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ناریا یسانش هرشح نمجنا ی همان

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èÒ ƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ éí èê ç èç æèîæ

ƱƹŹŵ ƽźĭŹŚĪƃ źŧř

ƶŤſŹ ƶƄě ƾŝŚƴŤūř ƽŚƷŹŚŤƟŹ ƽƹŹ ƽř

ƽ ƶŤƃ Źřƺų

Aphidoletes aphidimyza (Dip.: Cecidomyiidae)

źĭŹŚĪƃ Ʋſ Śŝ ƶƸūřƺƯ Źŵ

Orius laevigatus (Het.: Anthocoridae)

Ŷǀưů ƽźǀƘƯƝřźƇŚƋŹ

æ

ç ƹ

ƾſŶƴƸƯŚƋŹŶưůř* è

ƹ

é

ƽŹƺƃŚƗŶưůřƹ

è

æ ƵŚǀĭƵƹźĭ ƵŶĪƄƳřŵƾĪƃżě

į

ƱŚŬƳŻƱŚŬƳŻƵŚĮƄƳřŵƽŻŹƹŚƄĩ ç

ƵŶĪƄƷƹĦě į

ƱŚŬƳŻƱŚŬƳŻƵŚĮƄƳřŵƽĥƺƫƺƴĪţƺǀŝƹƽĥƺƫƺƿżǀƟ è

ƵŶĪƄƳřŵƾƘǀŞƏƖŝŚƴƯƹƽŻŹƹŚƄĩžƿŵźě

į ƵŚǀĭ

ƾƳŚŞƛŚŝƭƺƬƗƹƾĪƃżě

ũźĩƱřźƸţƵŚĮƄƳřŵ é

ƱřźƸţŶƳƺƫřŹŸŝƵŚǀĭŢĩźƃ

ƾĪǀƳƹźŤĪƫřŢƀ읹ŚŞţŚĪƯƩƺŘƀƯ*

hamidsarrafm@gmail.com

The effect of interaguild predation on avoidance behavior of the aphidophagous midge, Aphidoletes aphidimyza (Dip.: Cecidomyiidae) on its encounter with the predatory bug Orius laevigatus (Het.:

Anthocoridae)

H. R. S. Moayeri1,2&*, A. R. Mohandesi3&4 and A. Ashouri3

1. Department of Plant Protection, Faculty of Agriculture, University of Zanjan, Zanjan, Iran, 2. Research Institute of Physiology and Biotechnology, University of Zanjan, Zanjan, Iran, 3.Department of Plant Protection, Faculty of Horticulture and Plant Protection, College of Agriculture and Natural Resources, University of Tehran, Karaj, Iran, 4.

Gyah Bazr Alvand Corporation, Tehran, Iran.

*Corresponding author, E-mail: hamidsarrafm@gmail.com

ƵŶǀĪģ źĭŹŚĪƃƲſ Orius laevigatus (Fibber)

ƶƄěƹ źĭŹŚĪƃƽ Aphidoletes aphidimyza Rondani

ƶŤƃŻř į żǀƫŚū



Aphis gossypii Glover

 ƶŝ ƾƯƶƿŸƜţħźŤƄƯŹŚĪƃƱřƺƴƗ

Ŷƴƴĩ ƶƄěƾŝŚƴŤūřƽŚƷŹŚŤƟŹŻřƾųźŝƶƘƫŚƐƯƲƿřŹŵ

źĭŹŚĪƃƽ

ŚƷŹŶƴƳŚƯ ƮŴţƱŚĪƯśŚŴŤƳřƹƵŚǀĭŻřƱŶƃ

ƱƹŹŵźĭŹŚĪƃŚŝƶūřƺƯŹŵƽŹřŸĭ

ƶŤſŹ ƽř O. laevigatus

ŹřźƣƾŝŚƿŻŹřŵŹƺƯ

ŢƟźĭ ƶƄěƶĩŵřŵƱŚƄƳŪƿŚŤƳ

ƵŵŚƯƽŚƷ į A. aphidimyza

ƱƹŹŵźĭŹŚĪƃŻřƾƃŚƳźƐųŽŚƀůřƭŚĮƴƷŹŵ

ƶŤſŹ ũŚŤƳƽřźŝƽř

ƮŴţŻřŵƺų ƾƯƕŚƴŤƯřƱŚĪƯƱōŹŵƽŹřŸĭ

Ŷƴƴĩ ŚƷŹƱřżǀƯƲƿřźŝƱƹżƟř

ƶƄěƽŚƷƹŹLJƱŶƃ

ƶŤƃƽ ƌůŚŝƵŚǀĭŮƐſŻřŹřƺų Źƺ

źĭŹŚĪƃƲſ Ò

ÑÑ ƶŝ ƾƴƘƯŹƺƏ ƂǀŝƽŹřŵ ŵƺŝŶƷŚƃŻřźţ Ò

Ó ŚƷŹƶĩŵřŵƱŚƄƳƾſŹźŝƲƿř

ƹīźŝŮƐſŻřƱŶƃ

īźƯ ƱƹŹŵ ŹŚĪƃ źǀƯƹ

ƶŤſŹ ƶŤƃ ƽŶƃŹ ƪůřźƯ źǀŧŐţ ŢŰţ ƽř

į ƾƯ żǀƫŚū ŶƃŚŝ īźƯ ƱřżǀƯ ŚƷŹ ƹ źǀƯƹ

ƽŚƷƹŹLJ ƱŶƃ

A. aphidimyza

ƵŹƺěŹƺƌůŹŵ

į ƶŝƶŤƃƭƹŵƲſ

ţźţ Ŝǀ ÎÍ ƹ Ò æç ƵŹƺěŹƺƌůŹŵƹŶƇŹŵ

į ƭŹŚƸģƲſ ÐÎ ƹ Ò ÑÑ

ŵƺŝŶƇŹŵ ƶŤƟŚƿƲƿř ŹŚĪƃźƐųƱřżǀƯƲǀŝƍŚŞţŹřŚƷ

ƱƹŹŵźĭŹŚĪƃƎſƺţũŚŤƳƱŶƃ

ƶŤſŹ ƵŵŚƯƵŚĮŤƀƿŻśŚŴŤƳřƹƽř

ƽŚƷ

A. aphidimyza

ƮŴţŢƸū ƾƯŶǀƿŐţřŹƽŹřŸĭ

ŤūřƽŚƷŹŚŤƟŹŻƹźŝŹŵŹŚĪƃŢǀƘưūŹŚŤųŚſŢǀưƷřƶŝƹŶƴĩ ƶƄěƾŝŚƴ

į

ƶŤƃ ƶƳŚƯŚſĨƿŹŵŹřƺų

į ƱƹŹŵƽźĭŹŚĪƃ

ƶŤſŹ ƵŹŚƃřƽř ŵŹřŵ

ƽŶǀƬĩƱŚĭĥřƹ

ƶŤƃ į ƱŶƃŚƷŹŹŚŤƟŹƽźĭŹŚĪƃźƐųżǀƫŚū

Abstract

The predatory bug, Orius laevigatus (Fibber), and the predatory midge, Aphidoletes aphidimyza Rondani, belong to an aphidophagous guild, feeding on the cotton aphid Aphis gossypii Glover. In this

ŢƟŚƿŹŵ æé é æèîå ƁźƿŸě

æë æç æèîå

(2)

ÐÓ ƱřŹŚƨưƷƹƽźǀƘƯƝřźƇ

ƱƹŹŵįźĭŹŚĪƃźŧř

ƶŤſŹ įŚƷŹŚŤƟŹįƹŹįř

study some avoidance behavior of the predatory midge such as dropping from the plant and choosing an oviposition site in the face of interaguild (IG) predator O. laevigatus are investigated. The results indicate that A. aphidimyza females prefer not to lay eggs when they discover that their offspring would be at risk of being attacked by the IG predator. The experiment also shows that the dropping behavior of predatory midge larva, upon being spotted by the predatory bug (44.5%), is significantly higher than the control (6.5%). Both the dropping behavior and mortality of IG prey are found to be influenced by all developmental stages of the cotton aphid. The percentage of mortality and dropping rates of A.

aphidimyza larvae in the presence of 2nd instar nymphs of aphids were 10% and 12.5% and for 4th instar nymphs were 31% and 44.5%, respectively. These findings confirm the correlation between habitat choice of A. aphidimyza, in relation to predation risk for its offspring, and also underscore the population structure of prey for avoidance behavior strategy of the aphidophagous midge in an interaguild predation system.

Key words: cotton aphid, predation risk, dropping behavior

ƶƯŶƤƯ Źƺƌů ƱřźĭŹŚĪƃ

ƞƬŤŴƯ ƶŤſŹĨƿƶŝƢƬƘŤƯ

(guild)

ŢƀƿŻŹŵ ħźŤƄƯƾƳřƺų

ƲĪưƯ

Ţſř ŦƗŚŝ řƿ ŵŚŬ ƮƷźŝ Ƃƴĩ ŚƷ ƽ ƽŵŶƘŤƯ ŝǀ Ʋ Ʊō ŚƷ ŦůŚŞƯ Źŵ ƕƺƋƺƯ Ʋƿř Żř ƶĩ ŵƺƃ

ƭƺŝ ƱřƺƴƗ Śŝ ƾſŚƴƃ

źĭŹŚĪƃ ƽ ƱƹŹŵ ƶŤſŹ ř ƽ

(intraguild)

ƾƯ ŵŚƿ ŵƺƃ

(Lucas et al., 1998)

źĭŹŚĪƃ ƽ ƱƹŹŵ ƶŤſŹ ř ƽ ƗƺƳ ƾ ƶƐŝřŹ į źĭŹŚĪƃ ƽ Ţſř ƶĩ ŚƷźĭŹŚĪƃ Śŝ

ŵƺūƹ ŸƜţ ƿƶ Żř ƖŞƴƯ

ħźŤƄƯ ŹŚĪƃ Żř żǀƳźĮƿŶĪƿ

ŸƜţ ƿƶ ƾƯ Ŷƴƴĩ

(Polis et al., 1989)

Ʋſ źĭŹŚĪƃ

Orius laevigatus (Fibber)

ƿ Ī ƾ Żř ƮƸƯ źţ ƿ Ʋ ƳřźĭŹŚĪƃ ƾ Ţſř Żř ƶĩ

ƶƳƺĭ ưƷšŚƟōŻřƽŵŶƘŤƯƽŚƷ ƱƺĤ

źţ ƿ žĜ Ʒ Ś ƹ ƶŤƃ ŚƷ ƶƿŸƜţ Ư ƾ Ŷƴĩ řƿ Ʋ ĨƿƲſ Īƃ

źĭŹŚ

ƯƺưƗ ƾ Ţſř ƾƷŚĭƹ ŸƜţ ƿƶ į ƵƹLjƗƱřźĭŹŚĪƃźƿŚſŻřƱō

ƵŚǀĭšřźƄůźŝ

ƁŹřżĭżǀƳŹřƺų

ŢſřƵŶƃ

(Christensen et al., 2002; Tommasini et al., 2002; Van Lenteren et al., 2003;

Jakobsen et al., 2004)

ƶƄě į ƶŤƃ Źřƺų

Aphidoletes aphidimyza Rondani

żǀƳ źĭŹŚĪƃ ƽ

Ťųř ƇŚƈ ƾ Ţſř ƎƤƟ ƶĩ Źŵ ƶƬůźƯ į ƹŹLJ ƽ ƶƳƺĭ Żř Ʒ ƶŤƃ Żř ƾƠƬŤŴƯ ƽŚ

ƾƯ ƶƿŸƜţ ŚƷ

Ŷƴĩ

(Lucas et al., 1998; Lucas & Brodeur, 2001)

ƾţŚƘƫŚƐƯƹƁŹřżĭƱřźƿřŻřźĭŹŚĪƃƶƳƺĭƹŵźƷ

ŢƀƿŻ ƽƹŹ

Ʊō ƁŹƹźě ƹ ƾſŚƴƃ

Ţſř ƵŶƃ ƭŚŬƳř żǀƳ ŚƷ

(Labbafi, 1995; Madadi, 1999;

Erfan & Ostovan, 2004)

ƁŹřżĭ ƾƯƱŚƄƳƶĩŵŹřŵŵƺūƹƽŵŶƘŤƯƽŚƷ ŵ

ŶƷ ƹŹLJ ƶƄě ƽŚƷ ƶŤƃ Źřƺų ľLJƺưƘƯ Ǝſƺţ

Śſ ƿź ŚƷźĭŹŚĪƃ

ŶƴƳŚƯ

Orius spp.

ƹ

Chrysoperla vufilabris (Burmeister)

ŵŹƺƯ ƶƬưů Źřźƣ

Ư ƾ ĭǀ źƳ Ŷ

(Lucas & Brodeur, 2001; Christensen et al., 2002)

ŚƯř ŵƺūƹ ƾƿŚƷŹŚŤƟŹ

ŝŚƴŤūř ƾ



ŶƴƳŚƯ ƂƷŚĩ

ħźŰţ ƠŴƯ ƾ ƱŶƃ Źŵ Ż ƿź ŵŚƀūř ƶŤƃ ŚƷ ƹ Źřźƣ ƲŤƟźĭ Źŵ żĩźƯ ƴƬĩ ƾ ƶŤƃ ŚƷ



(3)

ƶƯŚƳ ƵźƄůƲưŬƳřƽ

ƱřźƿřƾſŚƴƃ èç

ç

 ÎÐîæ

ÐÔ

ƱřźĭŹŚĪƃ źƿŚſ Śŝ ƶūřƺƯ ƂƷŚĩ ƽřźŝ

ƶƄě Ʋƿř ƽŚƷƹŹLJ Źŵ ƵŶƷŚƄƯ

ƵŶƃ Ţſř

(Lucas & Brodeur, 2001)

ŚƷŹ ƾƿ ŻřśŚţźěƹ ƪŰƯ

ŸƜţ ƿƶ ƭŚĮƴƷŹŵƶĩŢſřƾŝŚƴŤūřƽŚƷŹŚŤƟŹƽŚƷŵƺưƳŻřƾĪƿ

Ɛų ŹŚĪƃ Żř ƾƃŚƳ ź

ƾƯ ƵŶƿŵ šřźƄů Żř ƾųźŝ Źŵ ƱŶƃ

ŵƺƃ ƶŝ ƱřƺƴƗ ƩŚŨƯ

 ŹLJ ƹ ƽŚƷ

ħŻƹŶƄƠĩ

ŢƠƷ ƶƐƤƳ ƽř



Coccinella septempunctata L.

 řźŝ ƽ ŹřźƟ Żř ŹŚĪƃźƐų

ƎſƺţƱŶƃ

źƿŚſ

ƱřźĭŹŚĪƃ

ŵƺų řŹ Żř ŮƐſ īźŝ ƶŝ ŗŚě ǀ Ʋ ŚƷŹ Ư ƾ Ŷƴƴĩ

(Sato et al., 2003)

ŵƺūƹ ƴģ ǀ Ʋ

ƳŚĪƯ ǀ ưƀ ƾ

 ƶŝ Ħƿƹ Ƶ Źŵ ţřźƄů ƾ ƶĩ ħźŰţ ưĩ ƾ ŶƳŹřŵ

ř ƿ Ʋ ƩŚưŤůř řŹ ƶŝ ŵƺūƹ Ư ƾ ŵŹƹō ƶĩ ƹŹLJ

ƶƄě į ƶŤƃ Źřƺų

A. aphidimyza

Ƴǀ ż ŝ ŶƳřƺŤ Żř řƿ Ʋ ƳŚĪƯ ǀ Ʈƀ ƶŝ ƱřƺƴƗ ƿ Ĩ ŹŚŤƟŹ ƾŝŚƴŤūř řźŝ ƽ

ŚƷŹ ƾƿ Żř ƶūřƺƯ ƳřźĭŹŚĪƃ ƲſŶƴƳŚƯƾ

O. laevigatus

ƵŵŚƠŤſř Ŷƴĩ ŻřƾĪƿƶǀƋźƟƲƿřƾſŹźŝƶĩ

ƷƹĦěƲƿřƝřŶƷř ƾƯƂ

ŶƃŚŝ ƲǀƴĤưƷ

 ƶĩŢſřƵŶƃƵŵřŵƱŚƄƳ ƵŵŚƯ

ŚƷ ƽ ƚƫŚŝ řƿ Ʋ ƶƄě ƮŴţ ŚƷ ƽ

ŵƺų řŹ ƶŝ šŹƺƇ ƽŵřźƠƳř ƿŚ Źŵ ƶŤſŵ ŚƷ ƽ Ĩģƺĩ Źŵ ƾĪƿŵżƳ ƴƬĩ ƾ ƶŤƃ ŚƷ Źřźƣ Ư ƾ ŶƴƷŵ

(Lucas et al., 1998)

ŝ ƢǀƤŰţ Ʋƿř Źŵ ƶĩ ƾţLJřƺſ Żř źĮƿŵ ƾĪƿ ƶ

Ʊō ƽřźŝ ƾŴſŚě ƩŚŞƳŵ

ƾƯ Śƿō ƶĩ Ţſř Ʋƿř ƮǀƃŚŝ ƌů

Źƺ źƿŚſ źĭŹŚĪƃ Ʊř ƶƄě ŢǀƫŚƘƟ ƱŚĪƯ Źŵ

į źĭŹŚĪƃ

A. aphidimyza

Ư ƾ ŶƳřƺţ Źŵ śŚŴŤƳř ƱŚĪƯ ƱřżǀƯƹ ƮŴţ ŹřŸĭ ƽ ƶŝƵŵŚƯšřźƄů

ŹŚŤƟŹĨƿƱřƺƴƗ

ƶƳŚƯŚſ Ĩƿ Źŵ ƾŝŚƴŤūř

į ƱƹŹŵ ƶŤſŹ ƶƄě ƲǀŝŚƯ ƽř

į ƶŤƃ ƶŤƃ ƹ źĭŹŚĪƃ Ʋſ Źřƺų

į żǀƫŚū

Aphis gossypii Glover

ţ Ő źǀŧ ŹřŸĭ ŶƃŚŝ ƴģźƷ Ÿ ƾƠƬŤŴƯ šŚƘƫŚƐƯ Ʊřźƿř Źŵ Ŷ ƽƹŹ

ƽźĭŹŚĪƃ

ƱƹŹŵ ƶŤſŹ ƵŶƃ ƭŚŬƳř ƽř

(Madadi et al., 2008; Hosseini et al., 2010; Tavoosi, 2011)

ƾƫƹ

ƱƹŹŵŹŚĪƃƾŝŚƴŤūřƽŚƷŹŚŤƟŹƽƹŹƵŶƿŶěƲƿřšřźŧř

ƶŤſŹ ƶŝŚţƽř ƲƿřƹŢſřƵŶƄƳƾſŹźŝƩŚů

ƲǀŤƀŴƳƽřźŝƂƷƹĦě

ƾƯƕƺƋƺƯƲƿřƶŝŹƺƄĩŹŵŹŚŝ

ŵŻřŵźě

ŵřƺƯ ƹ ƁƹŹ ŚƷ

ƁŹƹźě ƴƬĩ ƾ šřźƄů

ƴƬĩ ƾ ƶŤƃ į ƶŝżǀƫŚū ŶưŰƯƕŹřżƯŻřźĭŹŚĪƃƹŵźƷħźŤƄƯŹŚĪƃƱřƺƴƗ

ƖưūũźĩŵŚŝō

ƽŹƹō

ƾƿŚſŚƴƃŻřžěƹ

 ƹŹ ƽ ĭǀ ƵŚ ų ǀ ŹŚ ƲǀĮƳƮƣŹ ƁŹƹźě Ƶŵřŵ Ŷƃ ƶƄě į ƶŤƃ Źřƺų

A. aphidimyza

ƶŤƃƽƹŹ į ƹŹƺĩŸƯ Ʋſ źĭŹŚĪƃ

O. laevigatus

Ƴǀ ż ƢŝŚƐƯ Śŝ ƁƹŹ

Venzon et al. (2002)

ƁŹƹźě

(4)

ÐÕ ƱřŹŚƨưƷƹƽźǀƘƯƝřźƇ

ƱƹŹŵįźĭŹŚĪƃźŧř

ƶŤſŹ įŚƷŹŚŤƟŹįƹŹįř

ŶƳŶƃƵŵřŵ

šźěƺĩŢĩźƃŻřźĭŹŚĪƃƹŵźƷ

(Koppert)

ŶƃƶǀƸţŶƴƬƷ

ƶǀƫƹřŢǀƘưū

į ƶƄě į

źĭŹŚĪƃ æååå źĭŹŚĪƃƲſƹƵźǀƠƃ Òåå

ƵźƄů į ŵƺŝƪƯŚĩ ƁŹƹźě ƴƬĩ ƾ ƽŚƷ šřźƄů źĭŹŚĪƃ

ƹ ŹŚĪƃ Źŵ ŚƯŵ ƽ æ t çÒ ƶūŹŵ į ƀƬſ ǀ Žƺ ƹ ŢŝƺƏŹ æå t Òå ƹ śƹŚƴţ ŹƺƳ ƽ æÓ ŢƗŚſ

ŚƴƃƹŹ ƾƿ ƹ í ŢƗŚſ ŹŚţ ƿĪ ƾ ƭŚŬƳř Ŷƃ

ƂƿŚƯŻō Ʃƹř śŚŴŤƳř ƪŰƯ ƮŴţ ŹřŸĭ ƽ Ǝſƺţ ƶƄě ŚƷ ƽ ƶŤƃ ƹŹŹŵŹřƺų

Źŵ ƲſŚŝƾƿƹŹ

O. laevigatus

Ʋƿř ŚƯŻō ƿ Ƃ Źŵ ŚƷŶůřƹ ƽ ƶŤųŚſ ƵŶƃ Żř ƢƬţ ƶŝ ŵŚƘŝř Ñå

„ Ñå

„ Ñå ŤƳŚſ ƾ źŤƯ ƭŚŬƳř Ŷƃ

Źŵ ŵ ƿ ƵŹřƺ į ŹŚƴĩ ƽ řƿ Ʋ ŚƷŶůřƹ ſ ųřŹƺ ƾ ƶŝ źƐƣ æå ŤƳŚſ ƾ źŤƯ řƿ ŵŚŬ ƹ Ǝſƺţ Źƺţ ƽ æÒå ƂƯ

ƵŶƳŚƃƺě Ŷƃ źƷ ƿ Ĩ Żř ŚƷŶůřƹ ƽřŹřŵ ƹŵ ŵŶƗ ĭ ǀ ƵŚ ƹŵ ĭźŝ ƾ ų ǀ ŹŚ ƶŝ ƾĪƿ ƹ ŶƷŚƃ ƱřƺƴƗ

ƶŝƽźĮƿŵ ŹŚưǀţƱřƺƴƗ

ŵƺŝ Źŵ ƶĤƷŚǀĭźƷƲƿźƿŻŢưƀƣ

 ƟźƓ ƾ ƽƹŚů śō ƹ ƿ Ĩ ŚƯ ƿƖ ƺƃ ƿ ƵŶƴ

ŶƃƵŵřŵŹřźƣ Śţ

Żř ŹřźƟ šřźƄů ĭƺƬū ǀź ƽ ŵƺƃ Ƙŝ Ŷ Żř ƵŵŚƯō ƱŶƃ Ŷůřƹ ƽŚƷ ŚƯŻō ƿ Ƅ ƾ

 ƿ Ĩ

ŢƠū ƶƄě į ƚƫŚŝ

A. aphidimyza

źƷŹŵ Ŷůřƹ ŚƯŻō ƿ Ƅ ƾ ŚƷŹ Ŷƃ ř ƿ Ʋ ŚƯŻō ƿ Ƃ Źŵ ŜƫŚƣ ŭźƏ

ľLjƯŚĩ ƟŵŚƈţ ƾ źƿŻŜǀĩźţŚŝ ƹ

Źŵ šŶƯ ìç ŢƗŚſ Źŵ Òå ŹřźĪţ ƭŚŬƳř ƹ

Źŵ ƂƿŚƯŻōƱŚƿŚě

ŵřŶƘţ ƮŴţ ŚƷ ƽ ƶŤƃřŸĭ ƵŶƃ Ǝſƺţ ƶƄě ŚƷ ƽ ƚƫŚŝ ǀţƹŶƷŚƃƵŚǀĭƽƹŹ ŹŚư

ƁŹŚưƃ Ŷƃ

ĭ ǀ ƵŚ ţǀ ŹŚư ŵřŶƘţ Óå ŵŶƗ ƵŹƺě į Ʋſ ƭŹŚƸģ ƶŤƃ į

A. gossypii

ƿ Ĩ ŵŶƗ ƵŵŚƯ į ƚƫŚŝ

ŢƠū ĭ ǀź ƽ ƵŵźĪƳ ƽ ƶſ ƵŻƹŹ į

O. laevigatus

ĭǀ ƵŚ ŶƷŚƃ ŵřŶƘţ Óå ŵŶƗ ƵŹƺě į Ʋſ ƭŹŚƸģ ƶŤƃ į

A. gossypii

ƭƹŵƂƿŚƯŻō

ƾſŹźŝ ŚƷŹ ƱŶƃ Żř ŮƐſ īźŝ Ǝſƺţ ŚƷƹŹLJ ƽ ƶƄě į ƃ Ťƶ Źřƺų ƶŝ ŹŚŤƟŹĨƿƱřƺƴƗ

ƱōƾƿřŹŚĩƾŝŚƿŻŹřƹƾŝŚƴŤūř

řźŝ ƽ ƭŚŬƳř řƿ Ʋ ŚƯŻō ƿ Ƃ Żř ĭ ǀ ƷŚ ƶĤ ƽŚƷ ų ǀ ŹŚ Źŵ ƶƬůźƯ į ƹŵ ĭźŝ ƾ ƵŵŚƠŤſř Ŷƃ ƶĩ ŵ ƪųř

ƶƫƺƫ ƽŚƷ Īģƺĩ ƾ ƶŝ źƐƣ Ò ƹ ƕŚƠţŹř Õ ƾŤƳŚſ źŤƯ Źřźƣ Ƶŵřŵ Ŷƃ ŶƳŵƺŝƵ ŶůřƹƲƿř ƾƄƿŚƯŻōƽŚƷ

żǀƳ ƱƹŹŵ źƓ ƹ Ɲ īŹżŝ źţ ŵ ƿ źĮ ƽ ƶŝ źƐƣ ÎÔ ƾŤƳŚſ źŤƯ ƹ ƹŚů ƽ śō Źřźƣ ŤƟźĭ ƶŝŶƴ ƽŹƺƏ ƶĩ

ƶƄěƽŚƷƹŹLJ

į ƶŤƃ ŚƷŹŹŚŤƟŹŻƹźŝŻřžěŹřƺų

ƱŶƃ

 ƁŹŚưƃƪŝŚƣƹƵŵŚŤƟřƝƹźƓƲƿřƪųřŵ

ŶƴƃŚŝ ƿ Ĩ Ƃƃƺě ƤƬţ ƾ ƶƳřƺŤſř ř ƽ ƶŝ ƕŚƠţŹř ÏÒ ƹ źƐƣ ÏÍ ƾŤƳŚſ źŤƯ żǀƳ ƽƹŹ ƶƗƺưŬƯ į ơƺƟ

(5)

ƶƯŚƳ ƵźƄůƲưŬƳřƽ

ƱřźƿřƾſŚƴƃ èç

ç

 ÎÐîæ

ÐÖ

ƶŤƃřŸĭ Ŷƃ ƪŞƣ Żř ƕƹźƃ ŚƯŻō ƿ Ƃ ŵřŶƘţ Òå Ťƃ ƶ ƶŝ ſƹ ǀ ƶƬ į ƿ Ĩ ƮƬƣ Ư ƺ ƽ ƀŝ ǀ ŹŚ ŹŚŝ ƿ Ĩ ƹŹ ƽ

ĭǀ ƵŚ ƪƤŤƴƯ Ŷƃ ƹ ƶŝƱƺƯŻōŵŹƺƯƽŚƷŹŚưǀţŜǀĩźţ

ŹřźƣƶƀƿŚƤƯŵŹƺƯŶƷŚƃƶŝŢŞƀƳźƿŻŭźƃ

ŢƟźĭ

ţǀ ŹŚư æ ŵřŶƘţ Òå ŵŶƗ ƵŹƺě į Ʋſ ƭŹŚƸģ ƶŤƃ ƽ

A. gossypii

æå ŵŶƗ ƹŹLJ ƶƄě į ƶŤƃ Źřƺų

A. aphidimyza

ƿ Ĩ ŵŶƗ Ʋſ źĭŹŚĪƃ

O. laevigatus

ŶƷŚƃ æ ŵřŶƘţ Òå ŵŶƗ ƵŹƺě į Ʋſ ƭŹŚƸģ ƶŤƃ ƽ

A. gossypii

æå ŵŶƗ ƹŹLJ ƶƄě į ƶŤƃ Źřƺų

A. aphidimyza

ţǀ ŹŚư ç ŵřŶƘţ Òå ŵŶƗ ƵŹƺě į Ʋſ ƹŵ ƭ ƶŤƃ ƽ

A. gossypii

æå ŵŶƗ ƹŹLJ ƶƄě į ƶŤƃ Źřƺų

A. aphidimyza

ƿ Ĩ ŵŶƗ Ʋſ źĭŹŚĪƃ

O. laevigatus

ŶƷŚƃ ç řŶƘţ ŵ Òå ŵŶƗ ƵŹƺě į Ʋſ ƭƹŵ ƶŤƃ ƽ

A. gossypii

æå ŵŶƗ ƹŹLJ ƶƄě į ƶŤƃ Źřƺų

A. aphidimyza

ŚƯŻō ƿ Ƃ Śŝ ŚƷŹ ŻŚſ ƽ Ʋſ ŚƷ ƽ ƵŵŚƯ į ƚƫŚŝ ƕƹźƃ Ŷƃ Ʋſ ŚƷ ƽ ƚƫŚŝ ƮƷ

Ʋſ çÑ ŢƗŚſ ƪŞƣ

Żř ƕƹźƃ ŚƯŻō ƿ Ƃ ƹ Źŵ ƝƹźƓ ƁŹƹźě ƵřźưƷ Śŝ ŚƷŸƛŚĩ ƽ ģ ǀ Ʋ Źřŵ Įƴſźĭ ƾ Ƶŵřŵ ƵŶƃ ŶƳŵƺŝ

ŚƯŻō ƿ Ƃ Źŵ çå ŹřźĪţ ƭŚŬƳř Ŷƃ ƹ ŶƘŝ Żř ŢƃŸĭ çÑ ŢƗŚſ ŵřŶƘţ ŚƷƹŹLJ ƾƿ ƶĩ Żř ƹŹ ƽ īźŝ ŚƷ

ŚƷŹ

ŶƳŶƃ ƹ ŵřŶƘţ ŚƷƹŹLJ ƾƿ ƶĩ ƹŹ ƽ īźŝ ŚƷ ƵŶƳŻ ƵŶƳŚƯ ƹ źĭŹŚĪƃƲſƎſƺţŚƿ

ƶŤƄĩ ƵŶƃ

ŶƳŵƺŝ ŢŞŧ Ŷƃ ƲǀƴĤưƷ

 ƾŝŚƴŤūřŹŚŤƟŹƲƿřƾƿřŹŚĩƾŝŚƿŻŹřƹƶƀƿŚƤƯƽřźŝ

īźƯƱřżǀƯŹŵ

źǀƯƹ

ƶƄě ƽŚƷ

A. aphidimyza

ŚƷŹŹŚŤƟŹƝŸůŚŝ

ƾƗƺƴƈƯƎƿřźƃŹŵƾƸŝŚƄƯƂƿŚƯŻōŚƷƹŹLJƱŶƃ

ŶƃƭŚŬƳřƽźŤěƝźƓƪųřŵƹ

ƲƿŶŝ ŹƺƔƴƯ ŵƿ Ĩƀ ŚƷ ƽ ĭźŝ ƾ ų ǀ ŹŚ ƶŝ šŹƺƇ Ʊƺĭĥřƹ ƪųřŵ

źŤě ƽ ŵ ƿ Ƃ ŚƷ ƽ æå ŤƳŚſ ƾ źŤƯ ƽ ƹŹ ƽ śō ŹŚĭō æ Źřźƣ Ƶŵřŵ Ŷƃ ƶŝƱƺƯŻōŵŹƺƯƽŚƷŹŚưǀţƹŶƳ

źƿŻ ŭźƃ

 ƶĤƷŚǀĭ ƽƹŹ ƮƷ

ŹŚǀų ƽŚƷ ŝ

ľLjŞƣ ƶĩ ƾůźƃ Ś

ŢƃŸĭ Ʊō źĩŷ ƪųřŵ ƮƷ ƹ

ƽźŤě Ƃƿŵ

 ŶƳŶƃƶƀƿŚƤƯƮƷŚŝ

ţǀ ŹŚư æ ŵřŶƘţ çå ŵŶƗ ƹŹLJ Ʋſ ƭƺſ ƮƷ Ʋſ ƶƄě į

A. aphidimyza

Ñå ŵŶƗ ƵŹƺě į Ʋſ

ƭŹŚƸģ ƶŤƃ į

A. gossypii

ƿ Ĩ ŵŶƗ ƵŵŚƯ į ƚƫŚŝ Ʋſ źĭŹŚĪƃ

O. laevigatus

ţǀ ŹŚư ç ŵřŶƘţ çå ŵŶƗ ƹŹLJ Ʋſ ƭƺſ Ʒ Ʈ Ʋſ ƶƄě į

A. aphidimyza

Ñå ŵŶƗ ƵŹƺě į Ʋſ

ƭƹŵ ƶŤƃ į

A. gossypii

ƿ Ĩ ŵŶƗ ƵŵŚƯ į ƚƫŚŝ Ʋſ źĭŹŚĪƃ

O. laevigatus

(6)

ÑÍ ƱřŹŚƨưƷƹƽźǀƘƯƝřźƇ

ƱƹŹŵįźĭŹŚĪƃźŧř

ƶŤſŹ įŚƷŹŚŤƟŹįƹŹįř

ŹŵżǀƳƂƿŚƯŻōƲƿř çå

ƹŶƃƭŚŬƳřŹřźĪţ ŶƘŝ

Żř ŢƃŸĭ çÑ ŢƗŚſ ŵřŶƘţ ŚƷƹŹLJ ƽ ƶŤƄĩ ƵŶƃ į

ƶƄě į ƶŤƃ ĨƀƿŵƽƹŹŹřƺų

ƶĤƷŚǀĭƹƾĭźŝƽŚƷ

ŚƷ ƁŹŚưƃ ƶƀƿŚƤƯƹ ƃ Ŷ

żŬţ ƿƶ ƪǀƬŰţƹ ŹŚƯō ƽ

řźŝ ƽ żŬţ ƿƶ ƹ ƬŰţ ǀ ƪ ƝLjŤųř ŝǀ Ʋ ŵřŶƘţ ƮŴţ ŚƷ ƽ ƶŤƃřŸĭ ƵŶƃ Ǝſƺţ ƶƄě ŚƷ ƽ ƚƫŚŝ ƹ

řŹŚĩ ƾƿ ƳŚĪƯ ǀ Ʈƀ ŚƷŹ ƱŶƃ Żř ŮƐſ īźŝ Źŵ ƵŶƳŻ ƳŚƯ ƾ

ŚƷƹŹLJ Żř ŚƤƯ ƿ ƶƀ į Ư ǀ ĮƳŚ ǀ Ʋ ŚƷ ƶŝ Ĩưĩ

ƱƺƯŻō t

ƵŵŚƠŤſř Ŷƃ ƭƹŵƂƿŚƯŻōŹŵ řźŝ

ƽ żŬţ ƿƶ ƹ ƬŰţ ǀ ƪ ƝLjŤųř ŝǀ Ʋ ŶƇŹŵ ŹLJ ŚƷƹ ƽ ƵŵŚŤƟř

Żř

īźŝ ƵŶƳŻ ƵŶƳŚƯ ƹŹ ƽ īźŝ ƹ ŚƷƹŹLJ ƽ ƶŤƄĩ ƵŶƃ Żř ƱƺƯŻō ç

„ ç ƶƿŚěźŝƪƿŹƺŤĩŚƟ

į ŭźƏ

ľLjƯŚĩ źĭŹŚĪƃŮƐſƹŵŚŝƾƟŵŚƈţ

ŹƺƌůƭŶƗƹŹƺƌů

ƶŤƃƞƬŤŴƯƲǀƴſŻřŮƐſƹŵƹ

ƵŹƺě į ƭŹŚƸģƹƭƹŵƲſ

ŶƃƵŵŚƠŤſř

ƲǀĮƳŚǀƯ źǀƜŤƯ ƱƺƯŻōŚŝżǀƳƶŤƀŝřƹƽŚƷ

Ʈĩ źţ ƿ Ʋ ř ƝLjŤų

ƴƘƯ ƾ Źřŵ LSD

ŶƳŶƃƶƀƿŚƤƯ

ƵŵřŵƪǀƬŰţƹƶƿżŬţ

ŚŝŚƷ ƭźƳ ŹřżƟř

SAS 8.1

ŚƷŹřŵƺưƳƹŶƃƭŚŬƳř

ŹřżƟřƭźƳŚŝ

Microsoft Excel 2007

ŶƳŶƿŵźĭƮſŹ

ŚŤƳ ƿ Ū ŦŰŝƹ

żŬţ ƿƶ į ŹŚƯō ƽ Ƶŵřŵ ƩƹřƂƿŚƯŻōƽŚƷ ƱŚƄƳ

ŵřŵ ƶĩ ƶƄě ŚƷ ƽ ƵŵŚƯ ƽ ƚƫŚŝ ƾƳŚƷŚǀĭ ƶĩ Ʋſ

źĭŹŚĪƃ Ʊō ƽƹŹ Źƺƌů

ŵŹřŶƳ ƶŝ ŢŞƀƳ řŹ ƳŚĪƯ

ƾ ƶĩ Ʋſ źĭŹŚĪƃ Źƺƌů ŵŹřŵ ƶŝ ŹƺƏ

ƴƘƯ ƾ Źřŵ ƽ ƮŴţƽřźŝ ƾƯŮǀūźţƽŹřŸĭ

ŶƴƷŵ ÐÐ ÎÎ =

žt

ÖÕ Î,

=

ždf

ÍÍÍÎ Í

P <

ƪĪƃ æ

ƵŵřŵƲƿř ŵřŵƱŚƄƳŚƷ

ƶƧ Źƺƌů Ʋſ

O. laevigatus

ƶŝ ƱřƺƴƗ źĭŹŚĪƃ ƱƹŹŵ ƶŤſŹ ř ƽ Ư ƾ ŶƳřƺţ Źŵ

śŚŴŤƳř ƪŰƯ ƮŴţ ŹřŸĭ ƽ ƶƄě ŚƷ ƽ ŚƯ Ƶŵ į ƶŤƃ Źřƺų

A. aphidimyza

ŧŐţ ǀź ŹřŸĭ ŶƃŚŝ ř ƿ Ʋ ŤƳ ǀ ƶŬ

ƢŝŚƐƯ Śŝ ŚŤƳ ƿ Ū

Ruzicka & Havelka (1998)

Ţſř ƶĩ ƱŚƄƳ ŶƳŵřŵ ƶƄě ŚƷ ƽ

A. aphidimyza

Źŵ

ƬŰƯ ƾ ƶĩ ŚƷƹŹLJ ƽ ƽŹƺŤƫŚŝ

Chrysoperla carnea (Stephens)

ƹ ħŻƹŶƄƠĩ

C. septempunctata

Źƺƌů ŶƳŹřŵ

 Ʈĩ źţ ƮŴţ ŹřŸĭ ƽ Ư ƾ Ŷƴƴĩ ƲǀƴĤưƷ



Griffin & Yeargan (2002)

ƁŹřżĭżǀƳ

ŶƳŵźĩ

ƶĩ ħŻƹŶƄƠĩ

ŚƷ ƽ ƵŵŚƯ į

Colemegilla maculata (De Geer)

ƮŴţ ŚƷ ƽ ŵƺų řŹ Źŵ ƬŰƯ ƾ ƶĩ

Ʈĩ źţ ƿ Ʋ ƩŚưŤůř źĭŹŚĪƃ ƽ ŵƺūƹ ƶŤƃřŵ ŶƃŚŝ Źřźƣ Ư ƾ Ʒŵ ƴŶ ƶƴǀƯŻƲƿřŹŵƶĩƾţLJŚƤƯƶŝƽŹƹźƯ

ŢſřƵŶƃźƄŤƴƯ ƱŚƄƳ

Ư ƾ ŶƷŵ ƶĩ ƂǀŝŹŵ ƺƯźţ

ŵŹř ƱřźĭŹŚĪƃ

ƶŤƃ Źřƺų ŹŵŚƣ ŶƴŤƀƷ Źƺƌů

(7)

ƶƯŚƳ ƵźƄůƲưŬƳřƽ

ƱřźƿřƾſŚƴƃ èç

ç

 ÎÐîæ

ÑÎ

ƱřźĭŹŚĪƃ

ŵƺų řŹ ħŹŵ ƵŵźƧ ƹ Źŵ ƤƏŚƴƯ ƾ ƶĩ ƱřźĭŹŚĪƃ

ƮƷ ƶŤſŹ Źŵ Ʊō Śū Źƺƌů ŻřŶƳŹřŵ

ƮŴţ ŹřŸĭ ƽ ƕŚƴŤƯř Ŷƴƴĩ

(Doumbia et al., 1998; Ruzicka, 1998; Frechette et al., 2008)

ħźţ

ƪŰƯ ŜſŚƴƯ řźŝ ƽ ƮŴţ ŹřŸĭ ƽ ƶƄě ŚƷ ƽ ƚƫŚŝ ƶŝ ƶƐſřƹ į Źƺƌů ƱřźĭŹŚĪƃ

ƮƷ ƶŝ ƶŤſŹ źƔƳ

ƾƯ źěƾŝŚƴŤūřŹŚŤƟŹĨƿŶſŹ

ŶƃŚŝƶƴƿżƷ

 ħźţƶĩřźģ

ƱŚĪƯ ŜſŚƴƯ řźŝ ƽ ƮŴţ ƽŹřŸĭ ƲŤƟŚƿƹ

ŶƿŶūŜſŚƴƯ ƱŚĪƯ Ư

ƾ ŶƳřƺţ ƩLjŤųř ŦƗŚŝ

Źŵ ŢƠū ƿ ŝŚ ƾ ƭŚŬƳřźſ ƹ źĭŹŚĪƃ ƚƫŚŝ šřźƄů

ƂƷŚĩ ŵřŶƘţ ƮŴţ ƽŚƷ řŸĭ ƶŤƃ ƵŶƃ ŵƺƃ

(Havelka & Zemek, 1999)

ƪĪƃ æ ŚƤƯ ƿ ƶƀ į ŶƇŹŵ ƮŴţ ŹřŸĭ ƽ

SE)

t (

ƶƄě ŚƷ ƽ ƚƫŚŝ

A. aphidimyza

Źŵ Źƺƌů ƹ ƭŶƗ

Źƺƌů Ʋſ źĭŹŚĪƃ

O. laevigatus

Ɲƹźů źǀƛ ƶŝŚƄƯ ƲǀĮƳŚǀƯ ŚƷ ƱŚƄƳ ƵŶƴƷŵ į ŵƺūƹ

ƝLjŤųř ƴƘƯ ƾ Źřŵ Ţſř

t test

 åÒ å

á =

Fig. 1. Oviposition percentage (± SE) of the predatory midge females, A. aphidimyza, in the presence and absence of the predatory bug, O. laevigatus. Means with different letters are significantly different (t test, á = 0.05).

ŶƴƳŚƯ ƲǀƤƤŰƯ ƾųźŝ

Lima & Dill (1990)

ŶƳŶƤŤƘƯ ƶĩ źĭř Żř ƾƃŚƳ ƽźĭŹŚĪƃ źƐų

ƶƳƺĭ ƮƷƽŚƷ ŤſŹ ŶƃŚŝƽŶūƹƾưŤůƶ

 ƚƫŚŝŹƺƳŚū Żř

ŹŚŤƟŹ żƷźě ƿ ƶƴ į ƮŴţƭŶƗ ƽřźŝƽŹřŸĭ

ƾƯ ƵŵŚƠŤſř ŵƺų ƽŶƘŝ ƪƀƳ ƾƬƣřŶů ƒƠů

Ŷƴĩ ƶŝ

ƩŚŨƯ ƱřƺƴƗ

ŵŹƺƯ Źŵ ū

ǀ ūź ǀ ħź

Gryllus integer Scudder

ƱŚƄƳ Ƶŵřŵ ƵŶƃ Ţſř ŹƺƳŚūƲƿřƶĩ ŚƸƴţ

ƳŚƯŻ ƾ ƶĩ ŵŚƀūř ŵřźƟř ƮƷ ƶƳƺĭ

Źŵ ŰƯ ǀ Ǝ ƵřźưƷ Śŝ šƺŞĪƴƗ ŚĪƃ źĭŹ

Hololena nedra Chamberlin & Ivie

Źƺƌů ƶŤƃřŵ ŶƴƃŚŝ Żř

Ʊō ƪŰƯ Źƹŵ ƽ Ư ƾ ƴĩ Ŷ ţŹƺƇŹŵ ƾ ƶĩ źĭř šƺŞĪƴƗ ľLjŞƣ Żř ŵřźƟř ƮƷ ƶƳƺĭ į ū ǀ ūź ǀ ħź ŸƜţ ƿƶ ƵŵźĪƳ

(8)

ÑÏ ƱřŹŚƨưƷƹƽźǀƘƯƝřźƇ

ƱƹŹŵįźĭŹŚĪƃźŧř

ƶŤſŹ įŚƷŹŚŤƟŹįƹŹįř

ŶƃŚŝ ƹ ƿŚ ŵŚƀūř ŵřźƟř ƮƷ ƶƳƺĭ į ū ǀ ūź ǀ ħź Źŵ ŰƯ ǀ Ǝ ŵƺūƹ ƶŤƃřŶƳ

ŶƃŚŝ ū ǀ ūź ǀ ħź ơƺƟ Ʒ ǀ Ģ

ŹŚŤƟŹ źĭŹŚĪƃŶƋ ƽ

ŢŞƀƳ ƶŝ Ʊō ŚƄƳ Ʊ ưƳ ƾ ŶƷŵ

(Kortet & Hedrick, 2004)

ƶƿżŬţ Żř ƪƇŚů ŪƿŚŤƳ ƭƹŵ ƂƿŚƯŻō Źŵ

į Ƶŵřŵ žƳŚƿŹřƹ

ŵřŵ ƱŚƄƳ ŚƷ ƶƧ

ƮƷźŝ Ƃƴĩ

ƾƴƘƯ ŮƐſƹŵŹŵźĭŹŚĪƃƲǀŝƽŹřŵ

ŹƺƌůƭŶƗƹŹƺƌů

ƹ Ʋſƹŵ ƶŤƃƾĭŹƺě

ƵŹƺě į Ʋſ

ƭŹŚƸģƹƭƹŵ

ŵŹřŵŵƺūƹ

ƩƹŶū ƽŚƷ æ ƹ ç ƶŝ źĮƿŵšŹŚŞƗ

 ƲſŹƺƌůƭŶƗƹŹƺƌů

ƶƬůźƯƲǀƴĤưƷƹźĭŹŚĪƃ

į ƾƯŹŚĪƃƾƴſ

ŚƷŹŶƇŹŵŶƳřƺţ

ƶƄěƽŚƷƹŹLJƱŶƃ

į ƹźĭŹŚĪƃ

īźƯ ƱōźǀƯƹ ƎſƺţřŹŚƷ

O. laevigatus

ƶŝ ƾƴƘƯŹƺƏ ţŢŰţƽŹřŵ Ő

ŶƷŵŹřźƣźǀŧ

ƩƹŶū Î ƶƿżŬţ į ƶŤƃƾĭŹƺěƞƬŤŴƯƪůřźƯšřźŧřžƳŚƿŹřƹ

ſ ǀƴ ƾĭŹƺěƭƺſƹƭƹŵƲ

ƹ

Źƺƌů źĭŹŚĪƃ Ʋſ

ƮƷźŝ ƹ Ʊō Ƃƴĩ ŚƷŹ ŶƇŹŵ ƽƹŹ ŚƷ

ƶƄě ƽŚƷƹŹLJ ƱŶƃ

į

ŹŚǀųƵŚǀĭƽƹŹŻřźĭŹŚĪƃ

Table 1. Analysis of variance results for the effects of different aphid nymph stages (2nd and 4th instar nymphs) and the predatory bug presence and their interaction on the percentage of the predatory midge larvae dropping off from cucumber plant.

P F

MSE df

Source of variation

0.00 19.07

0.72 1

Aphid

0.00 42.65

1.60 1

Predator

0.00 20.48

0.77 1

Aphid × Predator

- -

0.04 76

Error

ƩƹŶū Ï ƶƿżŬţ į ƪůřźƯšřźŧřžƳŚƿŹřƹ

ƞƬŤŴƯ ƶŤƃƾĭŹƺě

ſ ǀƴ ƾĭŹƺěƭƺſƹƭƹŵƲ

ƹ

źĭŹŚĪƃ Ʋſ Źƺƌů

ƮƷźŝ ƹ Ʊō Ƃƴĩ īźƯ ŶƇŹŵ ƽƹŹ ŚƷ

ƶƄě ƽŚƷƹŹLJ źǀƯƹ

į

źĭŹŚĪƃ

Table 2. Analysis of variance results for the effects of different aphid nymph stages (2nd and 4th instar nymphs) and the predatory bug presence and their interaction on the percentage of the predatory midge larvae mortality.

P F

MSE df

Source of variation

0.00 101.71

2.02 1

Aphid

0.00 16.72

0.33 1

Predator

0.00 19.95

0.39 1

Aphid × Predator

- -

0.20 76

Error

(9)

ƶƯŚƳ ƵźƄůƲưŬƳřƽ

ƱřźƿřƾſŚƴƃ èç

ç

 ÎÐîæ

ÑÐ

ŚƤƯ ƿ ƶƀ į Ư ǀ ĮƳŚ ǀ Ʋ Ƶŵřŵ ŚƷ ƽ ů ƞƬŤŴƯƽŚƷŹŚưǀţŻřƪƇŚ ƱŚƄƳ

ŵřŵ ƶƧ Źƺƌů Ʋſ źĭŹŚĪƃ

ƽƹŹ Ư ǀ Ʊřż ŚƷŹ ƱŶƃ ŚƷƹŹLJ ƶƄěƽ į īźŝŻřźĭŹŚĪƃ

ƶŝŹŚǀųƵŚǀĭƽŚƷ

ƮƀǀƳŚĪƯĨƿƱřƺƴƗ

ƾŝŚƴŤūř ŧŐţ ǀź ƴƘƯ ƾ Źřŵ ƽ ƶŤƃřŵ Ţſř ÒÔ ÑÑ = F(3, 77)

 ÍÍÎ Í

P <

ƩƹŶū è ƴĤưƷ ǀ Ʋ

 ŚŤƳ ƿ Ū

ƱŚƄƳ ŵřŵ ƶĩ ŚƷƹŹLJ ƶƄěƽ į ƶŤƃ Źřƺų Źŵ Źƺƌů ƵŹƺě ŚƷ ƽ Ʋſ ƭŹŚƸģ ƶŤƃ ŝǀ Ƃ źţ Żř ƵŹƺě ŚƷ ƽ

Ʋſ ƭƹŵ

 ŵƺų řŹ Żř ŮƐſ īźŝ ŚƷŹ ŶƳŵźĩ ÒÔ ÑÑ = F(3, 77)

 ÍÍÍÎ Í

P <

ƩƹŶū è ƲƿřŹŵ

ƂƿŚƯŻō

 ƵŶƳŻ ƾƳŚƯ ŚƷƹŹLJ ƽ

A. aphidimyza

ƹŹ ƽ īźŝ ŚƷ ŹŚǀųƵŚǀĭƽ

Źŵ Źƺƌů źƷ ƿ Ĩ Żř Ʋǀƴſ

ƭƹŵ ƹ ƭŹŚƸģ ƾĭŹƺě ƶŤƃ ƝLjŤųřƮƷŚŝżǀƳ

ƴƘƯ ƾ Źřŵ ƽ Ţƃřŵ ÐÓ ÕÒ = F(3, 77)

 ÍÍÍÎ Í

P <

īźƯ ƹ ƶƄě ƽŚƷƹŹLJ źǀƯƹ

į ƶŤƃ źĭŹŚĪƃ Ǝſƺţ Źřƺų

O. laevigatus

ƭŹŚƸģ Ʋſ Źƺƌů Śŝ

ƶŤƃ ƾĭŹƺě

į ŵƹŶů żǀƫŚū

ƵŹƺě ƶŝ ŢŞƀƳ źŝřźŝ ƶſ

į ŵźĩ řŶǀě ƂƿřżƟř ƶŤƃ ƭƹŵ Ʋſ

ÖÎ ÑÓ = F(3, 77)

 ÍÍÍÎ Í

P <

ƩƹŶū æ

ƩƹŶū Ð Ưǀ ĮƳŚ ǀ Ʋ Źŵ ŶƇ

SE)

t (

ŚƷƹŹLJ ƽ ƶƄě į ƶŤƃ řƺų Ź ŚƷŹ ƵŶƃ Żř ŮƐſ

īźŝ ƶŤƄĩ ƵŶƃ

ƹŹ ƽ īźŝ ƹ ƵŶƳŻ ƵŶƳŚƯ ƹŹ ƽ

īźŝ źĭŹŚĪƃƲſŹƺƌůŹŵ

Table 3. Average percentage (± SE) of the dropping predatory midge larvae, in the presence of the predatory bug from the plant, and being lost to predation or present on the plant.

T2 C2

T1 C1

Response

12.5 ± 3.8 b 7.1 ± 1.9 b

44.5 ± 10.1 a 6.5 ± 1.1 b

Dropped off

10 ± 1.4 b 1.2 ± 0.3 c

31 ± 7.2 a 1 ± 0.4 c

Lost to predation

77.5 ± 4.7 b 91.7 ± 4.5 a

24.5 ± 2.7 c 92.5 ± 5 a

Present on leaf

C1, the predatory midge larvae + 4th instar nymphs of aphid.

C2, the predatory midge larvae + 2nd instar nymphs of aphid.

T1, the predatory midge larvae + 4th instar nymphs of aphid + the predatory bug.

T2, the predatory midge larvae + 2nd instar nymphs of aphid + the predatory bug.

Means followed by the same letter in each row are not significantly different (LSD test, á = 0.05).

ƶƀƿŚƤƯ į ŚŤƳ ƿ Ū ƪƇŚů Żř ƭŚŬƳřƂƿŚƯŻō

ĨƀƿŵƽƹŹƶŤƟźĭ

żǀƳƪƯŚĩƵŚǀĭƹƾĭźŝƽŚƷ

ƾƯ ƱŚƄƳ ƶĩ ŶƷŵ Óå ŚƷƹŹLJ ƽ ƶƄě į ƶŤƃ Źřƺų Źŵ Źƺƌů źƯ ƶƬů į ƭŹŚƸģ ĭŹƺě ƾ ƶŤƃ į

A. gossypii

ĨƀƿŵƽƹŹ

ƾĭźŝƽŚƷ Ǝſƺţ

Ʋſ źĭŹŚĪƃ ƶŤƄĩ ŶƳŶƃ Źŵ ƫŚů ƾ ƶĩ řƿ Ʋ Ư ǀ Ʊřż Źŵ

ƭŚŬƳřƂƿŚƯŻō

ƵŚǀĭƽƹŹƵŶƃ

ŵƹŶů çí ŵƺŝ ÒÕ Ô =

t

ÐÕ =

df

ÍÍÍÎ Í

P <

ƪĪƃ ç

ŶƴģźƷ ƶĩ ŵřŶƘţ ŚƷƹŹLJ ƽ ƶŤƄĩ ƵŶƃ Źŵ Źƺƌů ƵŹƺě ŚƷ ƽ Ʋſ ƭƹŵ ŹŵƶŤƃ řźƃ ƿ Ǝ ƀƿŵ Ĩ ƽŚƷ

(10)

ÑÑ ƱřŹŚƨưƷƹƽźǀƘƯƝřźƇ

ƱƹŹŵįźĭŹŚĪƃźŧř

ƶŤſŹ įŚƷŹŚŤƟŹįƹŹįř

ƾĭźŝ ŝǀ Ƃ źţ Żř ĭ ǀ ƵŚ ƪƯŚĩ ŵƺŝ ŚƯř ƝLjŤųř ƴƘƯ ƾ Źřŵ ƽ ŝǀ Ʋ Ʊō ŚƷ ŵ ƿ ƵŶ ŶƄƳ ÑÎ Î =

t

ÐÕ =

df

ÎÓÒ Í

P <

ƪĪƃ ç ƶŝŮƿŚŤƳƲƿřƶŝƶūƺţŚŝ

ƾƯźƔƳ ŶſŹ ƶƧ ƾŝŚƴŤūřŹŚŤƟŹ

ŚƷŹ ƱŶƃ ŚƷƹŹLJ ƽ

A. aphidimyza

Żř ŮƐſ īźŝ ƾƯ ŶƳřƺţ ƳŚĪƯ ǀ Ʈƀ źŧŒƯ ŝƶƄěƲƿřŵŹƺųźŝƂƷŚĩŹŵƽ źĭŹŚĪƃŚ

ƱƹŹŵ ƶŤſŹ ŵƺųƽř

 ƲſƾƴƘƿ

O. laevigatus

 ŶƃŚŝ ƶŝ ŹƺƏ ƶŝŚƄƯ



Yasuda & Kimura (2001)

żǀƳ

ƱŚƄƳ ŶƳŵřŵ ƶĩ ŚƷƹŹLJ ƽ ħŻƹŶƄƠĩ

ŢƠƷ ƶƐƤƳ ƽř Źŵ źŝřźŝ źĭŹŚĪƃ Ʊř ƱƹŹŵ ƶŤſŹ ř ƽ ŵƺų Śŝ ŚƷŹ

ƱŶƃ Żř ŮƐſ īźŝ ƾŝŚƴŤūř ŹŚŤƟŹ Ʋƿř Żř ƵŵŚƠŤſř ƹ

 ŵƺų řŹ Żř źƐų řƿ Ʋ ƱřźĭŹŚĪƃ

ŚƷŹ

ƾƯ ŶƳŻŚſ ƲǀƴĤưƷ

 ƶĩŵřŵƱŚƄƳƂƿŚƯŻō

Ʋſ ƴƬĩ ƾ ƶŤƃ ŚƷ ƶŝ ƹŵźƷƽřźŝħźŤƄƯŹŚĪƃƱřƺƴƗ

żǀƳ źĭŹŚĪƃ Ư

ƾ ŶƳřƺţ Źŵ ƳŚĪƯ ǀ Ʈƀ ƾŝŚƴŤūř ŚƷŹƹLJ ƽ

A. aphidimyza

źŧŒƯ ŶƃŚŝ ƹ Źƺƌů

ƵŹƺě ŚƷ ƽ Ʋſ ƭŹŚƸģ ƶŤƃ ŦƗŚŝ Ư ƾ ŵƺƃ ŚƷƹŹLJ ŝǀ Ƃ źţ ŵƺų řŹ Żř ŮƐſ īźŝ ŚƷŹ Ŷƴƴĩ ƶŝ źƔƳ

ƾƯ ſŹ Ŷ ƮƷźŝ ƾƴƘƯƂƴĩ

ƱƹŹŵźĭŹŚĪƃŹƺƌůƲǀŝŹřŵ

ƶŤſŹ ƶŤƃƾĭŹƺěƞƬŤŴƯƪůřźƯƹƽř

į

A. gossypii

źĭŹŚĪƃƲſŮǀūźţŹŵƱřƺŤŝřŹ

ƵŹƺěƶŝ į ƭƹŵƲſ ƵŹƺěƶŝŢŞƀƳ

į ƭŹŚƸģƲſ

ƶŤƃ

(Venzon et al., 2002)

ƂǀŝšŹŶƣŚƿƹ

ƵŹƺěƾƗŚƟŵźţ

į ƶĩŵźĩƺŬŤƀūƶŤƃƭŹŚƸģƲſ

ĪƃƲſŢſřƵŶƃŦƗŚŝ ŹƺƌůŹŵźĭŹŚ

ƵŹƺě į ƭŹŚƸģƲſ ƶŤƃ

ƂǀŝƱřżǀƯƶŝ

ƽŚƷƹŹLJŻřƽźţ

ƶƄě į ƶŤƃ ŶƴĩƶƿŸƜţŹřƺų ě

ǀ Ƃ řŻř ƿ Ʋ

Christensen et al. (2002)

ƱŚƄƳ ŶƳŵřŵ ƶĩ Źƺƌů ƶŤƃ į

A. gossypii

Źŵ Ư ǀ ƎŰ ŦƗŚŝ Ư ƾ ŵƺƃ ƶƧ Ʋſ źĭŹŚĪƃ

Orius majusculus (Reuter)

Ʈĩ źţ Żř ƹŹLJ

A. aphidimyza

ŸƜţ ƿƶ Ŷƴĩ ƴĤưƷ ǀ Ʋ



Lucas (2005)

żǀƳ ƱŚƄƳ ŵřŵ ŢſřƵ ƶĩ Ʈĩřźţ ƞƬŤŴƯƽŚƷ

Ƙưū ǀ Ţ ƶŤƃ ƽ

Macrosiphum euphorbiae (Thomas)

ƶŝ ƱřƺƴƗ ƖŞƴƯ řŸƛ ƾƿ ƽźĭŹŚĪƃ Źŵ

ƱƹŹŵ ƶŤſŹ ƽř ƹŹLJ ƽŹƺŤƫŚŝ

C. vufilabris

Żř ŚƷƹŹLJ ƽ ƶƄě į źĭŹŚĪƃ

A. aphidimyza

ţ Ő źǀŧ ŹřŸĭ

ƾƯ ŶƃŚŝ ƵƹLjƗ ƹŹ ƶĪƴƿř źŝ

Źŵ ƹŹ ƮƷ ƱřźĭŹŚĪƃ ƾƿ

ƶĪƫ Ĩƿ ƪųřŵ ƶŤſŹ

(patch)

ƾƯ Żƹźŝ Źŵ ŶƳřƺţ

ƱōƾŝŚƴŤūřƽŚƷŹŚŤƟŹ

ƯŚƷ Œ ƭŚǀěƂƤƳŶƃŚŝźŧ

ƱŚſŹ ƵŶƿŵŚƳŶƿŚŞƳƶƐŝřŹƲƿřŹŵżǀƳƾƿŚǀưǀƃƽŚƷ

ŵƺƃƶŤƟźĭ

 ƱŚƴģ ƶĩ

Chivers & Smith (1998)

ŵřŵƱŚƄƳ Ƶ ŶƳř ƶĩ ŚƷŹ ƱŶƃ ƱƺƯźƟ ŚƷ ƽ ƭLjƗř źƐų

(alarm pheromone)

ſƺţ Ǝ ƶŤƃ řźŝ ƽ ƾƯƱřźĭŹŚĪƃ

ƽřźŝƾƘŞƴƯŶƳřƺţ

ŚſŚƴƃ ƾƿ Ʊō ŵƺƃƾƤƬţŚƷ

ƂƿŚƯŻō ƶŤƃƎſƺţźƐųƭLjƗřƱƺƯźƟŮƃźţƅƺƈųŹŵƾƿŚƷ

ƹźĭŹŚĪƃƲſŹƺƌůŚŝŚƷ

ƶƫŚƐƯ į ŹŚŤƟŹ

A. aphidimyza

LjƗƲƿřŵƺūƹŚŝ ŗ

ƾƯƾƿŚƿƺŝƮ

ƾţōšŚƤǀƤŰţƽřźŝƾƗƺƋƺƯŶƳřƺţ

(11)

ƶƯŚƳ ƵźƄůƲưŬƳřƽ

ƱřźƿřƾſŚƴƃ èç

ç

 ÎÐîæ

ÑÒ

ŻƲƿřŹŵƾƬǀưĪţšŚƘƫŚƐƯŚŝŚţŶƃŚŝ šŚǀƿżūƹŢƣŵŚŝšLjƯŚƘţƲƿřŻřƽźĮƿŵŵŚƘŝřƶƴǀƯ

Ƃǀŝ ŵźǀĭŹřźƣšƹŚƌƣŵŹƺƯƽźţ

ƪĪƃ ç ƶƀƿŚƤƯ į ŶƇŹŵ ŹLJ ƹ ŚƷ ƽ ƶŤƄĩ ƵŶƃ ƽ

SE)

t (

ƶƄě į ƶŝźĭŹŚĪƃ

ƶƬǀſƹ ƽ źĭŹŚĪƃ

ƱƹŹŵ ƶŤſŹ ƽř

O. laevigatus

Źŵ Źƺƌů Ʋǀƴſ ƭŹŚƸģ (N4)

ƹ ƭƹŵ (N2)

ĭŹƺě ƾ ƶŤƃ

ƾĭźŝĨƀƿŵƹƪƯŚĩƵŚǀĭƽƹŹ

Ɲƹźů ƶŝŚƄƯ ƲǀĮƳŚǀƯ ŚƷ ƱŚƄƳ ƵŶƴƷŵ į ƭŶƗ ŵƺūƹ

ƝLjŤųř ƴƘƯ ƾ Źřŵ Ţſř

t test

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á =

Fig. 2. Comparison of mortality percentage (± SE) of the predatory midge larvae inflicted by interaguild predator, O. laevigatus in the presence of 4th (N4) and 2nd (N2) instar nymphs of aphid on the leaf disks or whole plant. Means with similar letters are not significantly different (t test, á = 0.05).

ŽŚĜſ ƽŹřżĭ ŢƿŚưů Żř ƵŵŚƠŤſř Śŝ ƢǀƤŰţ Ʋƿř

ĭ ƭŚŬƳř ƱřźƸţ ƵŚĮƄƳřŵ ƾƫŚƯ ƽŚƷ ŢƟź

Ţĩźƃ Żř

Koppert Biological System

ƶŝ ƮƷřźƟƪǀƫŵ

ƶǀƫƹřƖŞƴƯƱŵźĩ

į řŹźĪƄţƩŚưĩźĭŹŚĪƃšřźƄů

ŵŹřŵ ƿŚƣōŻř ƱŚ ƶŝŲſřŹƁŹōźŤĩŵƹƽŹŚǀƸƫřƲǀƀůźŤĩŵƾƴǀƀůƾŞŤŬƯźŤĩŵ

Ĩưĩƪǀƫŵ

ƽŚƷ

ƶǀƫƹřŭźƏƶŗřŹřŹŵƵŶƳŻŹř

į ŽŚĜſƢǀƤŰţƲƿř

ƾƯƽŹřżĭ

ŵƺƃ

ƖŝŚƴƯ

Chivers, D. P. & Smith, R. J. F. (1998) Chemical alarm signaling in aquatic predator-prey systems: a review and prospectus. Ecoscience 5, 338-352.

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ÑÓ ƱřŹŚƨưƷƹƽźǀƘƯƝřźƇ

ƱƹŹŵįźĭŹŚĪƃźŧř

ƶŤſŹ įŚƷŹŚŤƟŹįƹŹįř

Christensen, R. K., Enkegaard, A. & Brodsgaard, H. F. (2002) Intraspecific interactions among the predators Orius majusculus and Aphidoletes aphidomyza. International Organization for Biological Control (IOBC) Bulletin 25, 57-60.

Doumbia, M., Hemptinne, J. L. & Dixon, A. F. G. (1998) Assessment of patch quality by ladybirds: role of larval tracks. Oecologia 113(2), 197-202.

Erfan, D. & Ostovan, H. (2004) Species diversity of flower bugs (Family: Anthocoridae) in Shiraz region. Journal of Agricultural Sciences 11, 81-95. [In Persian with English summary].

Frechette, B., Larouche, F. & Lucas, E. (2008) Leucopis annulipes larvae (Diptera:

Chamaemyiidae) use a furtive predation strategy within aphid colonies. European Journal of Entomology 105(3), 399-403.

Griffin, M. L. & Yeargan, K. V. (2002) Oviposition site selection by the spotted ladybeetle Coleomegilla maculata (Coleoptera: Coccinellidae): choices among plant species.

Environmental Entomology 31, 107-111.

Havelka, J. & Zemek, R. (1999) Life table parameters and oviposition dynamics of various populations of the predacious gall-midge Aphidoletes aphidimyza. Entomologia Experimentalis et Applicata 91(3), 481-484.

Hosseini, M., Ashouri, A., Enkegaard, A., Weisser, W. W., Goldansaz, S. H., Nassiri Mahalati, M. & Sarraf Moayeri, H. R. (2010) Plant quality effects on intraguild predation between Orius laevigatus and Aphidoletes aphidimyza. Entomologia Experimentalis et Applicata 135, 208-216.

Jakobsen, L., Enkegaard, A. & Brodsgaard, H. F. (2004) Interactions between two polyphagous predators, Orius majusculus (Hemiptera: Anthocoridae) and Macrolophus caliginosus (Heteroptera: Miridae). Biocontrol Science and Technology 14(1), 17-24.

Kortet, R. & Hedrick, A. (2004) Detection of the spider predator, Hololena nedra by naïve juvenile field crickets (Gryllus integer) using indirect cues. Behaviour 141, 1189-1196.

Labbafi, Y. (1995) Biology of the predatory midge, Aphidoletes aphidimyza Rondani (Dip.;

Cecidomyiidae) and methods for its rearing in laboratory. M. Sc. Thesis. Department of Plant Protection, University of Tehran, Karaj, Iran. [In Persian with English summary].

Lima, S. L. & Dill, L. M. (1990) Behavioral decisions made under the risk of predation; a review and prospectus. Canadian Journal of Zoology 68(4), 619-640.

Lucas, E. (2005) Intraguild predation among aphidophagous predators. European Journal of Entomology 102, 351-364.

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ƱřźƿřƾſŚƴƃ èç

ç

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Lucas, E. & Brodeur, J. (2001) A fox in sheep's clothing: furtive predators benefit from the communal defense of their prey. Ecology 82(11), 3246-3250.

Lucas, E., Coderre, D. & Brodeur, J. (1998) Intraguild predation among aphid predators:

characterization and influence of extraguild prey density. Ecology 79, 1084-1092.

Madadi, H. (1999) Flower bugs of the genus Orius spp. Wolff (Heteroptera: Anthocoridae) in Karaj, Iran. M. Sc. Thesis. Department of Plant Protection, University of Tehran, Karaj, Iran. [In Persian with English summary].

Madadi, H., Enkegaard, A., Brødsgaard, H. F., Kharrazi-Pakdel, A., Ashouri, A., Mohaghegh-Neishabouri, J. (2008) Orius albidipennis (Heteroptera: Anthocoridae):

intraguild predation of and prey preference for Neoseiulus cucumeris (Acari:

Phytoseiidae) on different host plants. Entomologica Fennica 19, 32-40.

Polis, G. A., Myers, C. A. & Holt, R. D. (1989) The ecology and evolution of intraguild predation: potential competitors that eat each other. Annual Review of Ecology and Systematics 20, 297-330.

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Ruzicka, Z. & Havelka, J. (1998) Effects of oviposition deterring pheromone and allomones on Aphidoletes aphidimyza (Diptera: Cecidomyiidae). European Journal of Entomology 95(2), 211-216.

Sato, S., Dixon, A. F. G. & Yasuda, H. (2003) Effect of emigration on cannibalism and intraguild predation in aphidophagous ladybirds. Ecological Entomology 28, 628-633.

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ÑÕ ƱřŹŚƨưƷƹƽźǀƘƯƝřźƇ

ƱƹŹŵįźĭŹŚĪƃźŧř

ƶŤſŹ įŚƷŹŚŤƟŹįƹŹįř

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