria 2 France 3 Spa<strong>in</strong> 8 Portugal<br />
7<br />
Abutilon theophrasti Medik. R t adventive (econ.) x xx xx x<br />
yes Acer negundo L. F p subspont. xxx;<br />
(econ.)<br />
Scotland<br />
9<br />
Hungary<br />
5<br />
xxx x xxx<br />
Aesculus hippocastanum L. C p subspont. xxx<br />
Agave americana L. R j naturalized + M xxx xx<br />
yes Ailanthus altissima (Miller) P p naturalized xxx xxx xxx xxx xxx xxx xxx<br />
Allium paradoxum (M. v. Bieberste<strong>in</strong>) C g subspont. xx<br />
Amaranthus albus L. R t unknown x xxx<br />
Italy 6 Germany 4
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 152<br />
Fact<br />
Sheet<br />
Species Ecol. Life<br />
group form<br />
Status CH<br />
1<br />
Austria 2 France 3 Spa<strong>in</strong> 8 Portugal<br />
7<br />
Scotland<br />
9<br />
Hungary<br />
5<br />
Italy 6 Germany 4<br />
Amaranthus blitum L. R t unknown x x<br />
Amaranthus cruentus L. R t subspont. xxx<br />
Amaranthus deflexus L. R u unknown x xxx xxx xxx<br />
Amaranthus retroflexus L. R t unknown econ. x xxx xxx xxx<br />
yes Ambrosia artemisiifolia L. R t adventive xxx xx; econ. xxx x x xxx xxx<br />
yes Amorpha fruticosa L. H n subspont. x xx ++ M xxx xxx<br />
Artemisia annua L. R t adventive x<br />
yes Artemisia verlotiorum Lamotte R g unknown xxx xxx x x xxx<br />
Arundo donax L. H g subspont. xxx xx<br />
yes Asclepias syriaca L. C R g subspont. xx + M xxx<br />
yes Aster lanceolatus Willdenow C g subspont. xxx ++ M xx xxx Aster spp.<br />
Aster novae-angliae L. H g subspont.<br />
yes Aster novi-belgii L. H c subspont. xxx xxx<br />
Aubrieta deltoidea (L.) S c naturalized<br />
Avena sativa L. C t subspont. x<br />
Bidens bip<strong>in</strong>nata L. R t naturalized xxx<br />
Bidens connata Willdenow H t naturalized +++ A<br />
yes Bidens frondosa L. R t naturalized xxx xxx xx xxx xxx<br />
Bidens subalternans DC. R t adventive xx<br />
Brassica napus L. C u subspont. x xx<br />
yes Buddleja davidii Franchet P n naturalized xxx xx xxx xxx xx<br />
yes Bunias orientalis L. R h unknown x ++ C xxx<br />
Cerastium tomentosum L. C c subspont. xx<br />
Chenopodium ambrosioides L. R t adventive +++ M x xxx<br />
Chrysanthemum Segetum L. R t unknown xx<br />
Cicerbita macrophylla (Willdenow) C g subspont. xx<br />
yes Conyza canadensis (L.) P u unknown +++ C xxx xxx xxx xxx xxx<br />
yes Cornus sericea L. F n naturalized x<br />
Coronopus didymus (L.) R u adventive x xxx<br />
Cotoneaster horizontalis Decne C j subspont. xx<br />
Crepis nemauensis Gouan R t naturalized<br />
Cymbalaria muralis P. Gaertn., B. Mey. &<br />
Scherb.<br />
P c unknown xx<br />
yes Cyperus eragrostis Lamarck H h unknown xxx xxx xx<br />
yes Cyperus esculentus L. H g unknown x xxx<br />
Cyperus rotundus L. H g naturalized<br />
Datura stramonium L. R t unknown x x xxx xxx xxx<br />
yes Duchesnea <strong>in</strong>dica (Andrews) F h naturalized xx<br />
Eleus<strong>in</strong>e <strong>in</strong>dica (L.) R t naturalized xx x xxx<br />
yes Elodea canadensis Michaux E a unknown x xxx +++ C xxx xx xx xxx<br />
yes Elodea nuttallii (Planchon) E a naturalized xxx xx ++ C xx<br />
yes Epilobium ciliatum Raf<strong>in</strong>esque R h unknown xxx xxx x
10 > Plants – Planta 153<br />
Fact Species Ecol. Life Status CH Austria<br />
Sheet<br />
group form<br />
1<br />
2 France3 Spa<strong>in</strong>8 Portugal7<br />
Scotland9<br />
Hungary5<br />
Italy6 Germany4 Erigeron annuus (L.) R u naturalized xx xxx xxx<br />
Erigeron karv<strong>in</strong>skianus DC. P h unknown + A xx<br />
Euphorbia maculata L. R t unknown x x<br />
Euphorbia prostrata Aiton R t adventive x<br />
Ficus carica L. C p subspont. x<br />
Galega offic<strong>in</strong>alis L. G h subspont. ++ C<br />
yes Gal<strong>in</strong>soga ciliata (Raf<strong>in</strong>esque) R t unknown econ. x x xxx xxx<br />
yes Gal<strong>in</strong>soga parviflora Cavanilles R t unknown econ. x xx xxx xxx<br />
Gleditsia triacanthos L. C p subspont. xx<br />
Glyceria striata (Lamarck) H g unknown xx<br />
yes Helianthus tuberosus L. R g subspont. x xxx xxx xx xxx xxx xxx<br />
yes Heracleum mantegazzianum Sommier &<br />
Levier<br />
R h naturalized xxx xx; econ. xxx xxx xxx xxx<br />
yes Impatiens balfourii Hooker F. R t unknown + C<br />
yes Impatiens glandulifera Royle R t naturalized xxx xxx xxx xx xxx xxx xxx<br />
yes Impatiens parviflora DC. F t naturalized xxx xxx xxx<br />
Juncus tenuis Willdenow R h unknown ++ C<br />
Lagarosiphon major (Ridley) E a naturalized +++ A<br />
Lemna m<strong>in</strong>uta Humboldt et al. E a naturalized xxx<br />
Lepidium virg<strong>in</strong>icum L. R u unknown x x<br />
Ligustrum lucidum Aiton C i subspont. xx<br />
yes Lonicera henryi Hemsley F i subspont. x xx<br />
yes Lonicera japonica Thunberg F i naturalized xxx<br />
yes Ludwigia grandiflora (Michaux) E a naturalized xxx xxx<br />
yes Lup<strong>in</strong>us polyphyllus L<strong>in</strong>dley F h subspont. x xx<br />
Lycium barbarum L. C n subspont. ++ A<br />
Lysichiton americanus Hultén & H. St. John H g naturalized xxx<br />
Lysimachia punctata L. C H h subspont. xx<br />
yes Mahonia aquifolium (Pursh) F R j subspont. x xx<br />
Matricaria discoidea DC. R t unknown + C xx xxx<br />
Meconopsis cambrica (L.) R h unknown xx<br />
Medicago sativa L. C h subspont. x<br />
Mimulus guttatus DC. H g naturalized xx<br />
Morus alba L. C p subspont. ++ C<br />
Nicandra physalodes (L.) C R t subspont. xxx<br />
yes Oenothera biennis L. R u unknown xxx x x xxx<br />
Oenothera glazioviana Micheli R u unknown x x<br />
Opuntia imbricata (Haworth) S c unknown xxx<br />
Oxalis fontana Bunge R u unknown x<br />
Panicum capillare L. R t naturalized x x<br />
Panicum dichotomiflorum Michaux R t naturalized x xxx<br />
Parthenocissus qu<strong>in</strong>quefolia (L.) F p naturalized
An <strong>in</strong>ventory of <strong>alien</strong> <strong>species</strong> and their threat to biodiversity and economy <strong>in</strong> <strong>Switzerland</strong> FOEN 2006 154<br />
Fact<br />
Sheet<br />
Species Ecol. Life<br />
group form<br />
Parthenocissus tricuspidata (Siebold et<br />
Zuccar<strong>in</strong>i)<br />
Status CH<br />
1<br />
Austria 2 France 3 Spa<strong>in</strong> 8 Portugal<br />
7<br />
Scotland<br />
9<br />
Hungary<br />
5<br />
Italy 6 Germany 4<br />
C p subspont. xx xxx Parthen. spp.<br />
yes Paspalum dilatatum Poiret H g adventive xxx xxx xxx<br />
Paulownia tomentosa (Thunberg) F p subspont.<br />
Phalaris canariensis L. R t adventive x<br />
yes Phytolacca americana L. R h naturalized x xx xxx xxx<br />
Phytolacca esculenta Van Houtte R h naturalized<br />
Polygonum orientale L. R t subspont. x<br />
yes Polygonum polystachyum Meissner R g naturalized xxx<br />
yes Prunus laurocerasus L. F i naturalized x xx<br />
yes Prunus serot<strong>in</strong>a Ehrhart F p naturalized xxx xx xxx<br />
Pseudotsuga menziesii (Mirbel) F i naturalized xx<br />
yes Pueraria lobata (Willdenow) F R p subspont. x<br />
yes Reynoutria japonica Houttuyn R g naturalized xxx xxx;<br />
econ.<br />
yes Reynoutria sachal<strong>in</strong>ensis (F. Schmidt) R g subspont. xxx xx;<br />
(econ.)<br />
yes Rhus typh<strong>in</strong>a L. P p naturalized xxx<br />
xxx xxx x xxx xxx xxx<br />
xxx xx xxx xxx<br />
Ribes rubrum L. C n subspont. x<br />
yes Rob<strong>in</strong>ia pseudoacacia L. F p naturalized xxx xxx;<br />
econ.<br />
Rorippa austriaca (Crantz) H h adventive + C<br />
xxx xxx xx xxx xxx<br />
yes Rosa rugosa Thunberg R n subspont. xxx<br />
Rubia t<strong>in</strong>ctorum L. R h naturalized x<br />
yes Rubus armeniacus Focke F n naturalized xxx<br />
Rudbeckia hirta L. C u subspont. xxx<br />
yes Rudbeckia lac<strong>in</strong>iata L. C g subspont. xxx xxx<br />
Rumex longifolius DC. P h unknown<br />
Rumex patientia L. C R h subspont. + C<br />
Rumex thyrsiflorus F<strong>in</strong>gerhuth R h unknown ++ C<br />
Rumex confertus Willdenow P h naturalized<br />
Sedum spurium M. Bieberste<strong>in</strong> R c subspont. x<br />
Sedum hispanicum L. P u unknown<br />
yes Senecio <strong>in</strong>aequidens DC. R u naturalized xxx x xxx xxx xxx<br />
Senecio rupestris Waldste<strong>in</strong> et Kitaibel R u unknown x<br />
Solanum sublobatum Roemer et Schultes R t naturalized ++ M<br />
yes Solidago canadensis L. R g naturalized xxx xxx +++ C xxx xxx<br />
yes Solidago gigantea Aiton H g naturalized xxx xxx;<br />
(econ)<br />
Sorghum halepense (L.) R h unknown x xx x<br />
+++ C x xxx xxx xxx<br />
Symphoricarpos albus (L.) F n naturalized xxx<br />
Symphytum asperum Lepech<strong>in</strong> R h naturalized +++ C xx<br />
Syr<strong>in</strong>ga vulgaris L. C n subspont. xx
10 > Plants – Planta 155<br />
Fact<br />
Sheet<br />
Species Ecol. Life<br />
group form<br />
Status CH<br />
1<br />
Austria 2 France 3 Spa<strong>in</strong> 8 Portugal<br />
7<br />
Tanacetum parthenium (L.) C h subspont. xx<br />
Tetragonia tetragonioides (Pallas) C t subspont. ++ M x<br />
Trachycarpus fortunei (Hooker) F i subspont. x<br />
Trifolium <strong>in</strong>carnatum L. G u subspont. x<br />
Trifolium resup<strong>in</strong>atum L. R u naturalized x<br />
Ulex europaeus L. F n naturalized ++ A<br />
Vacc<strong>in</strong>ium macrocarpon Aiton H z naturalized<br />
Veronica filiformis Smith G h unknown xx<br />
Veronica peregr<strong>in</strong>a L. R t unknown + C x<br />
Veronica persica Poiret R u unknown xx xxx xx<br />
Xanthium sp<strong>in</strong>osum L. R t unknown + M xx xx<br />
xxx, <strong>in</strong>vasive <strong>species</strong> (black list); xx, potentially or moderately <strong>in</strong>vasive <strong>species</strong>; <strong>species</strong> <strong>in</strong> expansion, locally <strong>in</strong>vasive; x, <strong>species</strong> present; need to be followed up (watch list).<br />
+++ – For France: <strong>in</strong>vasive <strong>species</strong> <strong>in</strong> one sector only: M (Mediterranean area); A (Atlantic area); C (cont<strong>in</strong>ental area);<br />
++ – For France: potentially <strong>in</strong>vasive <strong>in</strong> one sector only: M, A or C;<br />
+ – For France: <strong>species</strong> present, which need to be followed up (Watch List), <strong>in</strong> one sector only: M, A or C.<br />
1 <strong>Switzerland</strong>: - CPS-SKEW (http://www.cps-skew.ch/)<br />
2 Austria: Essl, F. and W. Rabitsch (eds) (2002) Neobiota <strong>in</strong> Österreich. Federal Environment Agency, 432 pp.<br />
3 France: Muller, S. (2004) Plantes <strong>in</strong>vasives en France. Paris, Muséum National d’Histoire Naturelle. Patrimo<strong>in</strong>es naturels; 62, 176 pp.<br />
4 Germany: From a prelim<strong>in</strong>ary EPPO list of <strong>in</strong>vasive <strong>alien</strong> plants for the EPPO region (plants from Germany <strong>in</strong>clude mostly <strong>species</strong> <strong>in</strong> the category xxx)<br />
5 Hungary: <strong>Invasive</strong> <strong>alien</strong> <strong>species</strong> <strong>in</strong> Hungary. National Ecological Network No. 6 (the list <strong>in</strong>cludes only <strong>in</strong>vasive plant <strong>species</strong> <strong>in</strong> Hungarian protected areas)<br />
6 Italy: Laura Celesti, pers. comm. (2003). The <strong>species</strong> given for Italy are those most frequent <strong>in</strong> northern Italy<br />
7 Portugal: De Almeida, J.D. (1999) Flora exotica subespontanea de Portugal cont<strong>in</strong>ental. Universidade de Coimbra, 151 pp.<br />
8 Spa<strong>in</strong>: Dana, E.D., Sanz-Elorza, M. & E. Sobr<strong>in</strong>o (2001) Plant <strong>in</strong>vaders <strong>in</strong> Spa<strong>in</strong>, http://www.ual.es/personal/edana/<strong>alien</strong>plants/checklist.pdf<br />
9 Scotland: Welch, D. et al., (2001) An audit of <strong>alien</strong> <strong>species</strong> <strong>in</strong> Scotland. Scottish Natural Heritage Review No 139, 225 pp.<br />
> Fact sheets<br />
The fact sheets are available at http://www.environment-switzerland.ch/uw-0629-e.<br />
Scotland<br />
9<br />
Hungary<br />
5<br />
Italy 6 Germany 4