Due to the rise of monoculture in agricultural production, insect biodiversity has sharply declined in agroecosystems due to the reduction in host plant biodiversity. This includes declines in populations of native pollinators, which reduces the ambient pollination services received by crops and increases the reliance on managed pollinators like European honeybees (Apis mellifera). One way to combat this decline in pollination services is to develop crop varieties that have high attraction to pollinators, this can have a dual effect of inducing native pollinators to move deep into agricultural fields from adjacent habitats like woodland or meadow strips, as well as retaining managed honeybees within crop fields and improving their efficacy. Attraction can be determined by many traits including floral morphology, pigmentation, and chemical cues like fragrance. Under higher ambient temperature the floral temperature rises, resulting in several potential effects. A warmer flower is likely to emit a larger volume of volatile compounds, as well as to be an attractive resting place for ectothermic pollinators, and yet high floral temperature may reduce pollen viability. The goal of this study is to first assess the effects of floral traits, particularly morphology and pigmentation, on floral temperature relative to ambient conditions, and secondarily, how flower temperature differentials alongside floral traits may affect pollinator foraging preferences. Our findings indicate that floral morphology and pigmentation both influence floral temperature differentials, and that floral morphology, pigmentation, and temperature differentials influence visitation by native pollinators (bumblebees and sweat bees). Floral traits and their influence on temperature are potentially important targets for the improvement of pollinator visitation to sunflower cultivars and the improvement of yields.

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It is well known that animals can exert strong selective pressures on plant traits. However, studies on the evolutionary consequences of plant–animal interactions have mainly focused on understanding how these interactions shape trait means, while overlooking its potential direct effect on the variability among structures within a plant (e.g. flowers and fruits). The degree of within-plant variability can have strong fitness effects but few studies have evaluated its role as a potential target of selection. Here we reanalysed data on Ipomoea wolcottiana stigma–anther distance to test alternate mechanisms driving selection on the mean as well as on intra-individual variance in 2 years. We found strong negative selection acting on intra-individual variation but not on mean stigma–anther distance, suggesting independent direct selection on the latter. Our result suggests that intra-individual variance has the potential to be an important target of selection in nature, and that ignoring it could lead to the wrong characterisation of the selection regime. We highlight the need for future studies to consider patterns of selection on the mean as well as on intra-individual variance if we want to understand the full extent of plant–animal interactions as an evolutionary force in nature.

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Background: Pollen transfer via animals is necessary for reproduction by ~80 % of flowering plants, and most of these plants live in multispecies communities where they can share pollinators. While diffuse plant-pollinator interactions are increasingly recognized as the rule rather than the exception, their fitness consequences cannot be deduced from flower visitation alone, so other proxies, functionally closer to seed production and amenable for use in a broad variety of diverse communities, are necessary. Scope: We conceptually summarize how the study of pollen on stigmas of spent flowers can reflect key drivers and functional aspects of the plant-pollinator interaction (e.g. competition, facilitation or commensalism). We critically evaluate how variable visitation rates and other factors (pollinator pool and floral avoidance) can give rise to different relationships between heterospecific pollen and (1) conspecific pollen on the stigma and (2) conspecific tubes/grain in the style, revealing the complexity of potential interpretations. We advise on best practices for using these proxies, noting the assumptions and caveats involved in their use, and explicate what additional data are required to verify interpretation of given patterns. Conclusions: We conclude that characterizing pollen on stigmas of spent flowers provides an attainable indirect measure of pollination interactions, but given the complex processes of pollen transfer that generate patterns of conspecific-heterospecific pollen on stigmas these cannot alone determine whether competition or facilitation are the underlying drivers. Thus, functional tests are also needed to validate these hypotheses.

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Thesis (PhD)--Stellenbosch University, 2013. / Pollinators play a fundamental role in floral evolution. They can exert selection on the flowers they visit in a plethora of different ways, ranging from innate floral preferences to differences in body size and shape and behavioural elements such as flower constancy and learning capacity. Since different pollinators exhibit differences in these characters, shifts between pollinating species are often considered the most likely drivers of floral diversification. While many lines of evidence support this claim, numerous angiosperms pollinated by a single species also exhibit floral variation. Throughout my thesis, I explore and investigate floral diversification in such species in the absence of pollinator shifts. In Chapter 2, I investigate variation in the preference of conspecific male and female pollinators for the floral traits of a sexually deceptive daisy that comprises distinct floral forms. I show that its pollinator exhibits gender-specific variation in floral preferences, and that some floral forms have specialized on the male pollinator. This chapter thus illustrates the importance of intraspecific variation in pollinator preference for floral diversification, an underappreciated mechanism in this field of research. The innate preferences of pollinators are likely to have a genetic basis, especially innate preferences that govern mate choice. Genetic structure within the pollinators of sexually deceptive plants, which mimic female insects to achieve pollination, may thus provide an important source of selection on the plants they pollinate. This depends on an association between genetic divergence and divergent mate preferences, and I explore this intriguing idea in Chapter 3. While pollinators associated with sexually deceptive floral forms did exhibit significant genetic structuring, male pollinators from different phylogeographic clades all exhibited preference for the same sexually deceptive floral form, thus rejecting this hypothesis. Another behavioural attribute of pollinators that may affect floral evolution, particularly in deceptive plant species, is learning ability. Studies on sexually deceptive orchids often report that male pollinators tend to avoid sexually deceptive flowers with experience. In Chapter 4, I systematically investigate learning abilities within male pollinators and the costs they suffer on sexually deceptive floral forms that vary in deceptiveness. Results reveal a positive relationship between the level of floral deceptiveness and the 4 associated mating costs that deceived males suffer. Pollinator learning, however, appears to occur only on the most deceptive floral forms, suggesting a link between the costs suffered to the occurrence of learning. In Chapter 4, I systematically investigate learning abilities within male pollinators and the costs they suffer on sexually deceptive floral forms that vary in deceptiveness. Results reveal a positive relationship between the level of floral deceptiveness and the associated mating costs that deceived males suffer. Pollinator learning, however, appears to occur only on the most deceptive floral forms, suggesting a link between the costs suffered to the occurrence of learning. In Chapter 5, I explore the importance of florivory damage in a polymorphic daisy. Studies on floral evolution often overlook the significance of florivorous visits and focus only on pollinator-mediated selection. I show that floral polymorphism is maintained by antagonistic selection exerted by pollinators and florivores on the same floral traits. Lastly, I focus on evolutionary history to explore similarity in the patterns of South African angiosperm evolution and the pollinator species used throughout my thesis. Molecular dating shows this pollinator exhibits broadly congruent evolutionary patterns to these angiosperms, indicative of a shared biogeography. Taken together, my thesis demonstrates the vast impact of floral visitors, in particular pollinating insects, on the evolution of floral form. / My research was funded by the National Research Foundation of South Africa (NRF) and personal funding was provided by a NRF Innovation scholarship and merit bursaries from the Botany and Zoology department at Stellenbosch University. A WhiteSci Travel Grant and financial support from Prof. Erik Svensson at Lund University also allowed me to present parts of my research at international conference.

There is growing awareness about the value of preserving and restoring floral-rich habitats for the benefit of pollinators, especially native bees. The increasing demand for native plants in pollinator habitat restoration and other ecological landscaping applications, combined with the desire for more robust and predictable plant habits, have led to the selection and breeding of native cultivars. Yet, little is known about how these cultivated varieties differ from the native species in their ability to attract and support pollinators. I compared flower visitation by all insect pollinators to 12 native herbaceous plant species and 14 native cultivars in a replicated field experiment at two sites over two years. I classified insect pollinators during visual field observations into seven taxonomic and functional groups. I found seven native species to be visited significantly more frequently by all insect pollinators (combined) than their cultivars, four were visited equally, and one native cultivar was visited more frequently than the native species. Bees (both native and non-native) and moths/butterflies exhibited similar preferences, whereas flies showed no preference between the native species and the native cultivar. Our study shows that many insect pollinators prefer to forage on native species over cultivated varieties of the native species, but not always, and not exclusively. Some native cultivars may be comparable substitions for native species in pollinator habitat restoration projects, but all cultivars should be evaluated on an individual basis. Plant selection is integral to the value and success of pollinator habitat restorations, yet there is little consistency and overlap in pollinator planting recommendations and very little empirical data to support plant choice. Non peer-reviewed pollinator plant lists are widely available and are often region-specific, but they are typically based on anecdotal rather than empirical data and lack in specificity. To help close the gap between anecdotal and empirical data, and between practice and research, I reviewed the published literature on plant selection for pollinator habitat restoration. I explicitly reviewed and compared the value of native plant species, near-natives, non-natives and native cultivars. From there, I identified gaps in the literature that are most needed in practice and recommended basic strategies for practitioners to navigate plant lists and choose the best plants for a site's success.

Orientador: Felipe Wanderley Amorim / Resumo: As mudanças globais mediadas pela ação antrópica estão alterando a biodiversidade e os ecossistemas em um ritmo acelerado. Um dos principais impulsionadores dessas mudanças é a introdução de espécies exóticas em ecossistemas nativos. Entre os grupos de organismos afetados por este processo, o das plantas é reconhecido um dos mais preocupantes, uma vez que a produção primária limita o tamanho e a composição das comunidades e participa através de efeitos em cascata em interações multi-tróficas. Uma das principais relações ecológicas influenciada por esse efeito é a relação entre plantas e polinizadores, cujo papel é importante para estrutura e funcionamento das comunidades biológicas, não apenas porque as plantas fornecem recursos alimentares essenciais para muitos grupos de animais que visitam flores, mas também porque o sucesso reprodutivo da maioria das plantas com flores depende dos serviços bióticos fornecidos por estes animais. Neste contexto, a introdução de espécies de plantas exóticas invasoras pode ter impactos críticos nas interações planta-polinizador ao nível da comunidade, principalmente através da competição com espécies nativas. Como as interações planta-polinizador são cruciais para determinar a estrutura da comunidade, nesse estudo nós testamos como o crescimento rápido de uma gramínea invasora altera a composição das espécies de plantas nativas em um campo cerrado, juntamente com os impactos deste processo sobre a estrutura das interações planta-polinizador. ... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: The global change mediated by anthropic action is altering biodiversity and ecosystems in a fast pace. One major driver of these changes is the introduction of alien species in native ecosystems. Among the groups of organisms that are affected by this process, plants are recognized to be one of the most concern, since primary production limit global communities’ sizes and composition, and participate through cascade effects on multitrophic interactions. One crucial type of interaction that is influenced by this effect is the plant-pollinator relationship, which have an important role in the structure and functioning of biological communities, not only because plants provide essential food resources for many groups of animals that visit flowers, but also because the reproductive success of most flowering plants depends on the biotic services provided by animals. In this context, the introduction of invasive alien plant species may have critical impacts on plant-pollinator interactions at community level, mainly through competition with native species. Since plantpollinator interactions are determinants of community structure, here we evaluated how the rising of a fast-growing invasive alien grass species changes plant species composition of a Neotropical grassland community along with its impacts on the structure of plant-pollinator interactions. For this, we analyzed the changes in community composition and plantpollinator interactions over time, through the temporal turnover... (Complete abstract click electronic access below) / Mestre