In this paper, we present a conceptual framework for investigating ecological patterns and processes at regional to continental scales. Ecological phenomena operate across a range of scales (Figure 1), but the development of ecological theory of regions to continents lags behind that of finer scales. Better understanding of broad scales is needed because these are the extents over which many environmental problems have their causes and consequences. Our framework incorporates existing theories from other ecological subdisciplines and environmental disciplines, to promote broad-scale ecology as more general, integrative, and predictive.We define "macroscales" as regional to continental extents with distances spanning hundreds to thousands of kilometers (ie larger than landscapes; Urban et al. 1987). "Components" at these spatial scales (Figure 2) are biological (eg species, populations, communities), geophysical (eg climate, physiography, hydrology, geochemistry), and social (eg political systems, economies, cultures), and can span timescales ranging from days to millennia. When interacting with one another and with phenomena at other spatial or temporal scales, these components constitute a "macrosystem"; macrosystems ecology (MSE) is the study of such extensive and multiscaled systems. This perspective treats patterns and processes as dynamic and interactive, both within and across scales of time and space.n MotivationsThe emergence of MSE has been driven by three main factors: pressing societal needs for ecological predictions at these wider scales; the increasing focus on mechanistic studies that cover broad extents across a range of ecological subdisciplines; and a wealth of new methodological and technological capabilities that enable scientists to carry out such studies. These three interrelated issues will continue to shape the development of MSE.Ecologists are increasingly asked to address environmental problems and policies with causes and consequences that operate over broad extents (Clark et al. 2001;Peters et al. 2011;Liu et al. 2013). For example, scientists and policy makers are unsure how climate and land-use changes will influence the provision of multiple ecosystem services, at both local and regional scales (Qiu Macrosystems ecology is the study of diverse ecological phenomena at the scale of regions to continents and their interactions with phenomena at other scales. This emerging subdiscipline addresses ecological questions and environmental problems at these broad scales. Here, we describe this new field, show how it relates to modern ecological study, and highlight opportunities that stem from taking a macrosystems perspective. We present a hierarchical framework for investigating macrosystems at any level of ecological organization and in relation to broader and finer scales. Building on well-established theory and concepts from other subdisciplines of ecology, we identify feedbacks, linkages among distant regions, and interactions that cross scales of space and time as the most likely sou...

Abstract. Riparian zones are habitats of critical conservation concern worldwide, as they are known to filter agricultural contaminants, buffer landscapes against erosion, and provide habitat for high numbers of species. Here we test the generality of the notion that riparian habitats harbor more species than adjacent upland habitats. Using previously published data collected from seven continents and including taxa ranging from Antarctic soil invertebrates to tropical rain forest lianas and primates, we show that riparian habitats do not harbor higher numbers of species, but rather support significantly different species pools altogether. In this way, riparian habitats increase regional (␥-) richness across the globe by Ͼ50%, on average. Thus conservation planners can easily increase the number of species protected in a regional portfolio by simply including a river within terrestrial biodiversity reserves. Our analysis also suggests numerous possible improvements for future studies of species richness gradients across riparian and upland habitats. First, Ͻ15% of the studies in our analysis included estimates of more than one taxonomic group of interest. Second, within a given taxonomic group, studies employed variable methodologies and sampling areas in pursuit of richness and turnover estimates. Future analyses of species richness patterns in watersheds should aim to include a more comprehensive suite of taxonomic groups and should measure richness at multiple spatial scales.

As the permafrost region warms, its large organic carbon pool will be increasingly vulnerable to decomposition, combustion, and hydrologic export. Models predict that some portion of this release will be offset by increased production of Arctic and boreal biomass; however, the lack of robust estimates of net carbon balance increases the risk of further overshooting international emissions targets. Precise empirical or model-based assessments of the critical factors driving carbon balance are unlikely in the near future, so to address this gap, we present estimates from 98 permafrost-region experts of the response of biomass, wildfire, and hydrologic carbon flux to climate change. Results suggest that contrary to model projections, total permafrost-region biomass could decrease due to water stress and disturbance, factors that are not adequately incorporated in current models. Assessments indicate that end-of-the-century organic carbon release from Arctic rivers and collapsing coastlines could increase by 75% while carbon loss via burning could increase four-fold. Experts identified water balance, shifts in vegetation community, and permafrost degradation as the key sources of uncertainty in predicting future system response. In combination with previous findings, results suggest the permafrost region will become a carbon source to the atmosphere by 2100 regardless of warming scenario but that 65%-85% of permafrost carbon release can still be avoided if human emissions are actively reduced.

Nitrate (NO3 (-) ) export coupled with high inorganic nitrogen (N) concentrations in Alaskan streams suggests that N cycles of permafrost-influenced ecosystems are more open than expected for N-limited ecosystems. We tested the hypothesis that soil thaw depth governs inorganic N retention and removal in soils due to vertical patterns in the dominant N transformation pathways. Using an in situ, push-pull method, we estimated rates of inorganic N uptake and denitrification during snow melt, summer, and autumn, as depth of soil-stream flowpaths increased in the valley bottom of an arctic and a boreal catchment. Net NO3 (-) uptake declined sharply from snow melt to summer and decreased as a nonlinear function of thaw depth. Peak denitrification rate occurred during snow melt at the arctic site, in summer at the boreal site, and declined as a nonlinear function of thaw depth across both sites. Seasonal patterns in ammonium (NH4 (+) ) uptake were not significant, but low rates during the peak growing season suggest uptake that is balanced by mineralization. Despite rapid rates of hydrologic transport during snow melt runoff, rates of uptake and removal of inorganic N tended to exceed water residence time during snow melt, indicating potential for retention of N in valley bottom soils when flowpaths are shallow. Decreased reaction rates relative to water residence time in subsequent seasons suggest greater export of inorganic N as the soil-stream flowpath deepens due to thawing soils. Using seasonal thaw as a proxy for longer term deepening of the thaw layer caused by climate warming and permafrost degradation, these results suggest increasing potential for export of inorganic N from permafrost-influenced soils to streams.