and Evolution © Springer-Verlag 2000 Printed in Austria Convergent evolution a d adaptive radiation of beetle-pollinated angiosperms

Abstract. A literature review of 34 families of flowering plants containing at least one species pollinated primarily by beetles is presented. While the majority of species are represented by mag-noliids and basal monocotyledons specialized, bee-tle-pollinated systems have evolved independently in 14 families of eudicotyldons and six families of petaloid monocots. Four, overlapping modes of floral presentation i plants pollinated xclusively by beetles (Bilabiate, Brush, Chamber Blossom and Painted Bowl) are described. Chamber Blossoms and Painted Bowls are the two most common modes. Chamber Blossoms, found in magnoliids, primitive monocotyledons and in some families of woody eudicots, exploit the greatest diversity of beetle pollinators. Painted Bowls are restricted to petaloid monocots and a few families of eudicots dependent primarily on hairy species of Scarabaei-dae as pollen vectors. In contrast, generalist flowers pollinated by a combination of beetles and other animals are recorded in 22 families. Generalist systems are more likely to secrete nectar and exploit four beetle families absent in specialist flowers. Centers of diversity for species with specialized, beetle-pollinated systems are distributed through the wet tropics (centers for Brush and Chamber Blossoms) to warm temperate-Mediterranean zones (centers for Painted Bowls and a few Bilabiate flowers). It is unlikely that beetles were the first pollinators of angiosperms but specialized, beetle-pollinated flowers must have evolved by the mid-late Cretaceous to join pre-existing guilds of beetle-pollinated gymnosperms. The floras of Australia and western North America suggest that mutualistic interactions between beetles and flowers has been a continuous and labile trend in angiosperms with novel interactions evolving through the Tertiary