Elevational shifts in reproductive ecology indicate the climate response of a model chasmophyte, Rainer's bellflower (Campanula raineri).

Abstract

Elevation gradients provide 'natural experiments' for investigating plant climate change responses, advantageous for the study of protected species and life forms for which transplantation experiments are illegal or unfeasible, such as chasmophytes with perennial rhizomes pervading rock fissures. Elevational climatic differences impact mountain plant reproductive traits (pollen and seed quality, sexual vs. vegetative investment) and pollinator community composition; we investigated the reproductive ecology of a model chasmophyte, Campanula raineri Perp. (Campanulaceae), throughout its current elevational/climatic range to understand where sub-optimal conditions jeopardise survival. We hypothesised that: 1) reproductive fitness measures are positively correlated with elevation, indicative of the relationship between fitness and climate; 2) C. raineri, like other campanulas, is pollinated mainly by Hymenoptera; 3) potential pollinators shift with elevation. We measured pollen and seed quality, seed production, the relative investment in sexual vs. vegetative structures and vegetative (Grime's CSR) strategies at different elevations. Potential pollinators were assessed by combining molecular and morphological identification. Whereas CSR strategies were not linked to elevation, pollen and seed quality were positively correlated, as was seed production per fruit (Hypothesis 1 is supported). The main pollinators of C. raineri were Apidae, Andrenidae, Halictidae (Hymenoptera) and Syrphidae (Diptera), probably complemented by a range of occasional pollinators and visitors (Hypothesis 2 partially supported). Potential pollinator communities showed a taxonomic shift towards Diptera with elevation (particularly Anthomyiidae and Muscidae) and away from Hymenoptera (Hypothesis 3 was supported). Pollinator availability is maintained at all elevations by taxon replacement. However, reduced pollen quality and seed production at lower elevations suggest an impact of climate change on reproduction (especially <1200 m a.s.l., where seed germination was limited). Aside from guiding targeted conservation actions for C. raineri, our results highlight problems that may be common to mountain chasmophytes worldwide.