The presence of heterospecific pollen on stigmas of nectariferous and nectarless orchids and its consequences for their reproductive success

The widespread occurrence of nonorchid, heterospecific pollen grains on the stigmatic surfaces of a range of nectariferous and nectarless European orchids (Dacrylorhiza, Orchis, Goodyera, and Gymnadenia species) is reported for the first time, and the impact of heterospecific pollination on orchid reproductive success is experimentally investigated. There are three main components of stigmatic contamination by heterospecific pollen: the frequency of contamination, the diversity of foreign species present on the stigma, and the amount of pollen deposited. Six out of seven of the species examined have greater than 85% of stigmas contaminated with wind and insect-dispersed pollen. From one to nine insect-dispersed foreign pollen species are present per stigma, including pollen of members of the families Apiaceae, Asteraceae, Caryohpyllaceae, Ericaceae, and Primulaceae. Average loads per stigma vary from 13 to 176 grains, with individual stigma loads ranging from one to 909. Whether or not the orchid provides nectar has a major impact on these three components. Nectarless orchids have the greatest frequencies of contamination, diversity of species, and average load per stigma. Insect-dispersed pollen is deposited both by pollinators and visitors but, in spite of low levels of pollination, nectarless orchids still exhibit higher frequencies of heterospecific pollen contamination. The effect of the presence of heterospecific pollen on the reproductive success of orchids is tested in this study for the first time. Average-to-high, naturally occurring loads of heterospecific pollen derived from a mixture of Armeria maritima, Caltha palustris, Cochlearia officinalis, Cytisus scoparius, and Primula vulgaris and consisting of 50-250 grains per load are placed onto stigmas of Dactylorhiza purpurella which are subsequently self-pollinated with half of a pollinium. All pollinations produce capsules indicating that heterospecific pollen does not affect fruit set. Although experimental and control fruits are similar in size, they differ in total seed weight and composition. Total seed weight is reduced and the proportion of seeds with normal embryos decreased while the proportion of unfertilised ovules increased following pollination with heterospecific pollen, indicating a detrimental effect on fertilisation. Lower reproductive success caused by fertilisation failure is likely to be most severe in nectarless species because of their generally higher levels of contaminated stigmas. As nectarless orchids are known to have lower fruit set compared with nectariferous ones, this finding suggests that the reproductive success of nectarless orchids may be even lower than previously realised.