Pollination of Petaloid Geophytes by Monkey Beetles (Scarabaeidae: Rutelinae: Hopliini) in Southern Africa

Field observations, floral dissections, and pollen load analyses of insects indicate that pollination by hopliine beetles (Scarabaeidae: Rutelinae: Hopliini) has evolved convergently in many genera of herbaceous perennials in southern Africa. Beetle-pollinated flowers are identified by a suite of characters including a salver-to shallow bowl-shaped perianth and pigmentation emphasizing bright colors (red. orange, cream). Stereotyped "beetle marks" of either pale or dark color are frequently present at the bases of tepals or petals. These flowers are typically odorless and rarely offer nectar. Beetles, however, consume anthers and pollen, which are often a contrasting color from the perianth. Taxa that are pollinated by hopliine beetles include species in genera of the Hyacinthaceae (I)aubenya, Ornithogalum), Iridaceae (Aristea, Homeria, Ixia, Moraea, Ro-mulea, Sparaxis, Tritonia), and Hypoxidaceae (Spiloxene) in the monocots and Asteraceae (Arctotis, Ursinia), Campanulaceae (PrismcUocarpus, Wahlenbergia), and Droseraceae (Drosera) among the dicots. Hopliine pollinators include both male and female l>eetles in the genera Anisonyx, Anzsochelus, Heterochelis, Khoinu, Ijepisia, Lepithrix, Pacbycnema, and Peritrichia. These beetles visit flowers to consume pollen and possibly nectar, to compete for mates, and to copulate. Pollen is usually deposited on or between hairs on the exoskeleton. The suite of characters associated with beetle pollination in these herl)aceous geophytes is closer to that described in the herbaceous perennials