Significant ant pollination in two orchid species in the Alps as adaptation to the climate of the alpine zone?

\ Ants were shown to be significant pollinators of two orchid species in the alpine zone of the Alps. Repeated observations from several localities confirm the ant Formica lemani as pollinator of Chamorchis alpina whereas Formica exsecta is reported here for the first time as pollinator of Dactylorhiza viridis. These findings appear of great interest, as significant ant pollination of orchids is unknown so far from any other region or habitat type in the Holarctic. This raises the question if there are specific adaptations. The observations do not provide suggestions to adaptations of the Formica ants for pollinating orchids they simply followed their normal foraging behavior shown in any type of habitat. Yet, special adaptations are given by the two orchid species in developing pollination mechanisms more strongly involving ground-moving insects which are not inactivated by increased wind velocity and lower temperatures in the alpine zone. These are mainly beetles and ants. The pollination mechanisms and interactions with ants of both orchid species are described. Dactylorhiza viridis is outstanding among orchids in needing 20 to 30 minutes for the 90-degree forward bending of the pollinium after attachment to the insect's forehead whereas the same process takes place between 15 seconds and 3 to 5 minutes in other orchids. Forward bending of pollinia is required for precise placement of the pollen at the stigmatic surface of another flower. The very long bending time is an adaptation to the longer presence time of ground-moving insects at the same plant and aims to reduce the frequency of geitonogamy (self-fertilization). The high frequency of ant pollination in these orchids is a consequence of the high activity density of aggressive, predatory worker ants leading to a displacement of other pollinators. Attempts of ants to remove the fresh, strongly adhesive pollinia from their foreheads failed and a single ant head could carry up to eight pollinia.