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. 2019 Mar 26;116(13):6205-6210.
doi: 10.1073/pnas.1803122116. Epub 2019 Mar 8.

Applying modern coexistence theory to priority effects

Affiliations

Applying modern coexistence theory to priority effects

Tess Nahanni Grainger et al. Proc Natl Acad Sci U S A. .

Abstract

Modern coexistence theory is increasingly used to explain how differences between competing species lead to coexistence versus competitive exclusion. Although research testing this theory has focused on deterministic cases of competitive exclusion, in which the same species always wins, mounting evidence suggests that competitive exclusion is often historically contingent, such that whichever species happens to arrive first excludes the other. Coexistence theory predicts that historically contingent exclusion, known as priority effects, will occur when large destabilizing differences (positive frequency-dependent growth rates of competitors), combined with small fitness differences (differences in competitors' intrinsic growth rates and sensitivity to competition), create conditions under which neither species can invade an established population of its competitor. Here we extend the empirical application of modern coexistence theory to determine the conditions that promote priority effects. We conducted pairwise invasion tests with four strains of nectar-colonizing yeasts to determine how the destabilizing and fitness differences that drive priority effects are altered by two abiotic factors characterizing the nectar environment: sugar concentration and pH. We found that higher sugar concentrations increased the likelihood of priority effects by reducing fitness differences between competing species. In contrast, higher pH did not change the likelihood of priority effects, but instead made competition more neutral by bringing both fitness differences and destabilizing differences closer to zero. This study demonstrates how the empirical partitioning of priority effects into fitness and destabilizing components can elucidate the pathways through which environmental conditions shape competitive interactions.

Keywords: competition; fitness difference; invasion criterion; niche difference; stabilizing difference.

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Conflict of interest statement

The authors declare no conflict of interest.

Figures

Fig. 1.
Fig. 1.
Conceptual framework demonstrating how shifts in fitness differences and stabilizing differences across an environmental gradient alter competitive outcomes and the neutrality of competitive interactions. Circles represent a pair of species competing in three different environments, and squares represent a different pair of species competing in the same three environments. The solid arrowed line shows how positively correlated changes in fitness and stabilizing differences alter the outcome of competition (priority effects vs. competitive exclusion). The dashed arrowed line shows how negatively correlated changes in fitness and stabilizing differences change the neutrality of species interactions (distance to black star). The black star indicates the special case of neutrality in which competing species have equal fitness and no stabilizing differences (fitness difference = 1 and stabilizing difference = 0).
Fig. 2.
Fig. 2.
The effect of pH and sugar on (A) fitness differences and stabilizing differences, (B) the likelihood of priority effects vs. competitive exclusion, and (C) the neutrality of competitive interactions. Points in (A) show fitness differences and stabilizing differences for six species pairs competing in six nectar environments (two levels of sugar × three levels of pH). Four pairs were removed from this analysis because one species in each of these pairs had negative growth in monoculture (Methods). Error bars are 95% confidence intervals, so that error bars falling completely within the priority effects or exclusion zone indicate a significant outcome. Colored lines in B and C show mean values for each environment, calculated from points in A. Colored lines in B show the mean distance that species pairs competing in each nectar environment fall from the line that separates priority effects from competitive exclusion; lines below the black line indicate a greater likelihood of priority effects, and lines above the black line indicate a greater likelihood of competitive exclusion. Arcs in C show the mean distance that species pairs competing in each nectar environment fall from total neutrality (the special case of no fitness or stabilizing differences, indicated by the black star). Arcs close to the star indicate more neutral interactions, and arcs far from the star indicate less neutral interactions.
Fig. 3.
Fig. 3.
The effect of pH and sugar on (A) fitness differences and (B) stabilizing differences. Points show mean values in each environment, averaged across six species pairs, and error bars show 1 SE from the mean. Low sugar is shown in light colors (pink, light purple, and light blue) connected by a light gray line, and high sugar treatments are shown in dark colors (red, dark purple, and dark blue) connected by a dark gray line. The dashed lines at zero indicate the absence of fitness differences (A; note the log transformation) or stabilizing differences (B).

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