How is trait diversity in a community apportioned between and within coevolving species? Disruptive selection may result in either a few species with large intraspecific trait variation (ITV) or many species with different mean traits but little ITV. Similar questions arise in spatially structured communities: heterogeneous environments could result in either a few species that exhibit local adaptation or many species with different mean traits but little local adaptation. To date, theory has been well-equipped to either include ITV or to dynamically determine the number of coexisting species, but not both. Here, we devise a theoretical framework that combines these facets and apply it to the above questions of how trait variation is apportioned within and between species in unstructured and structured populations, using two simple models of Lotka-Volterra competition. For unstructured communities, we find that as the breadth of the resource spectrum increases, ITV goes from being unimportant to crucial for characterizing the community. For spatially structured communities on two patches, we find no local adaptation, symmetric local adaptation, or asymmetric local adaptation, depending on how much the patches differ. Our framework provides a general approach to incorporate ITV in models of eco-evolutionary community assembly.
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Synthesizing the effects of individual‐level variation on coexistence
Intraspecific trait variation (ITV) is a widespread feature of life, but it is an open question how ITV affects between‐species coexistence. Recent theoretical studies have produced contradictory results, with ITV promoting coexistence in some models and undermining coexistence in others. Here we review recent work and propose a new conceptual framework to explain how ITV affects coexistence between two species. We propose that all traits belong to one of two categories: niche traits and hierarchical traits. Niche traits determine an individual's location on a niche axis or trade‐off axis, such that changing an individual's trait makes it perform better in some circumstances and worse in others. Hierarchical traits represent cases where conspecifics with different traits have the same niche, but one performs better under all circumstances, such that there are winners and losers. Our framework makes predictions for how intraspecific variation in each type of trait affects coexistence by altering stabilizing mechanisms and fitness differences. For example, ITV in niche traits generally weakens the stabilizing mechanism, except when it generates a generalist–specialist trade‐off. On the other hand, hierarchical traits tend to impact competitors differently, such that ITV in one species will strengthen the stabilizing mechanism while ITV in the other species will weaken the mechanism. We re‐examine 10 studies on ITV and coexistence, along with four novel models, and show that our framework can explain why ITV promotes coexistence in some models and undermines coexistence in others. Overall, our framework reconciles what were previously considered to be contrasting results and provides both theoretical and empirical directions to study the effect of ITV on species coexistence.
more » « less- Award ID(s):
- 1754012
- PAR ID:
- 10362423
- Publisher / Repository:
- Wiley Blackwell (John Wiley & Sons)
- Date Published:
- Journal Name:
- Ecological Monographs
- Volume:
- 92
- Issue:
- 1
- ISSN:
- 0012-9615
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract Coexistence of species requires equalizing mechanisms that minimize fitness differences, which are balanced by stabilizing mechanisms that enhance negative intraspecific interactions versus interspecific ones. Here, we develop a simple theoretical framework that allows measuring the relative strength of intraspecific versus interspecific competition in dominance hierarchies. We use it to evaluate mechanisms promoting coexistence between two congeneric charr that compete for foraging positions, which strongly influence density‐dependent growth and survival.
Agonistic interactions (
n = 761) among 71 Dolly VardenSalvelinus malma and whitespotted charrSalvelinus leucomaenis were measured by snorkelling in two pools in the sympatric zone of a Hokkaido stream during two summers. Interspecific dominance hierarchies, analysed using three methods, were closely correlated with fish length but the species treated each other equally. Ranks for the most dominant fish in each pool, determined directly by knockout experiments, were also virtually identical to ranks by length.Similarly, exponential random graph modelling of the social networks provided no evidence that either species was dominant over the other. Instead, larger fish were more likely to win contests, especially over fish of the next lower ranks.
These results demonstrated that the two species were nearly ecological equivalents in accessing key resources in this sympatric zone. Nearly identical growth and stable densities over 4 years further supported this inference, although Dolly Varden were a minority (29% of the assemblage), a sign of some fitness difference.
Detailed foraging observations coupled with two concurrent studies revealed an effective stabilizing mechanism. Dolly Varden shifted to feeding directly from the benthos when drifting invertebrates declined, a behaviour enhanced by morphological character displacement, thereby partitioning food resources and enhancing intraspecific competition while avoiding agonistic encounters with whitespotted charr.
The plurality of evidence indicates that fitness differences between these ecologically equivalent species are small in this local assemblage, and balanced by resource partitioning, a modest stabilizing mechanism that promotes coexistence. The theoretical framework presented here is a useful tool to evaluate the strength of interspecific versus intraspecific competition, which combined with information on trade‐offs in ecological performance can contribute to a mechanistic understanding of species coexistence.
