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Abstract Gradient and scale are two key concepts in ecology and evolution that are closely related but inherently distinct. While scale commonly refers to the dimensional space of a specific ecological/evolutionary (eco–evo) issue, gradient measures the range of a given variable. Gradient and scale can jointly and interactively influence eco–evo patterns. Extensive previous research investigated how changing scales may affect the observation and interpretation of eco–evo patterns; however, relatively little attention has been paid to the role of changing gradients. Here, synthesizing recent research progress, we suggest that the role of scale in the emergence of ecological patterns should be evaluated in conjunction with considering the underlying environmental gradients. This is important because, in most studies, the range of the gradient is often part of its full potential range. The difference between sampled (partial) versus potential (full) environmental gradients may profoundly impact observed eco–evo patterns and alter scale–gradient relationships. Based on observations from both field and experimental studies, we illustrate the underlying features of gradients and how they may affect observed patterns, along with the linkages of these features to scales. Since sampled gradients often do not cover their full potential ranges, we discuss how the breadth and the starting and ending positions of key gradients may affect research design and data interpretation. We then outline potential approaches and related perspectives to better integrate gradient with scale in future studies. 

Abstract Grassland and other herbaceous communities cover significant portions of Earth's terrestrial surface and provide many critical services, such as carbon sequestration, wildlife habitat, and food production. Forecasts of global change impacts on these services will require predictive tools, such as process‐based dynamic vegetation models. Yet, model representation of herbaceous communities and ecosystems lags substantially behind that of tree communities and forests. The limited representation of herbaceous communities within models arises from two important knowledge gaps: first, our empirical understanding of the principles governing herbaceous vegetation dynamics is either incomplete or does not provide mechanistic information necessary to drive herbaceous community processes with models; second, current model structure and parameterization of grass and other herbaceous plant functional types limits the ability of models to predict outcomes of competition and growth for herbaceous vegetation. In this review, we provide direction for addressing these gaps by: (1) presenting a brief history of how vegetation dynamics have been developed and incorporated into earth system models, (2) reporting on a model simulation activity to evaluate current model capability to represent herbaceous vegetation dynamics and ecosystem function, and (3) detailing several ecological properties and phenomena that should be a focus for both empiricists and modelers to improve representation of herbaceous vegetation in models. Together, empiricists and modelers can improve representation of herbaceous ecosystem processes within models. In so doing, we will greatly enhance our ability to forecast future states of the earth system, which is of high importance given the rapid rate of environmental change on our planet. 

The three‐dimensional (3D) physical aspects of ecosystems are intrinsically linked to ecological processes. Here, we describe structural diversity as the volumetric capacity, physical arrangement, and identity/traits of biotic components in an ecosystem. Despite being recognized in earlier ecological studies, structural diversity has been largely overlooked due to an absence of not only a theoretical foundation but also effective measurement tools. We present a framework for conceptualizing structural diversity and suggest how to facilitate its broader incorporation into ecological theory and practice. We also discuss how the interplay of genetic and environmental factors underpin structural diversity, allowing for a potentially unique synthetic approach to explain ecosystem function. A practical approach is then proposed in which scientists can test the ecological role of structural diversity at biotic–environmental interfaces, along with examples of structural diversity research and future directions for integrating structural diversity into ecological theory and management across scales. 

As droughts become longer and more intense, impacts on terrestrial primary productivity are expected to increase progressively. Yet, some ecosystems appear to acclimate to multiyear drought, with constant or diminishing reductions in productivity as drought duration increases. We quantified the combined effects of drought duration and intensity on aboveground productivity in 74 grasslands and shrublands distributed globally. Ecosystem acclimation with multiyear drought was observed overall, except when droughts were extreme (i.e., ≤1-in-100-year likelihood of occurrence). Productivity losses after four consecutive years of extreme drought increased by ~2.5-fold compared with those of the first year. These results portend a foundational shift in ecosystem behavior if drought duration and intensity increase, from maintenance of reduced functioning over time to progressive and profound losses of productivity when droughts are extreme. 

During the 1930s Dust Bowl drought in the central United States, species with the C 3 photosynthetic pathway expanded throughout C 4 -dominated grasslands. This widespread increase in C 3 grasses during a decade of low rainfall and high temperatures is inconsistent with well-known traits of C 3 vs. C 4 pathways. Indeed, water use efficiency is generally lower, and photosynthesis is more sensitive to high temperatures in C 3 than C 4 species, consistent with the predominant distribution of C 3 grasslands in cooler environments and at higher latitudes globally. We experimentally imposed extreme drought for 4 y in mixed C 3 /C 4 grasslands in Kansas and Wyoming and, similar to Dust Bowl observations, also documented three- to fivefold increases in C 3 /C 4 biomass ratios. To explain these paradoxical responses, we first analyzed long-term climate records to show that under nominal conditions in the central United States, C 4 grasses dominate where precipitation and air temperature are strongly related (warmest months are wettest months). In contrast, C 3 grasses flourish where precipitation inputs are less strongly coupled to warm temperatures. We then show that during extreme drought years, precipitation–temperature relationships weaken, and the proportion of precipitation falling during cooler months increases. This shift in precipitation seasonality provides a mechanism for C 3 grasses to respond positively to multiyear drought, resolving the Dust Bowl paradox. Grasslands are globally important biomes and increasingly vulnerable to direct effects of climate extremes. Our findings highlight how extreme drought can indirectly alter precipitation seasonality and shift ecosystem phenology, affecting function in ways not predictable from key traits of C 3 and C 4 species.