Extreme events as shaping physiology, ecology, and evolution of plants: toward a unified definition and evaluation of their consequences.

Here we consider how extreme events, particularly climatic and biotic, affect the physiology, development, ecology and evolution of organisms, focusing on plants. The marked effects on organisms are of increasing interest for ecological prediction, given the natural and anthropogenic changes in spectra of extreme events being induced by global change. Yet there is currently a paucity of knowledge or even a common world-view of how extreme events shape individuals, communities and ecosystems. We propose that extreme events need be defined in terms of organismal responses of acclimation and of de-acclimation or hysteresis. From this definition we proceed to develop a number of hypotheses, including that fitness effects of extreme events occur primarily during recovery. We review evidence that, on the evolutionary time scale, selection is virtually absent except during extreme events; these drive strong directional selection, even to trait fixation and speciation. We describe a number of new tools, both conceptual and technological, that are now at hand or that merit rapid development. Contents I. Introduction 22 II. Moving to an organismally based definition of extreme events 22 III. Features to discern in extreme events 26 IV. Additional challenges in the study of extreme events 27 V. Evolutionary dimensions 29 VI. The mandate for new conceptual tools for ecological and evolutionary prediction 34 VII. Tools in hand, and tools needed, to study extreme events 35 VIII. Conclusions 37 Acknowledgements 37 References 38.