Although the concept of biodiversity emerged 30 years ago, patterns and processes influencing ecological diversity have been studied for more than a century. Historically, ecological processes tended to be considered as occurring in local habitats that were spatially homogeneous and temporally at equilibrium. Initially considered as a constraint to be avoided in ecological studies, spatial heterogeneity was progressively recognized as critical for biodiversity. This resulted, in the 1970s, in the emergence of a new discipline, landscape ecology, whose major goal is to understand how spatial and temporal heterogeneity influence biodiversity. To achieve this goal, researchers came to realize that a fundamental issue revolves around how they choose to conceptualize and measure heterogeneity. Indeed, observed landscape patterns and their apparent relationship with biodiversity often depend on the scale of observation and the model used to describe the landscape. Due to the strong influence of island biogeography, landscape ecology has focused primarily on spatial heterogeneity. Several landscape models were conceptualized, allowing for the prediction and testing of distinct but complementary effects of landscape heterogeneity on species diversity. We now have ample empirical evidence that patch structure, patch context, and mosaic heterogeneity all influence biodiversity. More recently, the increasing recognition of the role of temporal scale has led to the development of new conceptual frameworks acknowledging that landscapes are not only heterogeneous but also dynamic. The current challenge remains to truly integrate both spatial and temporal heterogeneity in studies on biodiversity. This integration is even more challenging when considering that biodiversity often responds to environmental changes with considerable time lags, and multiple drivers of global changes are interacting, resulting in non-additive and sometimes antagonistic effects. Recent technological advances in remote sensing, the availability of massive amounts of data, and long-term studies represent, however, very promising avenues to improve our understanding of how spatial and temporal heterogeneity influence biodiversity.
Juha Merilä and Ary A. Hoffmann
Changing climatic conditions have both direct and indirect influences on abiotic and biotic processes and represent a potent source of novel selection pressures for adaptive evolution. In addition, climate change can impact evolution by altering patterns of hybridization, changing population size, and altering patterns of gene flow in landscapes. Given that scientific evidence for rapid evolutionary adaptation to spatial variation in abiotic and biotic environmental conditions—analogous to that seen in changes brought by climate change—is ubiquitous, ongoing climate change is expected to have large and widespread evolutionary impacts on wild populations. However, phenotypic plasticity, migration, and various kinds of genetic and ecological constraints can preclude organisms from evolving much in response to climate change, and generalizations about the rate and magnitude of expected responses are difficult to make for a number of reasons.
First, the study of microevolutionary responses to climate change is a young field of investigation. While interest in evolutionary impacts of climate change goes back to early macroevolutionary (paleontological) studies focused on prehistoric climate changes, microevolutionary studies started only in the late 1980s. The discipline gained real momentum in the 2000s after the concept of climate change became of interest to the general public and funding organizations. As such, no general conclusions have yet emerged. Second, the complexity of biotic changes triggered by novel climatic conditions renders predictions about patterns and strength of natural selection difficult. Third, predictions are complicated also because the expression of genetic variability in traits of ecological importance varies with environmental conditions, affecting expected responses to climate-mediated selection.
There are now several examples where organisms have evolved in response to selection pressures associated with climate change, including changes in the timing of life history events and in the ability to tolerate abiotic and biotic stresses arising from climate change. However, there are also many examples where expected selection responses have not been detected. This may be partly explainable by methodological difficulties involved with detecting genetic changes, but also by various processes constraining evolution.
There are concerns that the rates of environmental changes are too fast to allow many, especially large and long-lived, organisms to maintain adaptedness. Theoretical studies suggest that maximal sustainable rates of evolutionary change are on the order of 0.1 haldanes (i.e., phenotypic standard deviations per generation) or less, whereas the rates expected under current climate change projections will often require faster adaptation. Hence, widespread maladaptation and extinctions are expected. These concerns are compounded by the expectation that the amount of genetic variation harbored by populations and available for selection will be reduced by habitat destruction and fragmentation caused by human activities, although in some cases this may be countered by hybridization. Rates of adaptation will also depend on patterns of gene flow and the steepness of climatic gradients. Theoretical studies also suggest that phenotypic plasticity (i.e., nongenetic phenotypic changes) can affect evolutionary genetic changes, but relevant empirical evidence is still scarce. While all of these factors point to a high level of uncertainty around evolutionary changes, it is nevertheless important to consider evolutionary resilience in enhancing the ability of organisms to adapt to climate change.