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Kordas 2011 Journal of Experimental Marine Biology and Ecology-1.Pdf Journal of Experimental Marine Biology and Ecology 400 (2011) 218–226 Contents lists available at ScienceDirect Journal of Experimental Marine Biology and Ecology journal homepage: www.elsevier.com/locate/jembe Community ecology in a warming world: The influence of temperature on interspecific interactions in marine systems Rebecca L. Kordas a,⁎, Christopher D.G. Harley a, Mary I. O'Connor a,b a Department of Zoology, University of British Columbia, Vancouver, BC, V6T 1Z4, Canada b National Center for Ecological Analysis and Synthesis, 735 State St, Suite 300, Santa Barbara, CA 93101, United States article info abstract Keywords: Ecological patterns are determined by the interplay between abiotic factors and interactions among species. Species interaction As the Earth's climate warms, interactions such as competition, predation, and mutualism are changing due to Temperature shifts in per capita interaction strength and the relative abundance of interacting species. Changes in Climate Change Ecology interspecific relationships, in turn, can drive important local-scale changes in community dynamics, Community Ecology biodiversity, and ecosystem functioning, and can potentially alter large-scale patterns of distribution and Metabolic Ecology abundance. In many cases, the importance of indirect effects of warming, mediated by changing species interactions, will be greater—albeit less well understood—than direct effects in determining the community- and ecosystem-level outcomes of global climate change. Despite considerable community-specific idiosyncrasy, ecological theory and a growing body of data suggest that certain general trends are emerging at local scales: positive interactions tend to become more prevalent with warming, and top trophic levels are disproportionately vulnerable. In addition, important ecological changes result when the geographic overlap between species changes, and when the seasonal timing of life history events of interacting species falls into or out of synchrony. We assess the degree to which such changes are predictable, and urge advancement on several high priority questions surrounding the relationships between temperature and community ecology. An improved understanding of how assemblages of multiple, interacting species will respond to climate change is imperative if we hope to effectively prepare for and adapt to its effects. Crown Copyright © 2011 Published by Elsevier B.V. All rights reserved. Contents 1. Introduction .............................................................. 218 2. The biological importance of temperature ................................................ 219 3. Interspecific variation in thermal sensitivity ............................................... 220 4. Incorporating time and space: phenology and biogeography........................................ 222 5. The search for generality ........................................................ 223 6. Future research priorities ........................................................ 224 Acknowledgements ............................................................. 224 References ................................................................. 225 1. Introduction into organismal survival, growth, and reproduction, environmental temperature plays a large role in determining when and where Temperature is one of the most fundamental determinants of species—particularly ectothermic species—can survive and thrive biological patterns and processes. Many decades of laboratory-based (Wethey, 1983; Thomas et al., 2000; Hochachka and Somero, 2002). research have demonstrated that variation in temperature has Indeed, variation in temperature explains much of the spatial and important and easily measured effects on biochemical and physio- temporal patterns we observe in the distribution and abundance of logical rates. Because biochemical and physiological rates translate species around the world (Hutchins, 1947). Although long recognized as biologically important, environmental temperature is currently being addressed with renewed vigor as ⁎ Corresponding author. Tel.: +1 778 862 2000; fax: +1 604 822 2416. anthropogenic climate change alters patterns of mean and extreme E-mail address: [email protected] (R.L. Kordas). temperatures across the globe. Climate models suggest that the average 0022-0981/$ – see front matter. Crown Copyright © 2011 Published by Elsevier B.V. All rights reserved. doi:10.1016/j.jembe.2011.02.029 R.L. Kordas et al. / Journal of Experimental Marine Biology and Ecology 400 (2011) 218–226 219 temperature of the surface of the earth will warm by 1.7–4.4 °C by the Temperature end of the current century, with increases in mean temperatures and in the frequency and magnitude of extreme temperature events (IPCC, 2007). The magnitude of these projected changes varies from place to place (see Fig. 1). The broad-brush effects of warming are already Biochemical reaction rates observable across a wide variety of systems and taxa, with shifts in the distribution and abundance of species and the timing of life history events occurring largely as one would predict over spatial (e.g. lati- tudinal and altitudinal) and temporal (e.g., seasonal) thermal gradients (Sagarin et al., 1999; Parmesan and Yohe, 2003; Southward et al., 1995, Maintenance Maximum 2005; Helmuth et al., 2006a; Mieszkowska et al., 2007). However, not metabolic metabolic every species has responded as predicted (e.g. Hawkins et al., 2009), and rate rate for the vast majority of species little to no data on responses to temperature exist. To better understand which species are shifting and why, and the ecological impacts of temperature changes of different Metabolic magnitudes, tests of climate impacts must link processes from the scope for climatological and biophysical to the physiological and demographic to activity produce a more refined understanding of how environmental temper- ature influences body temperature and thereby the distribution and abundance of species (Helmuth, 2009). Resource Resource Resource It has long been known, however, that temperature is not the sole requirements acquisition availability determinant of where a species can live and how well it will perform. For example, Darwin (1959) recognized that many distributional patterns across thermal gradients seemed to depend more on interactions among species than upon the direct effects of temper- ature, an observation that has since received extensive observational and experimental support (Connell, 1961; MacArthur, 1972). The current theory holds that a species' response to spatial or temporal Individual variation in temperature will depend both on direct effects on the growth and individual- and population-level attributes of that species and on reproduction indirect effects mediated by changes in the distribution, abundance, and behavior of competitors, predators, parasites, and mutualists (Dunson and Travis, 1991; Davis et al., 1998; Sanford, 1999; Hawkins et al., 2009; Johnson et al., in press; Wernberg et al., in press). Thus, although general patterns of change may be robust and predictable Population (e.g. Barry et al., 1995; Parmesan and Yohe, 2003), accurate growth and predictions regarding the consequences of warming for particular size species or ecosystems of interest often remain elusive. A significant challenge in this era of global change is to improve our Fig. 2. The pathway by which temperature as a physical phenomenon influences the ecology of individuals and populations. predictive power with regards to the ecologically important conse- quences of climatic warming. To accomplish this, we must integrate single species, ecophysiological/population-level approaches and mul- can be formulated and tested, and a theory of climate change ecology tispecies, community- and ecosystem-level research into a single can progress. Here, we consider biological effects of temperature change framework so that general hypotheses regarding the effects of warming across levels of organization from enzymes to ecosystems to determine how much is known about the potential effects of temperature on complex groups of interacting species. We begin with a brief review of how temperature affects basic metabolic processes, and then explore how differences in these responses among species affect species interactions. Next, we consider how differences in physiological responses across different species can influence the overall effect of temperature on ecological communities. Finally, we outline possible frameworks for generalization of the impacts of temperature on ecological systems, and consider broader implications of these gener- alities for climate change and biogeographic patters in marine systems. We do not intend to present an exhaustive review of the ever-expanding literature on climate change. Rather, we aim to highlight the ways in which warming will influence species interactions, and the ways in which species interactions will determine the outcome of warming. 2. The biological importance of temperature Temperature is one of the most important factors affecting biological processes in poikilotherms (see Fig. 2 for a summary). The link between temperature and biological processes is kinetic; as Fig. 1. Projected surface temperature changes for the late 21st century relative to the period 1980–1999. The panels show the multi-AOGCM average projections for the A1B temperature rises and atoms become more energetic, processes such SRES scenarios averaged over 2090–2099 (IPCC, 2007). as diffusion
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