Floral morphology can play a key role in mechanically guiding pollinators towards reproductive structures, particularly when visibility is limited at night, but the functional significance of morphological traits has seldom been considered in this context. Here we describe a remarkably intricate pollination mechanism in the hawkmoth-pollinated African grassland orchid Habenaria clavata, and also document aspects of the reproductive success and chemical ecology of this pollination system. The flowers are pollinated by several short-tongued hawkmoths, particularly Basiothia schenki, which was the most frequent visitor and occurred at all sites. Moths are probably attracted by the strong scent, which was dominated by several oxygenated aromatics that also elicited strong electrophysiological responses from antennae of B. schenki. Apart from the white rostellum lobes and stigma, which serve as a visual guide to the spur entrance, the flower parts are entirely green and indistinguishable from leaves in terms of spectral information. Using motion-activated video cameras we established that the leading edges of the forewings of foraging hawkmoths contact the two upwardly curving petal lobes, and that hawkmoths are then apparently mechanically guided down onto the reproductive structures. Pollinaria are attached in an unusual place – among hairs on the ventral surface of the thorax, between the middle legs – and are brushed over the protruding stigma lobes when the proboscis is fully inserted in the 41-mm-long spur. These results highlight how multiple traits (morphology, spectral reflectance and scent) can act synergistically to ensure transfer of pollen among flowers.

Background and Aims The temporal dimensions of floral adaptation to pollinators are not yet well understood, partly because we lack accurate information on the diel rhythms of flower visitation for many pollinators. We investigated whether diel patterns of pollinator visitation to flowers of the African woodland orchid Bonatea polypodantha are synchronised with rhythms of floral anthesis, scent emission and nectar availability. Methods Direct observations and motion-activated cameras were used to identify pollinators of B. polypodantha and to document their activity periods. The timing of pollinaria removal from flowers, emission of scent and availability of nectar were also measured. Results We found that B. polypodantha is pollinated exclusively by short-tongued hawkmoths. Pollinaria of the orchid are affixed between the labial palps of the moths and brush over the protruding stigmatic arms. The flowers also receive visits by long-tongued hawkmoths, but these act as nectar thieves. Tracking of pollinaria removal from flowers confirmed that pollination occurs only at night. Camera footage revealed a striking crepuscular pattern of foraging by short-tongued hawkmoths with peaks of activity during the twilight periods at dusk and at dawn. In contrast, long-tongued hawkmoths were found to visit flowers throughout the night. Flowers of B. polypodantha exhibit unimodal peaks of anthesis, scent emission (dominated by nitrogenous aromatics) and nectar availability before or around dusk. Conclusions Flowers of B. polypodantha are pollinated exclusively by short-tongued hawkmoths which show crepuscular foraging activity at dusk and dawn. Floral phenophases of the orchid are closely synchronised with the peak of pollinator activity at dusk.

1. Diel rhythms of foraging activity by animal flower visitors can reflect niche partitioning and are considered an important component of selection on floral traits. However, it has been notoriously difficult to obtain objective information on the patterns of flower visitation by crepuscular and nocturnal insects.2. Motion-activated cameras were used for field-based studies of hawkmoth foraging behaviour on six African plant species.3. The results showed that short-tongued hawkmoth species forage mainly around dusk and then sporadically throughout the night, whereas long-tongued hawkmoth species feed consistently throughout the night, with a peak shortly before midnight.4. These results provide the first quantitative estimates of diel patterns of interactions between multiple hawkmoth and plant species and, when combined with qualitative reports from other studies, suggest that differences in diel activity between the two main hawkmoth functional groups (short-and long-tongued) are consistent across the Old and New Worlds.

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