Animals navigate landscapes based on perceived risks vs. rewards, as inferred from features of the landscape. In the wild, knowing how strongly animal movement is directed by landscape features is ...difficult to ascertain but widespread disturbances such as wildfires can serve as natural experiments. We tested the hypothesis that wildfires homogenize the risk/reward landscape, causing movement to become less directed, given that fires reduce landscape complexity as habitat structures (e.g., tree cover, dense brush) are burned. We used satellite imagery of a research reserve in Northern California to count and categorize paths made primarily by mule deer (Odocoileus hemionus) in grasslands. Specifically, we compared pre-wildfire (August 2014) and post-wildfire (September 2018) image history layers among locations that were or were not impacted by wildfire (i.e., a Before/After Control/Impact design). Wildfire significantly altered spatial patterns of deer movement: more new paths were gained and more old paths were lost in areas of the reserve that were impacted by wildfire; movement patterns became less directed in response to fire, suggesting that the risk/reward landscape became more homogenous, as hypothesized. We found evidence to suggest that wildfire affects deer populations at spatial scales beyond their scale of direct impact and raises the interesting possibility that deer perceive risks and rewards at different spatial scales. In conclusion, our study provides an example of how animals integrate spatial information from the environment to make movement decisions, setting the stage for future work on the broader ecological implications for populations, communities, and ecosystems, an emerging interest in ecology.
A rich body of knowledge links biodiversity to ecosystem functioning (BEF), but it is primarily focused on small scales. We review the current theory and identify six expectations for scale ...dependence in the BEF relationship: (1) a nonlinear change in the slope of the BEF relationship with spatial scale; (2) a scale‐dependent relationship between ecosystem stability and spatial extent; (3) coexistence within and among sites will result in a positive BEF relationship at larger scales; (4) temporal autocorrelation in environmental variability affects species turnover and thus the change in BEF slope with scale; (5) connectivity in metacommunities generates nonlinear BEF and stability relationships by affecting population synchrony at local and regional scales; (6) spatial scaling in food web structure and diversity will generate scale dependence in ecosystem functioning. We suggest directions for synthesis that combine approaches in metaecosystem and metacommunity ecology and integrate cross‐scale feedbacks. Tests of this theory may combine remote sensing with a generation of networked experiments that assess effects at multiple scales. We also show how anthropogenic land cover change may alter the scaling of the BEF relationship. New research on the role of scale in BEF will guide policy linking the goals of managing biodiversity and ecosystems.
We address the challenge of scale for biodiversity and ecosystem functioning (BEF) research. We review current theory and identify six expectations for scale dependence in the BEF relationship. We suggest directions for synthesis that combine theoretical and empirical methods and suggest their application to human transformed landscapes.
AbstractCompetition drives evolutionary change across taxa, but our understanding of how competitive differences among species directs the evolution of interspecific interactions remains incomplete. ...Verbal models assume that interspecific competition will select for reducing a species' sensitivity to competition with their opponent; however, they do not consider the potential for other demographic components of competitive ability to evolve, specifically, interspecific effects, intraspecific interactions, and intrinsic growth rates. To better understand how competitive ability evolves, we set out to explore how each component has evolved and whether their evolution has been constrained by trade-offs. By setting sympatric and allopatric populations of an annual grass in competition with a dominant invader, we demonstrate (1) that in response to interspecific competition, populations can evolve increased competitive ability through either reduced interspecific or, surprisingly, reduced intraspecific competition; (2) that trade-offs do not always constrain the evolution of competitive ability but rather that parameters may correlate in ways that mutually beget higher competitive ability; and (3) that the evolution of one species can influence the competitive ability of its opponent, a consequence of how competitive ability is defined ecologically. Overall, our results reveal the complexity with which demographic components evolve in response to interspecific competition and the impact past evolution can have on present-day interactions.
‘Filtering’, or the reduction in species diversity that occurs because not all species can persist in all locations, is thought to unfold hierarchically, controlled by the environment at large scales ...and competition at small scales. However, the ecological effects of competition and the environment are not independent, and observational approaches preclude investigation into their interplay. We use a demographic approach with 30 plant species to experimentally test: (i) the effect of competition on species persistence in two soil moisture environments, and (ii) the effect of environmental conditions on mechanisms underlying competitive coexistence. We find that competitors cause differential species persistence across environments even when effects are lacking in the absence of competition, and that the traits which determine persistence depend on the competitive environment. If our study had been observational and trait-based, we would have erroneously concluded that the environment filters species with low biomass, shallow roots and small seeds. Changing environmental conditions generated idiosyncratic effects on coexistence outcomes, increasing competitive exclusion of some species while promoting coexistence of others. Our results highlight the importance of considering environmental filtering in the light of, rather than in isolation from, competition, and challenge community assembly models and approaches to projecting future species distributions.
Evolutionary biologists since Darwin have hypothesized that closely related species compete more intensely and are therefore less likely to coexist. However, recent theory posits that species diverge ...in two ways: either through the evolution of ‘stabilizing differences’ that promote coexistence by causing individuals to compete more strongly with conspecifics than individuals of other species, or through the evolution of ‘fitness differences’ that cause species to differ in competitive ability and lead to exclusion of the weaker competitor. We tested macroevolutionary patterns of divergence by competing pairs of annual plant species that differ in their phylogenetic relationships, and in whether they have historically occurred in the same region or different regions (sympatric versus allopatric occurrence). For sympatrically occurring species pairs, stabilizing differences rapidly increased with phylogenetic distance. However, fitness differences also increased with phylogenetic distance, resulting in coexistence outcomes that were unpredictable based on phylogenetic relationships. For allopatric species, stabilizing differences showed no trend with phylogenetic distance, whereas fitness differences increased, causing coexistence to become less likely among distant relatives. Our results illustrate the role of species' historical interactions in shaping how phylogenetic relationships structure competitive dynamics, and offer an explanation for the evolution of invasion potential of non-native species.
Character displacement is one of the most studied phenomena in evolutionary biology, yet research has narrowly focused on demonstrating whether or not displacement has occurred. We propose a new ...experimental approach, adopted from the coexistence literature, that directly measures interspecific competition among sympatric and allopatric populations of species. Doing so allows increased ability to (i) test predictions of character displacement without biases inherent to character-centric tests, (ii) quantify its effect on the stability of coexistence, (iii) resolve the phenotypic pathways through which competitive divergence is achieved, and (iv) perform comparative tests. Our approach extends research to forms of character displacement not readily identified by past methods and will lead to a broader understanding of its consequences for community structure.
Recent evaluations of tests of character displacement suggest major gaps in our understanding.
We advocate for adopting an experimental approach from the ecological literature to allow researchers to test long-standing theory about when, where, and how character displacement occurs.
The approach adopts an invasibility criterion, and in practice is assessed by measuring the population growth rate of a species when rare competing against a background of individuals of another species from either sympatric or allopatric populations.
We show how adopting this approach will provide unbiased demonstrations of whether character displacement is present, as well as reveal its evolutionary underpinnings and ecological consequences.
Ecology is a science of scale, which guides our description of both ecological processes and patterns, but we lack a systematic understanding of how process scale and pattern scale are connected. ...Recent calls for synthesis between population ecology, community ecology, and ecosystem ecology motivate the integration of phenomena at multiple organizational levels. Furthermore, many studies leave out the scaling of a critical process: species interactions, which may be non‐local through movement or foraging and must be distinguished from dispersal scales. Here, we use simulations to explore the consequences of three different process scales (species interactions, dispersal, and the environment) on emergent patterns of biodiversity, ecosystem functioning, and their relationship, in a spatially‐explicit landscape and stable equilibrium setting. A major result of our study is that the spatial scales of dispersal and species interactions have opposite effects: a larger dispersal scale homogenizes spatial biomass patterns, while a larger interaction scale amplifies their heterogeneity. Interestingly, the specific scale at which dispersal and interaction scales begin to influence landscape patterns depends on the scale of environmental heterogeneity – in other words, the scale of one process allows important scales to emerge in other processes. This interplay between process scales, i.e. a situation where no single process dominates, can only occur when the environment is heterogeneous and the scale of dispersal small. Finally, contrary to our expectations, we observe that the spatial scale of ecological processes is more clearly reflected in landscape patterns (i.e. distribution of local outcomes) than in global patterns such as species–area relationships (SARs) or large‐scale biodiversity–functioning relationships. Overall we conclude that long‐range interactions often act differently and even in opposite ways to dispersal, and that the landscape patterns that emerge from the interplay of long‐ranged interactions, dispersal and environmental heterogeneity are not well captured by often‐used metrics like the SAR.
Ecological theory posits that dispersal among habitat patches links local communities and is a key “regional” process that maintains biological diversity. However, manipulations required to ...experimentally test regional processes are infeasible for most systems, and thus more work is needed to detect the scales at which regional processes manifest and their overall effect on diversity. In a Californian grassland, a hotspot for global biodiversity, we used a seed vacuum to increase dispersal at spatial scales varying from1mto 10 km while maintaining a realistic spatial structure of species pools and environmental conditions. We found that dispersal limitation has a profound influence on diversity; species richness increased with the spatial scale of seed mixing, doubling in plots that received seed from large (≥5 km) compared with small (≤5 m) scales. This increase in diversity corresponded to an increase in how well species distributions were explained by environmental conditions, from modest at small scales (R² = 0.34) to strong at large scales (R² = 0.52). Responses to the spatial scale of seed mixing were nonlinear, with no differences below 5 m or above 5 km. Nonlinearities were explained by homogeneity of environmental conditions below 5 m and by a lack of additional variation in the species pool above 5 km. Our approach of manipulating natural communities at different spatial scales reveals (i) nonlinear transitions in the importance of environmental sorting and dispersal, and (ii) the negative effects of dispersal limitation on local diversity, consistent with previous research suggesting that large numbers of species are headed toward regional extinction.
ABSTRACT
Biological insurance theory predicts that, in a variable environment, aggregate ecosystem properties will vary less in more diverse communities because declines in the performance or ...abundance of some species or phenotypes will be offset, at least partly, by smoother declines or increases in others. During the past two decades, ecology has accumulated strong evidence for the stabilising effect of biodiversity on ecosystem functioning. As biological insurance is reaching the stage of a mature theory, it is critical to revisit and clarify its conceptual foundations to guide future developments, applications and measurements. In this review, we first clarify the connections between the insurance and portfolio concepts that have been used in ecology and the economic concepts that inspired them. Doing so points to gaps and mismatches between ecology and economics that could be filled profitably by new theoretical developments and new management applications. Second, we discuss some fundamental issues in biological insurance theory that have remained unnoticed so far and that emerge from some of its recent applications. In particular, we draw a clear distinction between the two effects embedded in biological insurance theory, i.e. the effects of biodiversity on the mean and variability of ecosystem properties. This distinction allows explicit consideration of trade‐offs between the mean and stability of ecosystem processes and services. We also review applications of biological insurance theory in ecosystem management. Finally, we provide a synthetic conceptual framework that unifies the various approaches across disciplines, and we suggest new ways in which biological insurance theory could be extended to address new issues in ecology and ecosystem management. Exciting future challenges include linking the effects of biodiversity on ecosystem functioning and stability, incorporating multiple functions and feedbacks, developing new approaches to partition biodiversity effects across scales, extending biological insurance theory to complex interaction networks, and developing new applications to biodiversity and ecosystem management.
Species interact with the physical world in complex ways, and life-history strategies could cause species to differ in how they experience the connectedness of the same landscape. As a consequence, ...dispersal limitation might be present but not captured by distance-based measures of connectivity. To test these ideas, we surveyed plant communities that live on discrete patches of serpentine habitat embedded within an invaded nonserpentine habitat matrix. Species in these communities differ in dispersal mode (gravity, animal, or wind); thus we used satellite imagery to quantify landscape features that might differentially influence connectivity for some dispersal- mode groups over others (surface streams, animal paths). Our data yielded two key insights: first, dispersal limitation appeared to be absent using a conventional distance-based measure of connectivity, but emerged after considering forms of landscape connectivity relevant to each dispersal mode. Second, the landscape variables that emerged as most important to each dispersal mode were generally consistent with our predictions based on species’ putative dispersal vectors, but also included unexpected interactive effects. For example, the richness of animal-dispersed species was positively associated with animal connectivity when patches were close in space, but when patches were isolated, animals had a strong negative effect. This finding alludes to the reduced ability of animals to disperse seeds between suitable patches in invaded landscapes because of increased inter-patch distances. Real landscapes include complex spatial flows of energy and matter, which, as our work demonstrates, sets up ecological opportunity for organisms to differ in how they disperse in a common landscape.
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