Pollination, floral deception and evolutionary processes in Eulophia (Orchidaceae) and its allies.
Orchids provide a model system for addressing evolutionary and ecological questions both because of their species diversity, and because the packaging of their pollen into pollinia facilitates the estimation of male and female pollination success. This thesis focuses on the ecology and evolution of pollination systems in the African orchid genus Eulophia, with an emphasis on deceptive pollination, mechanisms promoting cross-pollination, and pollinatordriven speciation. Pollination in the deceptive species E. zeyheriana is shown to depend on flower colour and proximity to the rewarding model species, Wahlenbergia cuspidata (Campanulacae). This study demonstrates the functional importance of colour matching between model and mimic in a floral Batesian mimicry system, as well as the importance of facilitation by the rewarding model [chapter 2]. The pollinaria of the vast majority of Eulophia and Acrolophia species undergo reconfiguration following removal by pollinators, similar to the phenomena first described by Darwin in some European orchids and which he hypothesised to be adaptations to limit pollinator mediated self-pollination. In chapter 3, a less common mechanism – anther cap retention – is described for E. foliosa. Observations of reconfiguration times were compared to the respective visit times by pollinators in a number of orchids (including Eulophia and Acrolophia) and asclepiads. In 18 of 19 species, pollinarium reconfiguration times exceed the average visit times, providing empirical support for Darwin’s cross-pollination hypothesis [chapter 4]. All of the 25 species of Eulophia examined are deceptive, but two of the three species in the small, closely related Cape genus Acrolophia examined in chapter 5 are rewarding. This translates into very high levels of pollen transfer efficiency in the rewarding A. cochlearis relative to the deceptive A. capensis and species of Eulophia. In addition, A. cochlearis exhibits high rates of pollinator-mediated self-pollination, as quantified using a novel method based on levels of inbreeding depression during embryo development. In chapter 6 the evolutionary divergence of long- and short-spurred forms of E. parviflora in response to different pollinators is investigated. This shows that divergence has occurred in floral morphology, scent chemistry and flowering phenology and that this can be attributed to adaptations to the respective bee and beetle pollinators of each form. This thesis also includes case histories of bee pollination in an additional five Eulophia species, and beetle-pollination in two other species of Eulophia with dense inflorescences and slow pollinarium reconfiguration [chapter 7]. In addition, four taxa were found to undergo auto-pollination [chapter 8]. The main conclusions of this thesis are that pollination of food-deceptive species can be enhanced by spatial proximity to, and floral colour matching with, sympatric rewarding species; that selection strongly favours traits that promote cross-pollination; that pollinatorshifts can drive speciation; and that floral adaptations for bee-, beetle-, and auto-pollination are found in South African representatives of Eulophia.