Thorn-like prickles and heterophylly in Cyanea: adaptations to extinct avian browsers on Hawaii?

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Abstract
The evolution of thorn-like structures in plants on oceanic islands that lack mammalian and reptilian herbivores is puzzling, as is their tendency toward juvenile-adult leaf dimorphism. We propose that these traits arose in Cyanea (Campanulaceae) on Hawaii as mechanical and visual defenses against herbivory by flightless geese and goose-like ducks that were extirpated by Polynesians within the last 1600 years. A chloroplast DNA phylogeny indicates that thorn-like prickles evolved at least four times and leaf dimorphism at least three times during the last 3.7 million years. The incidence of both traits increases from Oahu eastward toward younger islands, paralleling the distribution of avian species apparently adapted for browsing. The effectiveness of visual defenses against avian browsers (once dominant on many oceanic islands, based on the vagility of their ancestors) may provide a general explanation for insular heterophylly: the other islands on which this previously unexplained phenomenon is marked (New Zealand, New Caledonia, Madagascar, Mascarene Islands) are exactly those on which one or more large flightless avian browsers evolved.