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The Role of Macrozamin and Cycasin in Cycads (Cycadales) as Antiherbivore Defenses

Citlalli Castillo-Guevara and Victor Rico-Gray
The Journal of the Torrey Botanical Society
Vol. 130, No. 3 (Jul. - Sep., 2003), pp. 206-217
Published by: Torrey Botanical Society
DOI: 10.2307/3557555
Stable URL: http://www.jstor.org/stable/3557555
Page Count: 12
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The Role of Macrozamin and Cycasin in Cycads (Cycadales) as Antiherbivore Defenses
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Abstract

Macrozamin and cycasin are very toxic azoxyglycosides of the Cycadales. Caterpillars (Lepidoptera) and weevils (Coleoptera) feed on cycad root, stem, leaf, and reproductive tissues. Azoxyglycosides may have played an important ecological role as antiherbivore defenses. We evaluated the association between herbivory and the amount of azoxyglycosides in the Cycadales using phylogenetic independent contrasts. We hypothesized that herbivory types should be related to the presence of macrozamin and cycasin, thus herbivory should be lower in species with higher concentrations of azoxyglycosides. We gathered information available on the literature of these two characters as well as life form, geographic distribution, height, and seed volume for the majority of cycad species, in order to assess correlated evolution and control for possible allometric effects. Herbivory types and macrozamin were negatively correlated, suggesting a possible defensive function for macrozamin against herbivores. No significant correlation was observed between cycasin percent and herbivory type. However, when analysed using phylogenetic independent contrasts and thus removing the historical effect, the association did not hold. This suggests that the presence of metabolites in plants may have evolved for some other reason, and has been mantained among cycads perhaps by phylogenetic inertia. The presence of macrozamin should then be explained as an exaptation, playing today an important role in defense against herbivores. Furthermore, this analysis showed that macrozamin has independently and repeatedly (Bowenia, Macrozamia, Stangeria) increased over evolutionary time.

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