The ant-pollination system of Cytinus hypocistis (Cytinaceae), a Mediterranean root holoparasite

The genus Cytinus is composed of rootless, stemless and leafless parasites whose flowers are only visible during the reproductive period when they arise from the host tissues. Most of the taxa occur in Madagascar and South Africa, where mammal pollination has been suggested for one species. There is only one species in the Mediterranean region, and its pollination system has been unknown. Here, a long-term field observation study is combined with experimental pollination treatments in order to assess the pollination biology and reproductive system in the Mediterranean species Cytinus hypocistis. Field studies were carried out in six populations in southern Spain over 4 years. Temporal and spatial patterns of variation in the composition and behaviour of floral visitors were characterized. Pollen loads and pollen viability were observed, and exclusion and controlled-pollination treatments were also conducted. Cytinus hypocistis is a self-compatible monoecious species that relies on insects for seed production. Ants were the main visitors, accounting for 97.4 % of total floral visits, and exclusion experiments showed that they act as true pollinators. They consistently touched reproductive organs, carried large pollen loads and transported viable pollen, although the different ant species observed in the flowers differed in their pollination effectiveness. The abundance of flying visitors was surprisingly low, and only the fly Oplisa aterrima contributed to fruit production and cross-pollination. Mutualistic services by ant are essential for the pollination of Cytinus hypocistis. Although this parasite does not exhibit typical features of the 'ant-pollination syndrome', many other characteristics indicate that it is evolving to a more specialized ant-pollination system. The striking interspecific differences in the pollination systems of Mediterranean Cytinus (ant-pollinated) and some South African Cytinus (mammal-pollinated) make this genus an excellent model to investigate the divergent evolution of pollination systems in broadly disjunct areas.