How the daisy got its spots... and why

Dark spots on flower petals are common across many angiosperm plant families and occur on flowers such as some lilies, orchids, and daisies. Much research has been done on the physiological and behavioral mechanisms for how these spots attract pollinators. But have you ever wondered what these spots are composed of, how they develop, or how they only appear on some but not all of the ray florets?

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