Species employ multiple strategies to deal with stressful environments, but these strategies often incur costs. Aphids frequently utilize multiple predator avoidance strategies, including attracting mutualist ants for protection and dispersing by producing winged forms. While both strategies can be physiologically costly, the magnitudes of these costs have not been previously compared. In this study, we experimentally manipulated ant attendance in the field and measured the individual and interactive effects of ant attendance and wing formation on body size and reproduction of the ant-tended aphid Cinara schwarzii (Wilson) (Hemiptera: Aphididae). Aphid adult body size was smaller in the presence of ants (18%), but controlling for body size, there were no differences in embryo number or size. In contrast, wing formation did not affect adult body size but strongly reduced embryo number (46%) and size (8%). Although ant attendance reduced C. schwarzii wing formation, ant attendance and wing formation acted independently on aphid body size and reproduction. For comparison, we confirmed that the manipulation of ant presence had no effect on body size or reproduction of the untended co-existing congener Cinara solitaria (Gillette and Palmer) (Hemiptera: Aphididae). Complementing our empirical study, a meta-analysis of 78 responses from 24 publications showed that wing formation consistently and significantly reduces aphid body size and reproduction (37%), while the effects of ant attendance showed a mean positive effect (9%) that did not significantly differ from zero. Together, our empirical study and meta-analysis provide strong evidence for costs of wing formation but not ant attendance for aphids.
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