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Pollinator-Mediated Selection and the Evolution of Floral Traits Inorchids

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Pollinator-Mediated Selection and the Evolution of Floral Traits Inorchids Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1677 Pollinator-mediated selection and the evolution of floral traits in orchids JUDITH TRUNSCHKE ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 ISBN 978-91-513-0352-9 UPPSALA urn:nbn:se:uu:diva-349139 2018 Dissertation presented at Uppsala University to be publicly examined in Zootissalen, Villavägen 9, Uppsala, Thursday, 14 June 2018 at 10:00 for the degree of Doctor of Philosophy. The examination will be conducted in English. Faculty examiner: Professor Allan G. Ellis (Stellenbosch University). Abstract Trunschke, J. 2018. Pollinator-mediated selection and the evolution of floral traits in orchids. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1677. 43 pp. Uppsala: Acta Universitatis Upsaliensis. ISBN 978-91-513-0352-9. In this thesis, I combined manipulations of traits and pollination environment with analysis of phenotypic selection to examine causes of variation in strength and mode of selection on floral traits, and I conducted a reciprocal sowing experiment to test for local adaptation in germination success. I tested the following predictions (1) the opportunity for selection, and the strength of pollinator-mediated and net selection increase with increasing pollen limitation, (2) the effects of traits affecting pollinator attraction and traits affecting pollination efficiency are non-additive and this leads to pollinator-mediated correlational selection, (3) the effects of spur length on pollen removal, pollen receipt, and female fitness differ between populations with short-tongued and populations with long-tongued pollinators, and (4) local adaptation at the stage of germination contributes to the maintenance of ecotypes growing in grasslands and woodlands, respectively. A study including natural populations of 12 orchid species that varied widely in pollen limitation showed that opportunity for selection, pollinator-mediated selection and net selection were all positively related to pollen limitation, whereas non-pollinator-mediated selection was not. In the moth-pollinated orchid Platanthera bifolia, experimental reductions of plant height and spur length decreased pollen removal, pollen receipt and fruit production, but non-additive effects were not detected. Effects of plant height translated into pollinator-mediated selection for taller plants via female fitness, but there was no current pollinator-mediated selection on spur length. An experiment using artificial nectar spurs demonstrated that in P. bifolia pollen receipt saturated at shorter spur length in a population with short-tongued pollinators than in a population with a long-tongued pollinator. Effects of spur length on pollen receipt did not translate into current pollinator-mediated selection indicating that also plants with the shortest spurs for the most part received sufficient pollen for full seed set. Reciprocal sowing of seeds from grassland and woodland populations detected no evidence of local adaptation at the germination stage between ecotypes of P. bifolia. Taken together, the results illustrate how a combination of trait manipulation and analysis of strength and causes of selection can throw light on both the functional and adaptive significance of trait variation within and among natural populations. Keywords: Among-population variation, Ecotype, Field experiment, Floral evolution, Germination, Local adaptation, Orchidaceae, Platanthera bifolia, Phenotypic selection, Pollination, Reciprocal transplant, Trait manipulation Judith Trunschke, Department of Ecology and Genetics, Plant Ecology and Evolution, Norbyvägen 18 D, Uppsala University, SE-752 36 Uppsala, Sweden. © Judith Trunschke 2018 ISSN 1651-6214 ISBN 978-91-513-0352-9 urn:nbn:se:uu:diva-349139 (http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-349139) “If you don´t climb the mountain, you can´t see the view” (Harvey Mackay) For my parents and my sister List of Papers This thesis is based on the following papers, which are referred to in the text by their Roman numerals. I Trunschke, J., Sletvold, N. and Ågren, J. (2017) Interaction intensity and pollinator-mediated selection. New Phytologist, 214: 1381-1389 II Trunschke, J., Sletvold, N. and Ågren, J. The independent and com- bined effects of floral traits distinguishing two pollination ecotypes of a moth-pollinated orchid. (Manuscript) III Trunschke, J., Sletvold, N. and Ågren, J. Pollinator-mediated selec- tion and effects of spur length on pollination success in short-spurred and long-spurred populations of the orchid Platanthera bifolia. (Manuscript) IV Trunschke, J., Sletvold, N. and Ågren, J. No evidence for adaptive population differentiation during seed germination between wood- land and grassland populations of the orchid Platanthera bifolia. (Manuscript) Reprints were made with permission from the respective publishers. Contents Introduction ..................................................................................................... 9 Biodiversity and the importance of ecology for evolution .................... 9 Quantifying phenotypic selection ........................................................ 10 Opportunity for selection and strength of selection ............................. 11 The importance of experimental manipulations for understanding selection ............................................................................................... 12 Pollinator-mediated selection ............................................................... 13 Traits important for pollinator attraction and pollination efficiency ... 14 Aims of the thesis ................................................................................. 16 Material and Methods .................................................................................... 17 Deceptive and rewarding orchids ......................................................... 17 Platanthera bifolia and its ecotypes .................................................... 18 Pollen limitation and strength of pollinator-mediated selection (I) ..... 19 Manipulation of plant height and spur length, and correlational selection (II) ......................................................................................... 20 Manipulation of spur length and pollinator-mediated selection (III) ..... 22 Reciprocal transplant (IV) .................................................................... 23 Results and Discussion .................................................................................. 24 Pollen limitation and strength of pollinator-mediated selection (I) ..... 24 Effects of plant height and spur length on pollinator interactions (II) ... 26 Pollinator-mediated selection and the effects of spur length on pollination success (III) ........................................................................ 27 Local adaptation in germination success (IV) ..................................... 29 Conclusions .......................................................................................... 31 Svensk sammanfattning ................................................................................. 32 Acknowledgements ....................................................................................... 36 References ..................................................................................................... 38 Introduction Biodiversity and the importance of ecology for evolution Eight point seven million is an impressively large number. It is the currently estimated number of species inhabiting our planet (Mora et al. 2011), and a numerical expression of the tremendous diversity of living forms and func- tions surrounding us. Understanding the processes generating and maintain- ing differences among populations and ultimately speciation are fundamental question addressed by evolutionary biologists. Many plant and animal species are widely distributed, occur in a variety of habitats and show substantial variation among populations. Local adapta- tion has been documented for both plants and animals across several spatial scales (Leimu and Fischer 2008, Hereford 2009). However, in most cases the selective agents and the traits contributing to local adaptation are poorly known (Wade and Kalisz 1990, MacColl 2011, Caruso et al. 2017). Selection and adaptive differentiation can act through several different components of fitness and at different stages in the life-history of an organ- ism (Mojica and Kelly 2010, Postma and Ågren 2016). Many studies have quantified selection and adaptation based on estimates of survival, mating success and fecundity, while few attempted measuring selection through life- time fitness by including also the earliest stages of the life cycle (germina- tion and seedling establishment; McGraw and Antonovics 1983, Schemske 1984, Campbell 1998, Postma and Ågren 2016). Studying the mechanisms shaping and maintaining among-population variation in natural populations provides essential insights to the processes of divergence on a micro- evolutionary scale and contributes to our understanding of macro- evolutionary patterns. In my thesis, I combine studies of phenotypic selection with experimental manipulations of traits and selection environment in natural populations of animal-pollinated plants to examine the functional and adaptive significance of differences in trait expression and mechanisms responsible for current selection. In the paragraphs below, I present the aims and
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