Weed seed choice by carabid beetles (Coleoptera : Carabidae): Linking field measurements with laboratory diet assessments

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Weed seed choice by carabid beetles (Coleoptera : Carabidae): Linking field measurements with laboratory

diet assessments

Sandrine Petit, Aline Boursault, David Bohan

To cite this version:

Sandrine Petit, Aline Boursault, David Bohan. Weed seed choice by carabid beetles (Coleoptera :

Carabidae): Linking field measurements with laboratory diet assessments. European Journal of En-

tomology, Czech Entomological Society, 2014, 111 (5), pp.615-620. �10.14411/eje.2014.086�. �hal-

02635407�

INTRODUCTION

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REFERENCES

A

LIGNIER

A., M

EISS

H., P

ETIT

S. & R

EBOUD

X9DULDWLRQRI SRVWGLVSHUVDOZHHGVHHGSUHGDWLRQDFFRUGLQJWRZHHGVSHFLHV VSDFHDQGWLPH²J. Plant Dis. Protect. 21±

B

^AARS

0$&DWFKHVLQSLWIDOOWUDSVLQUHODWLRQWRPHDQGHQ- VLWLHVRIFDUDELGEHHWOHV²Oecologia 41±

B

OHAN

D.A., B

OHAN

A., G

LEN

D.M., S

YMONDSON

W.O.C. & W

ILT

-

SHIRE

C.6SDWLDOG\QDPLFVRISUHGDWLRQE\FDUDELGEHHWOHV RQVOXJV²J. Anim. Ecol. 69±

B

^OHAN

D.A., B

^OURSAULT

A., B

^ROOKS

D. & P

^ETIT

S.1DWLRQ- DOVFDOHUHJXODWLRQRIWKHZHHGVHHGEDQNE\FDUDELGSUHGDWRUV

²J. Appl. Ecol. 48±

B

ROOKS

D.R., S

TORKEY

J., C

LARK

S.J., F

IRBANK

L.G., P

ETIT

S. &

W

OIWOD

I.P. 2012: 7URSKLFOLQNVEHWZHHQIXQFWLRQDOJURXSVRI

arable plants and beetles are stable at a national scale.

²J.

Anim. Ecol. 81: 4–13.

C

ARDINA

J., N

ORQUAY

H.M., S

TINNER

B.R. & M

C

C

ARTNEY

D.A.

1996: 3RVWGLVSHUVDO SUHGDWLRQ RI YHOYHWOHDI Abutilon theo- phrastiVHHGV²Weed Sci. 44: 534–539.

C

HESSON

-0HDVXULQJSUHIHUHQFHLQVHOHFWLYHSUHGDWLRQ² Ecology 59±

C^RAWLEY

M.J. 2013: The R Book, 2nd ed.-RKQ:LOH\DQG6RQV

&KLFKHVWHUSS

D

AVIS

A.S. & R

AGHU

S:HLJKLQJDELRWLFDQGELRWLFLQÀX- HQFHVRQZHHGVHHGSUHGDWLRQ²Weed Res. 50±

D

^AVIS

A.S., D

^EADLOW

D., S

^CHUTTE

B.J. & W

^ESTERMAN

P.R.

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G

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H.R. & C

LAUDIO

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RATTON

C. 6HHG SUHGDWLRQ LQ- FUHDVHVZLWKJURXQGEHHWOHGLYHUVLW\LQD:LVFRQVLQ86$SR- WDWRDJURHFRV\VWHP²Agric. Ecosyst. Environ. 137±

G

^OLDSMITH

H. T

^OFT

S9DULDEOHGHJUHHVRIJUDQLYRU\DQG SK\WRSKDJ\LQLQVHFWLYRURXVFDUDELGEHHWOHV²Pedobiologia

41±

H

ONEK

A., M

ARTINKOVA

Z. & J

AROSIK

V. 2003: *URXQG EHHWOHV

&DUDELGDHDVVHHGSUHGDWRUVEur. J. Entomol. 100±

H

^ONEK

A., S

^ASKA

P. & M

^ARTINKOVA

Z6HDVRQDOYDULDWLRQLQ VHHGSUHGDWLRQE\DGXOWFDUDELGEHHWOHV²Entomol. Exp. Appl.

118±

H

ONEK

A., M

ARTINKOVA

Z., S

ASKA

P. & P

EKAR

S. 6L]HDQG WD[RQRPLFFRQVWUDLQWVGHWHUPLQHWKHVHHGSUHIHUHQFHVRI&DUD ELGDH&ROHRSWHUD²Basic Appl. Ecol. 8±

K

ROMP

B. &DUDELGEHHWOHVLQVXVWDLQDEOHDJULFXOWXUHDUH- YLHZRQSHVWFRQWUROHI¿FDF\FXOWLYDWLRQLPSDFWVDQGHQKDQFH- PHQW²Agric. Ecosyst. Environ. 74±

L

^UNDGREN

J.G. & R

^OSENTRATER

K.A.7KHVWUHQJWKRIVHHGV DQG WKHLU GHVWUXFWLRQ E\ JUDQLYRURXV LQVHFWV ² Arthropod- Plant Interact. 1±

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B.F.J., M

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OOK

L.M.$QDO\VLVRIDVHOHF- WLYHSUHGDWLRQH[SHULPHQW²Am. Nat. 106±

M

^AUCHLINE

A.L., W

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R.J.3RVWGLVSHUVDOVHHGSUHGDWLRQRIQRQWDUJHWZHHGVLQ DUDEOHFURSV²Weed Res. 45±

M

EISS

H., L

AGADEC

L., M

UNIER

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N., W

ALDHARDT

R. & P

ETIT

S.:HHGVHHGSUHGDWLRQLQFUHDVHVZLWKYHJHWDWLRQFRYHU LQ SHUHQQLDO IRUDJH FURSV ² Agric. Ecosyst. Environ. 138

±

M

ENALLED

F.D., S

MITH

R.G., D

AUER

J.T. & F

OX

T.B.,PSDFW RIDJULFXOWXUDOPDQDJHPHQWRQFDUDELGFRPPXQLWLHVDQGZHHG VHHGSUHGDWLRQ²Agric. Ecosyst. Environ. 118±

O’R

OURKE

M.E., H

EGGENSTALLER

A.H., L

IEBMAN

M. & R

ICE

M.E.

3RVWGLVSHUVDO ZHHG VHHG SUHGDWLRQ E\ LQYHUWHEUDWHV LQ FRQYHQWLRQDODQGORZH[WHUQDOLQSXWFURSURWDWLRQV\VWHPV² Agric. Ecosyst. Environ. 116±

R

OBERTS

J.H. & K

ILPATRICK

J.M. 3UHGDWRU IHHGLQJ SUHIHU- HQFHV IRU D EHQWKLF VWUHDP ¿VK (IIHFWV RI YLVLEOH LQMHFWHG PDUNV²J. Freshw. Ecol. 19 ±.

S

^ASKA

P., V

^ANDER

W

^ERF

W.,

^DE

V

^RIES

E. & W

^ESTERMAN

P.R.

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Entomol. Res. 98±

T

^HOMAS

C.F.G., P

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$QQXDOORVVHVRIZHHGVHHGVGXHWRSUHGDWLRQLQRUJDQLF FHUHDO¿HOGV²J. Appl. Ecol. 40±

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