About

Jonathan Levine is the J.N. Allison Professor of Environmental Studies and serves as the Department Chair of Ecology & Evolutionary Biology at Princeton University. His research focuses on population, community, and global change ecology. Levine has contributed to understanding the effects of rapid evolution on species coexistence, the mechanisms of species coexistence in complex communities, and how rapid evolution influences plant population spread in fragmented landscapes. His work also explores how novel competitors shape species' responses to climate change and the role of plant functional traits in species coexistence.

Research topics

• Biology
• Ecology
• Computer Science
• Environmental science
• Geology
• Atmospheric sciences
• Microbiology

Selected publications

• Seasonal evolution of <i>Drosophila melanogaster</i> abdominal pigmentation is associated with a multifarious selective landscape

  Evolution · 2026-03-11

  articleOpen access

  Pigmentation has been widely studied by evolutionary biologists due to both ease of measurement and relationship to fitness. Drosophila melanogaster pigmentation has represented a particularly useful avenue of investigation, as extensive genetic tools have enabled the characterization of the trait's complex architecture. Drosophila pigmentation also varies predictably across space and time in wild populations, suggesting pigmentation is a component of adaptation to local environmental conditions. Despite this, the impact of D. melanogaster pigmentation on fitness, and the environmental factors that drive the evolution of pigmentation, are not well understood. To address this gap, we experimentally evolved replicated D. melanogaster populations in field mesocosms to determine whether and how pigmentation evolves in response to environmental variation. We found that pigmentation rapidly and predictably adapted to a direct manipulation of temperature, supportive of melanization playing a role in thermoregulation. However, we also determined that pigmentation responded adaptively to direct manipulations of numerous additional factors, including intraspecific competition, diet, and the microbiome. These findings suggest that the selective landscape acting on pigmentation is complex and multifaceted, and that patterns of melanization may be driven, at least in part, by indirect selection due to correlations with other fitness-related traits.

  PublisherDOI

• Variable impacts of land-based climate mitigation on habitat area for vertebrate diversity

  Science · 2025-01-23 · 12 citations

  articleSenior author

  Pathways to achieving net zero carbon emissions commonly involve deploying reforestation, afforestation, and bioenergy crops across millions of hectares of land. It is often assumed that by helping to mitigate climate change, these strategies indirectly benefit biodiversity. Here, we modeled the climate and habitat requirements of 14,234 vertebrate species and show that the impact of these strategies on species' habitat area tends not to arise through climate mitigation, but rather through habitat conversion. Across locations, reforestation tends to provide species more habitat through both land-cover change and climate mitigation, whereas habitat loss from afforestation and bioenergy cropping typically outweighs the climate mitigation benefits. This work shows how and where land-based mitigation strategies can be deployed without inadvertently reducing the area of habitat for global biodiversity.

  PublisherDOI

• Trait diversity in plant communities maintained by competition for water and light

  Ecological Monographs · 2025-02-01 · 4 citations

  articleOpen access

  Abstract Ecological communities frequently exhibit remarkable taxonomic and trait diversity, and this diversity is consistently shown to regulate ecosystem function and resilience. However, ecologists lack a synthetic theory for how this diversity is maintained when species compete for limited resources, hampering our ability to project the future of biodiversity under climate change. Water‐limited plant communities are an ideal system in which to study these questions given (1) the diversity of hydraulic traits they exhibit, (2) the importance of this diversity for ecosystem productivity and drought resilience, and (3) forecast changes to precipitation and evapotranspiration under climate change. We developed an analytically tractable model of water and light competition in age‐structured perennial plant communities and demonstrated that high diversity is maintained through phenological division of the time between storms. We modeled a system where water arrives in the form of intermittent storms, between which plants consume the limited pool of soil water until it becomes dry enough that they must physiologically shut down to avoid embolism. Competition occurs because individuals, by consuming the shared water pool, cause their competitors to shut down earlier, harming their long‐term growth and reproduction. When total precipitation is low, plants in the model compete only for water. However, increases in precipitation can cause the canopy to close and individuals to begin competing for light. Variation among species in the minimum soil water content at which they can sustain growth without embolizing leads to emergent phenological variation, as species will shut down at varying points between storm events. When this variation is paired with a trade‐off such that species that shut down early are compensated by faster biomass accumulation, higher fecundity, or lower mortality, there is no limit to the number that can coexist. These results are robust to variation in both total precipitation and the time between storms. The model therefore offers a plausible explanation for how hydraulic trait diversity is maintained in a wide array of natural systems. More broadly, this work illustrates how the phenological division of an apparently singular resource can emerge because of common trade‐offs and ultimately foster high taxonomic and trait diversity.

  PublisherOA PDFDOI

• Proximity-centred accessibility – A conceptual debate involving planning practitioners worldwide

  Cities · 2025-08-26 · 6 citations

  articleOpen access

  In recent years, the concept of proximity has garnered increasing attention in both transportation research and practice, albeit under various terms and interpretations. Among these, the concept of the 15-minute city has catalysed attention in planning practice, with recent evolution to the x-minute city and city of proximities . In research, proximity-centred accessibility has been offered as an umbrella term to express the ability to reach activities and destinations at short distances. Regardless of the terminology used, the essence of proximity lies in the ease with which one can access desired activities and destinations within reasonable travel times, independent of speed-enhancing transport modes most notably through walking. This research investigates the nuanced meanings ascribed to proximity-centred accessibility by planning practitioners globally, spanning diverse regional and local contexts. For this, we used an online survey, disseminated among over 9000 practitioners from 22 countries across 5 continents, which generated over 1300 responses. The survey explored the preferred terms for proximity-centred accessibility and their definitions, specifically emphasizing time and distance thresholds and the identification of relevant activities. By juxtaposing our findings with an earlier survey of accessibility researchers, this study also contributes to the groundwork for a conceptual framework for proximity-centred accessibility. Our findings affirm a relatively consistent interpretation of proximity among global planning practitioners, predominantly extending up to 1600 m, in accordance with earlier results for accessibility researchers. Despite some relevant dissimilarities among practitioners from megacities compared to their smaller city counterparts, or in specific countries (most notably the Netherlands), the distance that is considered proximate is the attribute that generates the most consistent results across different contexts. Also consistent was the relevance of short distances (up to 15 min walking) for activities such as primary and pre-primary schools, playgrounds, parks, food shopping, and pharmacies, reinforcing the importance of proximity to basic and caregiving activities. No term was found to be consistently meaningful across different contexts, although terms like local and neighbourhood accessibility and walking/pedestrian, or cycling accessibility, show higher preference in the global sample. • Consistent Distance Threshold: Most practitioners regard distances up to 1,600 meters as proximate, aligning closely with accessibility researchers’ interpretations. This threshold was robust across diverse geographic contexts. • Time-Based Proximity: A 15-minute walking threshold was commonly associated with access to essential daily activities, particularly caregiving and basic services (e.g., schools, parks, pharmacies, food shopping). • Terminology Diversity: No single term emerged as universally accepted. However, "local accessibility," "neighbourhood accessibility," and terms emphasizing walking or cycling gained relative prominence. • Contextual Nuances: Minor variations were observed across urban scales (e.g., megacities vs. smaller cities) and national contexts (e.g., the Netherlands), though these did not substantially alter the overall patterns.

  PublisherDOI

• Seasonal evolution of <i>Drosophila melanogaster</i> abdominal pigmentation is associated with a multifarious selective landscape

  bioRxiv (Cold Spring Harbor Laboratory) · 2025-08-20

  preprintOpen access

  Abstract Pigmentation has been widely studied by evolutionary biologists due to both ease of measure and relationship to fitness. Drosophila melanogaster pigmentation has represented a particularly useful avenue of investigation, as extensive genetic tools have enabled the characterization of the trait’s complex architecture. Drosophila pigmentation also varies predictably across space and time in wild populations, suggesting pigmentation is a component of adaptation to local environmental conditions. Despite this, the impact of D. melanogaster pigmentation on fitness, and the environmental factors that drive the evolution of pigmentation, are not well understood. To address this gap, we experimentally evolved replicated D. melanogaster populations in field mesocosms to determine whether and how pigmentation evolves in response to environmental variation. We found that pigmentation rapidly and predictably adapted to a direct manipulation of temperature, supportive of melanization playing a role in thermoregulation. However, we also determined that pigmentation responded adaptively to direct manipulations of numerous additional factors, including intraspecific competition, diet, and the microbiome. These findings suggest that the selective landscape acting on pigmentation is complex and multifaceted, and that patterns of melanization may be driven, at least in part, by indirect selection due to correlations with other fitness-related traits.

  PublisherOA PDFDOI

• Competitor‐induced plasticity modifies the interactions and predicted competitive outcomes between annual plants

  Ecology · 2025-05-01 · 3 citations

  articleOpen access

  The competitive effect of one individual on another can have impacts beyond just reductions in performance. Because species plastically respond to their environment, competition can also induce changes in species traits, and in turn, these modified traits can then affect interactions with yet other individuals. In this context, plasticity is often argued to favor species coexistence by increasing the niche differentiation between species, though experimental evidence for this hypothesis that explicitly projects competitive outcomes is largely lacking. Here, we transiently subjected four annual plant species to early-season intraspecific or interspecific competition to explicitly induce plastic responses and then examined the response of these individuals to competitors faced later in life. Competing with nearby individuals early in the growing season tended to amplify the sensitivity of individuals to competition, and particularly so for interspecific competition, but the strength of this effect depended on the identity of the focal species. This increase in interspecific relative to intraspecific competition caused plasticity to decrease the predicted likelihood of pairwise coexistence. By combining recent theory with a new experimental approach, we provide a pathway toward integrating phenotypic plasticity into our quantitative understanding of coexistence.

  PublisherOA PDFDOI

• Spatial clustering reveals the impact of higher-order interactions in a diverse annual plant community

  bioRxiv (Cold Spring Harbor Laboratory) · 2025-07-18

  preprintOpen access

  Abstract Spatial patterns are widespread in ecology, but their effects on species interactions remain unresolved, especially in diverse communities. In principle, the degree of spatial clustering could alter the concentration of higher-order interactions, which occur when one (or more) species modifies competition between two others. When species are well mixed, heterospecific neighbors have ample opportunity to modify a competitor’s interactions with other species. In contrast, species clustering can reduce the concentration of interspecific higher-order interactions. In a field experiment with annual grassland plants in California, we manipulated the spatial arrangement — but not the number or identity — of two competitors and measured how they jointly affected a focal individual. We found that focal plants produced more seeds when their competitors were clustered than when they were mixed. These results suggest that interspecific higher-order interactions generally had a stronger competitive (or weaker facilitative) effect than intraspecific ones. However, the effect of clustering varied across species. Larger differences in focal fecundity were correlated with competitors that had greater differences in size and/or functional traits between the spatial arrangements. Additionally, a competitive hierarchy among our study species predicted the effects of clustered versus mixed competitors on focal seed production. Altogether, our work suggests that the spatial arrangement of competitors changes the realized strength of competition in diverse plant communities by modifying the concentration of higher-order interactions. Given the extensive variation in spatial aggregation in plant communities, this mechanism is likely to be a powerful but underappreciated force shaping competition in nature. Significance Statement Plant species coexist in remarkably diverse assemblages throughout the world. Spatial patterns, including aggregation and intermixing, are also widespread in these communities. One potentially underappreciated mechanism that may structure the spatial dynamics of plant communities is interactions that uniquely occur in diverse systems, often called higher-order interactions. Here, we experimentally demonstrated that spatially mediated higher-order interactions operate among annual plants. These higher-order interactions, and their associated changes in competitor size and functional traits, were correlated with the competitive imbalance between competitors. Because both spatial aggregation and competitive hierarchies are widespread in nature, higher-order interactions emerging from their combination may be a more common driver of biodiversity patterns in plant communities than previously thought.

  PublisherDOI

• Rigorous validation of ecological models against empirical time series

  Nature Ecology & Evolution · 2025-10-27 · 3 citations

  articleSenior author
  PublisherDOI

• Linking relative suitability to probability of occurrence in presence‐only species distribution models: Implications for global change projections

  Methods in Ecology and Evolution · 2025-02-17 · 2 citations

  articleOpen accessSenior author

  Abstract Our ecological understanding of how biodiversity will respond to global change is to a large extent based on projections from presence‐only species distribution models. Despite the incredible utility of these models, we know that the predictions they generate can be heavily influenced by user decisions about model structure or parameter choices. Here, we test how the function used to convert relative suitability to probably of occurrence in presence‐only species distributions models can affect predictions of both the magnitude and location of biodiversity change. We used MaxEnt models to create maps of relative suitability for 354 avian species under both current climate conditions and climate conditions in the year 1981. In a back‐casting analysis we tested how well three functions relating relative suitability to probably of occurrence perform in recovering observed changes in range size: (1) a logistic curve with informed, species‐specific, prevalence values, (2) the default logistic curve, or (3) a commonly used statistical threshold. We then quantified the implications of these functions for projections of species' future range shifts with climate change. We found that using either the default logistic function or a common threshold function for habitat suitability tends to (1) estimate larger effects of past climate change on species range size than observed in the time‐series data, (2) inflate projections of how much future climate change will impact species range size and (3) potentially misidentify the locations of greatest range expansion or contraction. We further provide a mathematical basis for these biases, suggesting their general applicability to other systems. Last, we show that these biases can be avoided by analysing proportional rather than absolute changes in range with climate change, and by abandoning the use of habitat suitability thresholds. Incorporating these practices can facilitate a more predictive use of species distribution models when forecasting the response of biodiversity to global change.

  PublisherOA PDFDOI

• Author response for "Competition for time: Evidence for an overlooked, diversity‐maintaining competitive mechanism"

  2024-01-02

  peer-reviewSenior author
  PublisherDOI

Recent grants

• Environmental Variation, Dormancy, and Rare Plant Persistence in Invaded Habitats

  NSF · $516k · 2003–2008

• Collaborative Research: Niche and Neutral Controls over the Coexistence of Serpentine Annual Plants

  NSF · $338k · 2008–2013

• Collaborative Research: Higher order interactions, functional trait changes, and species coexistence in an annual plant community

  NSF · $541k · 2020–2025

Frequent coauthors

• Mark D. Bertness

  Brown University

  27 shared

• Nathan J. B. Kraft

  University of California, Los Angeles

  22 shared

• Óscar Godoy

  Estación Biológica de Doñana

  20 shared

• George H. Leonard

  19 shared

• Aubrey O. Ingraham

  Dartmouth College

  18 shared

• Paul R. Schmidt

  18 shared

• Peter B. Adler

  Utah State University

  17 shared

• Stephen W. Pacala

  Princeton University

  17 shared

Labs

• Levine LabPI

Education

• Ph.D., Integrative Biology

  University of California, Berkeley

  2001

• B.Sc., Biology

  Brown University

  1995