Abstract
An apparent contradiction in the ecology of cycad plants is that their seeds are known to be highly poisonous, and yet they seem well adapted for seed dispersal by animals, as shown by their visually conspicuous seed cones and large seeds presented within a brightly colored fleshy “fruit” of sarcotesta. We tested if this sarcotesta could function as a reward for cycad seed dispersal fauna, by establishing if the toxic compound cycasin, known from the seeds, is absent from the sarcotesta. The Australian cycads Macrozamia miquelii and Cycas ophiolitica were tested (N = 10 individuals per species) using gas chromatography / mass spectrometry. Cycasin was detected at 0.34 % (fresh weight) in seed endosperm of M. miquelii and 0.28 % (fresh weight) in seed endosperm of C. ophiolitica. Cycasin was absent from the sarcotesta of the same propagules (none detected in the case of M. miquelii, and trace quantities detected in sarcotesta of only four of the ten C. ophiolitica propagules). This laboratory finding was supported by field observations of native animals eating the sarcotesta of these cycads but discarding the toxic seed intact. These results suggest cycads are adapted for dispersal fauna capable of swallowing the large, heavy propagules whole, digesting the non-toxic sarcotesta flesh internally, and then voiding the toxic seed intact. Megafauna species such as extant emus or cassowaries, or extinct Pleistocene megafauna such as Genyornis, are plausible candidates for such dispersal. Cycads are an ancient lineage, and the possible antiquity of their megafaunal seed dispersal adaptations are discussed.
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Acknowledgments
The GCMS method for the detection of cycasin was developed by Mr Grahame Kervan (UQ School of Land, Crop and Food Sciences). Our sincere thanks to him, as well as Dr Leslie Force and Dr Jennifer Waanders (UQ School of Land, Crop and Food Sciences) and Dr Susanne Schmidt (UQ School of Biological Sciences) for their assistance in the implementation of this GC/MS technique, and the interpretation of the results. We also thank Dr Chris Moore for suggesting some improvements to the draft and and giving us the benefit of considerable expertise in analytical chemistry. This study would not have been possible without the kind support Dr Glen Kisby (Oregon Health and Science University) in generously providing a pure standard of the toxic compound cycasin. We thank Dr James Gilardi (World Parrot Trust) for help in interpreting his published data (Gilardi and Toft 2012) on the toxicity of seeds and fruits in the diet of Amazonian parrots. The collection of wild cycad seed discussed in this paper was undertaken in conjunction with the Queensland Government Department of Environment and Resource Management scientific purposes permit number WITK06373009.
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Hall, J.A., Walter, G.H. Relative Seed and Fruit Toxicity of the Australian Cycads Macrozamia miquelii and Cycas ophiolitica: Further Evidence for a Megafaunal Seed Dispersal Syndrome in Cycads, and Its Possible Antiquity. J Chem Ecol 40, 860–868 (2014). https://doi.org/10.1007/s10886-014-0490-5
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DOI: https://doi.org/10.1007/s10886-014-0490-5