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The evolution of antiherbivore defenses and their contribution to species coexistence in the tropical tree genus Inga
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Edited by Anurag A. Agrawal, Cornell University, Ithaca, NY, and accepted by the Editorial Board July 27, 2009 (received for review May 1, 2009)

Abstract
Plants and their herbivores constitute more than half of the organisms in tropical forests. Therefore, a better understanding of the evolution of plant defenses against their herbivores may be central for our understanding of tropical biodiversity. Here, we address the evolution of antiherbivore defenses and their possible contribution to coexistence in the Neotropical tree genus Inga (Fabaceae). Inga has >300 species, has radiated recently, and is frequently one of the most diverse and abundant genera at a given site. For 37 species from Panama and Peru we characterized developmental, ant, and chemical defenses against herbivores. We found extensive variation in defenses, but little evidence of phylogenetic signal. Furthermore, in a multivariate analysis, developmental, ant, and chemical defenses varied independently (were orthogonal) and appear to have evolved independently of each other. Our results are consistent with strong selection for divergent defensive traits, presumably mediated by herbivores. In an analysis of community assembly, we found that Inga species co-occurring as neighbors are more different in antiherbivore defenses than random, suggesting that possessing a rare defense phenotype increases fitness. These results imply that interactions with herbivores may be an important axis of niche differentiation that permits the coexistence of many species of Inga within a single site. Interactions between plants and their herbivores likely play a key role in the generation and maintenance of the conspicuously high plant diversity in the tropics.
Footnotes
- 1To whom correspondence should be addressed. E-mail: kursar{at}biology.utah.edu
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Author contributions: T.A.K., K.G.D., J.L., R.T.P., and P.D.C. designed research; T.A.K., K.G.D., J.L., R.T.P., J.E.R., M.G.W., E.T.M., C.D., R.M., and P.D.C. performed research; K.G.D., J.L., R.T.P., J.E.R., and P.D.C. analyzed data; and T.A.K., K.G.D., J.L., R.T.P., and P.D.C. wrote the paper.
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The authors declare no conflict of interest.
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This article is a PNAS Direct Submission. A.A.A. is a guest editor invited by the Editorial Board.
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Data deposition: The sequences reported in this paper have been deposited in the GenBank database (accession nos. FJ974145, FJ974151, FJ974164, FJ974180, FJ974241, FJ974279, FJ974351, FJ974409, FJ974438, FJ974449, FJ974509, FJ974628, FJ974673, FJ974683, FJ974713, FJ974870, FJ974883, FJ974910, FJ974975, FJ974988, FJ975003, FJ975011, FJ975039, GQ118709–GQ118890, GQ871263–GQ871278, and GQ892055).
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This article contains supporting information online at www.pnas.org/cgi/content/full/0904786106/DCSupplemental.
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