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A Geography of Lake Carbon Cycling Limnology and Oceanography Letters, 3: 49-56 http://www.diva-portal.org This is the published version of a paper published in . Citation for the original published paper (version of record): Seekell, D A., Lapierre, J., Cheruvelil, K S. (2018) A geography of lake carbon cycling Limnology and Oceanography Letters, 3: 49-56 https://doi.org/10.1002/lol2.10078 Access to the published version may require subscription. N.B. When citing this work, cite the original published paper. Permanent link to this version: http://urn.kb.se/resolve?urn=urn:nbn:se:umu:diva-154436 Limnology and Oceanography Letters 3, 2018, 49–56 VC 2018 The Author. Limnology and Oceanography Letters published by Wiley Periodicals, Inc. on behalf of Association for the Sciences of Limnology and Oceanography doi: 10.1002/lol2.10078 SPECIAL ISSUE-ESSAY A geography of lake carbon cycling David A. Seekell ,1,2 Jean-Franc¸ois Lapierre ,3,4* Kendra Spence Cheruvelil 5,6 1Department of Ecology and Environmental Science, Umea˚ University, Umea˚, Sweden; 2Climate Impacts Research Centre, Umea˚ University, Umea˚, Sweden; 3Departement de sciences biologiques, Universite de Montreal, Montreal, Quebec, Canada; 4Groupe de Recherche Interuniversitaire en Limnologie et en Environnement Aquatique (GRIL); 5Department of Fisheries and Wildlife, Michigan State University, East Lansing, Michigan; 6Lyman Briggs College, Michigan State University, East Lansing, Michigan Scientific Significance Statement Carbon cycling in lakes is highly variable among lakes within regions, and across regions and continents, but the underly- ing causes of this variation among lakes and regions are not well understood. In this essay, we propose two main mecha- nisms that operate at the regional scale and contribute to broad-scale interlake variation in carbon cycling. This essay sets the foundation for a geographic understanding of lake carbon cycling, which facilitates developing testable hypotheses to improve estimates of the role of inland waters in global elemental cycles. Lake carbon cycling is influenced by external factors con- climate, land use/land cover) as compared to that of individ- trolling the transfer of mass (e.g. carbon, nutrient) and energy ual lake ecosystems (100’s–1000’s km2 vs < 1km2;Lapierre (e.g. light, heat) on lake processes (e.g. primary production, et al. 2015; Cael and Seekell 2016). Historically, broad-scale respiration, greenhouse gases emissions), either directly or studies of lake carbon cycling exceeded the logistic abilities of through the terrestrial landscape (Leavitt et al. 2009; Lapierre individual lab groups and were outside the mandate of most et al. 2015). Although these transfers of mass and energy governmental lake monitoring programs. However, publicly should affect all lakes similarly, studies of lake carbon cycling accessible geographic and limnological data originating from conducted in regions with contrasting climate, land cover, individual studies and from data synthesis efforts are increas- and land use suggest that how strongly limnological first ingly available for broad extents across wide environmental principles are expressed, and their relative importance to eco- gradients to develop understanding of the geographic pat- system functions, vary geographically. This fact may be due terns in lake carbon cycling (e.g., Soranno et al. 2017). to the mismatch in spatial scales that characterize the geo- Landscape- and global-scale limnology have received graphic factors behind these contrasting patterns (e.g. increased attention in recent years. For example, landscape *Correspondence: [email protected] Author Contribution Statement: DAS and JFL are co-leads of this paper; they developed the conceptual framework and wrote large sections of the essay. KSC contributed ideas, feedback, and text throughout the manuscript development, writing, and revising process. All authors read and approved the paper. Data Availability Statement: Data are available at the following repositories or tables in published articles: https://figshare.com/articles/Latitudinal_gra- dients_in_phosphorus_and_vegetation_biomass_for_Sweden_and_Denmark/5614588, https://figshare.com/articles/Institutions_studying_lake_carbon_ cycling/5612968, http://doi.org/10.7927/H40Z715X, http://doi.org/10.1029/2006JD007377, http://doi.org/10.7927/H4NP22DQ, and Imhoff and Bou- noua 2006. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. This article is an invited paper to the Special Issue: Carbon cycling in inland waters Edited by: Emily Stanley and Paul del Giorgio 49 Seekell et al. Geography of lake carbon cycling limnology has quantified and explained variation in ecosys- 2015a). Therefore, even the most basic relationships between tem states and lake responses to multi-scaled pressures (e.g. carbon, nutrients, and lake ecosystem processes may show Soranno et al. 2010) and global-scale limnology has devel- unexpected patterns depending on where the study was con- oped global rate estimates for carbon exchange between sur- ducted. We argue that these conflicting results may be due face waters and the atmosphere that can be compared with to regional-scale differences in geographic context, which other aspects of global elemental cycles (Downing 2009). drives broad-scale heterogeneity in patterns of lake carbon Less well-known are the fundamental limnological patterns cycling. at an intermediate, regional scale, where non-linear and Landscape limnology sets the context for resolving con- threshold relationships between lake processes and their in- flicting findings among studies conducted in different areas lake and landscape drivers often emerge and have major of the globe. This subdiscipline of limnology has formalized implications for broad-scale understanding of lake carbon the importance of the regional scale (e.g., Cheruvelil et al. cycling. Moreover, the mechanisms responsible for the pres- 2013) and the concept of cross-scale interactions (which ence of geographically varying relationships in aquatic eco- occur when the drivers interact across spatial scales or study systems remain unknown. Therefore, we argue that such regions (Soranno et al. 2014) for improved understanding of geographically varying relationships could have profound aquatic ecosystems. However, the examples that have been implications for broad scale understanding of lake carbon so far have not included carbon, nor have they described cycling, and that these implications have seldom been how geographic context may alter fundamental ecosystem explored. functions. We propose two regional-scaled mechanisms that In this essay, we propose a geographic framework for can lead to such geographic variation in lake carbon cycling. studying lake carbon cycling that can connect within- First, different regions may have a DOC frequency distribu- ecosystem carbon processes with their broader-scaled drivers, tion that fall on either side of a threshold relationship including climate, land cover, and human activities. We between DOC and an aquatic process (Fig. 1a). In this con- present two regional-scale mechanisms related to thresholds text we define threshold as a point across which there is a that we hypothesize can give rise to geographically varying change in the direction of a relationship or a fundamental relationships between lake carbon cycling and its drivers, shift in ecosystem function in response to a driver. For which can account for the often contrasting results from the example, a threshold may be observed as a ratio shifting literature. Finally, we present a way to conceptualize and from above to below 1 or as a net rate shifting from a posi- delineate testable, hypothetical regions of lake carbon tive to a negative value (e.g. P : R ratio or net ecosystem pro- cycling that could account for the above contrasting rela- duction; blue line on Fig. 1), or as a unimodal relationship tionships and help to synthesize studies of carbon in widely whereby the direction of a trend shifts with changing DOC different regions. concentration (red line in Fig. 1). Second, threshold relation- ships may occur when geographic gradients intersect and The role of geographic context in lake carbon alter the balance between production, delivery, and process- cycling ing of carbon and nutrients to and within lakes. This phe- There are many variables driving the processes involved nomenon may be particularly relevant when natural and in lake carbon cycling, but a few of them play a central role. human gradients intersect (Fig. 1b). Within our geographic Along with phosphorus concentration and lake size, dis- framework of lake carbon cycling, we discuss examples based solved organic carbon (DOC) concentration is commonly on relationships of key lake processes with DOC and used to compare, predict, and generalize patterns of lake car- nutrients, and argue that these ideas may be applicable for bon cycling (Williamson et al. 1999; Prairie 2008). DOC is other limnological patterns and processes. the largest carbon pool in the water column and it strongly Mechanism 1: Threshold relationships explained by influences diverse physical, chemical, and biological charac- regional heterogeneity in dissolved organic carbon teristics (Prairie 2008). In particular, DOC is colored and has concentrations a primary role in limiting light (Carpenter et al. 1998; Wil- Lakes often have nonlinear relationships with drivers that liamson et al.
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