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Past Outstanding Paper Award Recipients


The 2020 award goes to Ed Tekwa and collaborators Eli Fenichel, Simon Levin, and Malin Pinsky.

In their PNAS paper “Path-dependent institutions drive alternative stable states in conservation”, the authors use both a new theoretical model and empirical analysis of 217 managed fisheries to demonstrate that ecological and institutional factors can produce historical contingencies which determine dynamics. Their model suggests that, if institutional response rate is slow relative to the ecological response, alternative stable states are possible. Harvest rates form a path-dependent pitchfork bifurcation so that similar institutions may end up in either an overharvesting or conservation stable state. The paper therefore offers a new explanation for the different statuses of fisheries with similar management, and new approaches for switching from overharvesting to conservation.


The 2019 award goes to Bo Zhang and collaborators for their paper entitled “Carrying capacity in a heterogeneous environment with habitat connectivity” published in Ecology Letters in 2017 (20: 1118-1128).

This paper uses a delightful mixture of mathematical analysis, numerical computations, and experimental work to understand a fundamental question in spatial population biology: “How do spatial heterogeneity in environmental factors and movement patterns determine the regional abundance of a population?” The paper builds on a classical, theoretical prediction (Holt 1985 TPB) that diffusive movement into sink habitats can, counter-intuitively, increase regional equilibrium abundance. Extending this classical work, Bo Zhang and her co-authors apply a mixture of analytical and computational approaches to PDE models (and their spatially discrete counterparts) of consumer-resource interactions. Consistent with earlier theory, they demonstrate that diffusive movement of consumers can increase regional equilibrium abundance in spatially heterogeneous environments. However, contrary to predictions of earlier theoretical work, they demonstrate that lower regional equilibrium abundances are achieved in spatially heterogeneous environments than in their homogeneous counterparts. Remarkably, Bo Zhang and her collaborators empirically demonstrate both of these theoretical predictions using yeast metapopulations feeding on the amino acid tryptophan as the renewable resource. The committee and I believe this work will inspire much future work in spatial ecology at the interface of theory and experiment.

The committee also would like to recognize that the important contributions that other applicants for this prize have made to theoretical ecology. Learning about their accomplishments over the past weeks has been simultaneously inspiring and humbling.


The 2018 award goes to Colin T. Kremer and Christopher A. Klausmeier for their paper entitled “Species packing in eco-evolutionary models of seasonally fluctuating environments” published in Ecology Letters in 2017 (20: 1158-1168).

This paper examines the effects of species’ trait evolution on coexistence under environmental fluctuations and related limiting similarity patterns. It finds that many basic expectations are resilient to evolution, but also the emergence of unexpected results,
such as alternative evolutionary trajectories and stable states of the community, and that subtle properties of environmental fluctuations have a strong influence on the resulting community.

We also gave an honorable mention to Katherine Scranton and Priyanga Amarasekare, for their paper “Predicting phenological shifts in a changing climate” published in PNAS in 2017.


The 2017 winners are Andrew D. Letten, Po-Ju Ke, and Tadashi Fukami for their paper entitled “Linking modern coexistence theory and contemporary niche theory” published in Ecological Monographs (2017, 87: 161-177). This paper synthesizes the two dominant frameworks describing the niche’s role in species coexistence: modern coexistence theory and contemporary niche theory. Despite shared goals, the compatibility and complementarity of these two frameworks has received remarkably little attention. The authors demonstrate, for a general consumer-resource model, how one can translate the three criteria for species coexistence of contemporary niche theory (i.e. resource supply ratio, impact niche, and requirement niche) into the stabilizing and equalizing processes of modern coexistence theory. The paper is written in accessible prose, and draws useful connections with empirical work. By presenting the two frameworks side by side, this paper clarifies the scope and direction of existing research on these two dominant approaches to species coexistence.

We received a number of outstanding nominations for the 2017 award. Thanks to everyone who submitted nominations. The committee enjoyed reading and discussion all of the papers.


The first co-recipient is Gyorgy Barabas and co-authors for their paper entitled “Sensitivity analysis of coexistence in ecological communities: theory and application” (Ecology Letters 2014, 17:1479-1494).  This paper extends the tools of sensitivity analysis to the community level, and in doing so develops an important new perspective on the analysis of theoretical and applied models. First, it gives a method for assessing extinction risk in communities facing environmental change. Second, much like standard linear stability analysis, it can be used as part of the theoretician’s toolbox that can aid understanding model behavior better. Finally, on the broadest level, it provides a new formulation of the classical concepts of niche differentiation and limiting similarity, one that retains the original intuition behind the terms but avoids some of the pitfalls of earlier approaches.

The second co-recipient is Simon Stump and Peter Chesson for their paper entitled “Distance-responsive predation is not necessary for the Janzen-Connell hypothesis” (Theoretical Population Biology 2015, 106:60-70).  This paper uses mathematical models to test three of the core claims of the Janzen-Conell hypothesis – a major hypotheses for explaining tropical biodiversity.  Using an elegant site occupancy model, the authors show that each of these claims are problematic.  For example, many studies on the Janzen-Connell hypothesis have focused on testing whether predators are distance-responsive, but the authors show that distance-responsive predations are actually less able to promote coexistence than more widely dispersing predators.  Additionally, this paper resolves contradictory results in both the theoretical and empirical literature, and suggests ways to improve experimental testing.  

A number of impressive papers were nominated for the prize this year, and the section executive appreciated the opportunity to read and evaluate all of the nominated papers.  We would also like to note that one of our officers removed herself from the final decision-making process due to her role as a coauthor on one of the papers.  The remainder of the section executive felt strongly that this paper should not be overlooked for the award due to her involvement.


 G.A. Maciel, F. Lutscher (2013) How individual movement response to habitat edges affects population persistence and spatial spread. American Naturalist 182(1): 42-52

This paper addresses a basic question motivated by ecological dynamics — how behavior at habitat edges affects population persistence and spatial spread — and comprehensively presents and analyzes a set of broadly applicable tools for incorporating edge behavior. In doing so, Maciel & Lutscher nicely balance tractable models with biological realism and clearly describe how their modeling insights fit into the existing theoretical and empirical context. In addition to advancing the long-standing theory of spatial population dynamics, the findings clearly contribute to the biological understanding of the effect of a variety of movement behaviors, such as faster movement through unfavorable habitat enhancing population persistence and an intermediate strength of patch-quality-based preference enhancing the rate of population spread, with a strong mechanistic understanding for why these dynamics arise.

We had seventeen outstanding nominees for the best paper award this year; many thanks to everyone who submitted nominations. Please join us in congratulating Gabriel Maciel and Frithjof Lutscher on their achievement.


The winners of the 2014 award for the best paper in theoretical ecology are Jiang Jiang and Don DeAngelis, for their 2013 paper “Strong species-environment feedback shapes plant community assembly along environmental gradients,” published in the journal Ecology and Evolution (3: 4119–4128).

In their clearly-written paper, the authors make direct linkages to problems in plant ecology, while building a general theoretical model that addresses a key issue, not just in plant ecology, of feedbacks between organisms and their environment. Through well-designed analyses of an elegant model, they found that “ecological engineers” (species that modify the environment to their own benefit) can affect the diversity of the competitive community they inhabit, and that the direction of this effect depends critically on the extent to which the community is closed to immigration and on the spatial heterogeneity of the environment. These novel results should are likely to foster further theoretical research and generate some fine hypotheses that will motivate experimental and field studies.

Please join me in congratulating Jiang and Don!


The Theoretical Ecology Section is pleased to award the 2013 prize for an outstanding theoretical ecology paper to “Nitrogen and Phosphorus Limitation over Long-term Ecosystem Development in Terrestrial Ecosystems” by Duncan Menge, Lars Hedin, and Stephen Pacala, published in PLoS One (2012) vol. 7(8) e42045.

This paper formulates a mathematical model for looking at nitrogen (N) and phosphorus (P) dynamics over multiple timescales. Analyzing their model through techniques of timescale separation, the authors determine whether N or P is more likely to limit net primary productivity (NPP) at short, intermediate, and long timescales. While some of their results provide support for already well-established ideas on NPP limitation, others are counterintuitive, thereby nicely demonstrating the importance of quantitative models for understanding dynamics at the level of the ecosystem. Finally, the work is elegant in that it nicely integrates disparate observations into one cohesive framework.


The Theoretical Ecology Section is pleased to award the 2012 prize for an outstanding paper to “Comparing the qualitatively different effects of rapidly evolving and rapidly induced defences have on predator-prey interactions” by Michael Cortez, published in Ecology Letters (2011) 14:202-209.

This paper unifies models of the ecological impacts of plasticity and rapid evolution, and extends these models to acheive general insights about similarities and differences between heritable and non-heritable trait variation. The results were not intuitive; induced defences stabilize or syncrhonize fluctuations (depending on the rate of induction), whereas rapid evolution can lead to a broader range of dynamics. These results were obtained through mathematical innovation, specifically, application of slow-fast theory to make a general analysis tractable.

Congratulations to the authors!


The recipients of the 2011 award are Heather Berkley, Bruce Kendall, Satoshi Mitarai, and David Siegel for their paper entitled “Turbulent dispersal promotes species coexistence”, published in Ecology Letters 13:360-371.

This paper uses spatially explicit simulations and analytical approximations to demonstrate that, for species with dispersed larvae and sessile adults, stochastic larval dispersal driven by environmental turbulence can result in decorrelated settlement patterns that enable long-term coexistence of competing species.  The paper demonstrates how the nuanced interplay between species traits and their abiotic environment can drive population-scale processes, and illustrates how multiple sources of variation contribute differently to long-term community dynamics.

Congratulations to the authors!


James O’Dwyer and Jessica Green (2009), for their paper entitled “Field theory for biogeography: A spatially explicit model for predicting patterns of biodiversity,” found in Ecology Letters 13: 87-95.

Using mathematical methods more commonly found in quantum physics, the authors derive the species-area relationship for a spatially explicit neutral model that includes dispersal and generalizes a previous prediction about beta diversity.

Congratulations to the authors!


Last updated on: September 1, 2020