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Creators/Authors contains: "Kraft, Nathan J. B."

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  1. Abstract

    Contemporary studies of species coexistence are underpinned by deterministic models that assume that competing species have continuous (i.e., noninteger) densities, live in infinitely large landscapes, and coexist over infinite time horizons. By contrast, in nature, species are composed of discrete individuals subject to demographic stochasticity and occur in habitats of finite size where extinctions occur in finite time. One consequence of these discrepancies is that metrics of species’ coexistence derived from deterministic theory may be unreliable predictors of the duration of species coexistence in nature. These coexistence metrics include invasion growth rates and niche and fitness differences, which are now commonly applied in theoretical and empirical studies of species coexistence. In this study, we tested the efficacy of deterministic coexistence metrics on the duration of species coexistence in a finite world. We introduce new theoretical and computational methods to estimate coexistence times in stochastic counterparts of classic deterministic models of competition. Importantly, we parameterized this model using experimental field data for 90 pairwise combinations of 18 species of annual plants, allowing us to derive biologically informed estimates of coexistence times for a natural system. Strikingly, we found that for species expected to deterministically coexist, community sizes containing only 10 individuals had predicted coexistence times of more than 1000 years. We also found that invasion growth rates explained 60% of the variation in intrinsic coexistence times, reinforcing their general usefulness in studies of coexistence. However, only by integrating information on both invasion growth rates and species' equilibrium population sizes could most (>99%) of the variation in species coexistence times be explained. This integration was achieved with demographically uncoupled single‐species models solely determined by the invasion growth rates and equilibrium population sizes. Moreover, because of a complex relationship between niche overlap/fitness differences and equilibrium population sizes, increasing niche overlap and increasing fitness differences did not always result in decreasing coexistence times, as deterministic theory would predict. Nevertheless, our results tend to support the informed use of deterministic theory for understanding the duration of species’ coexistence while highlighting the need to incorporate information on species' equilibrium population sizes in addition to invasion growth rates.

     
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  2. Abstract

    Turnover in species composition and the dominant functional strategies in plant communities across environmental gradients is a common pattern across biomes, and is often assumed to reflect shifts in trait optima. However, the extent to which community‐wide trait turnover patterns reflect changes in how plant traits affect the vital rates that ultimately determine fitness remain unclear.

    We tested whether shifts in the community‐weighted means of four key functional traits across an environmental gradient in a southern California grassland reflect variation in how these traits affect species' germination and fecundity across the landscape.

    We asked whether models that included trait–environment interactions help explain variation in two key vital rates (germination rates and fecundity), as well as an integrative measure of fitness incorporating both vital rates (the product of germination rate and fecundity). To do so, we planted seeds of 17 annual plant species at 16 sites in cleared patches with no competitors, and quantified the lifetime seed production of 1360 individuals. We also measured community composition and a variety of abiotic variables across the same sites. This allowed us to evaluate whether observed shifts in community‐weighted mean traits matched the direction of any trait–environment interactions detected in the plant performance experiment.

    We found that commonly measured plant functional traits do help explain variation in species responses to the environment—for example, high‐SLA species had a demographic advantage (higher germination rates and fecundity) in sites with high soil Ca:Mg levels, while low‐SLA species had an advantage in low Ca:Mg soils. We also found that shifts in community‐weighted mean traits often reflect the direction of these trait–environment interactions, though not all trait–environment relationships at the community level reflect changes in optimal trait values across these gradients.

    Synthesis. Our results show how shifts in trait–fitness relationships can give rise to turnover in plant phenotypes across environmental gradients, a fundamental pattern in ecology. We highlight the value of plant functional traits in predicting species responses to environmental variation, and emphasise the need for more widespread study of trait–performance relationships to improve predictions of community responses to global change.

     
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  3. Abstract

    When species simultaneously compete with two or more species of competitor, higher‐order interactions (HOIs) can lead to emergent properties not present when species interact in isolated pairs. To extend ecological theory to multi‐competitor communities, ecologists must confront the challenges of measuring and interpreting HOIs in models of competition fit to data from nature. Such efforts are hindered by the fact that different studies use different definitions, and these definitions have unclear relationships to one another. Here, we propose a distinction between ‘soft’ HOIs, which identify possible interaction modification by competitors, and ‘hard’ HOIs, which identify interactions uniquely emerging in systems with three or more competitors. We show how these two classes of HOI differ in their motivation and interpretation, as well as the tests one uses to identify them in models fit to data. We then show how to operationalise this structure of definitions by analysing the results of a simulated competition experiment underlain by a consumer resource model. In the course of doing so, we clarify the challenges of interpreting HOIs in nature, and suggest a more precise framing of this research endeavour to catalyse further investigations.

     
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  4. Quaternary climate change reduced and homogenized angiosperm tree diversity across large landscapes worldwide. 
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  5. Abstract

    Recent work has shown that evaluating functional trait distinctiveness, the average trait distance of a species to other species in a community offers promising insights into biodiversity dynamics and ecosystem functioning. However, the ecological mechanisms underlying the emergence and persistence of functionally distinct species are poorly understood. Here, we address the issue by considering a heterogeneous fitness landscape whereby functional dimensions encompass peaks representing trait combinations yielding positive population growth rates in a community. We identify four ecological cases contributing to the emergence and persistence of functionally distinct species. First, environmental heterogeneity or alternative phenotypic designs can drive positive population growth of functionally distinct species. Second, sink populations with negative population growth can deviate from local fitness peaks and be functionally distinct. Third, species found at the margin of the fitness landscape can persist but be functionally distinct. Fourth, biotic interactions (positive or negative) can dynamically alter the fitness landscape. We offer examples of these four cases and guidelines to distinguish between them. In addition to these deterministic processes, we explore how stochastic dispersal limitation can yield functional distinctiveness. Our framework offers a novel perspective on the relationship between fitness landscape heterogeneity and the functional composition of ecological assemblages.

     
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  6. Abstract

    Plant species can show considerable morphological and functional variation along environmental gradients. This intraspecific trait variation (ITV) can have important consequences for community assembly, biotic interactions, ecosystem functions and responses to global change. However, directly measuring ITV across many species and wide geographic areas is often infeasible. Thus, a method to predict spatial variation in a species’ functional traits could be valuable.

    We measured specific leaf area (SLA), height and leaf area (LA) of grasses across California, covering 59 species at 230 sampling locations. We asked how these traits change along climate gradients within each species and used machine learning to predict local trait values for any species at any location based on phylogenetic position, local climate and that species’ mean traits. We then examined how much these local predictions alter patterns of assemblage‐level trait variation across the state.

    Most species exhibited higher SLA and grew taller at higher temperatures and produced larger leaves in drier conditions. The random forests predicted spatial variation in functional traits very accurately, with correlations up to 0.97. Because trait records were spatially biased towards warmer areas, and these areas tend to have higher SLA individuals within each species, species means of SLA were upwardly biased. As a result, using species means over‐estimates SLA in the cooler regions of the state. Our results also suggest that height may be substantially under‐predicted in the warmest areas.

    Synthesis. Using only species mean traits to characterize the functional composition of communities risks introducing substantial error into trait‐based estimates of ecosystem properties including decomposition rates or NPP. The high performance of random forests in predicting local trait values provides a way forward for estimating high‐resolution patterns of ITV without a massive data collection effort.

     
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  7. Abstract

    Many plant species exhibit strong association with topographic habitats at local scales. However, the historical biogeographic and physiological drivers of habitat specialization are still poorly understood, and there is a need for relatively easy‐to‐measure predictors of species habitat niche breadth. Here, we explore whether species geographic range, climatic envelope, or intraspecific variability in leaf traits is related to the degree of habitat specialization in a hyperdiverse tropical tree community in Amazonian Ecuador. Contrary to our expectations, we find no effect of the size of species geographic ranges, the diversity of climate a species experiences across its range, or intraspecific variability in leaf traits in predicting topographic habitat association in the ~300 most common tropical tree species in a 25‐ha tropical forest plot. In addition, there was no phylogenetic signal to habitat specialization. We conclude that species geographic range size, climatic niche breadth, and intraspecific variability in leaf traits fail to capture the habitat specialization patterns observed in this highly diverse tropical forest.

     
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  8. Abstract

    Latitudinal and elevational richness gradients have received much attention from ecologists but there is little consensus on underlying causes. One possible proximate cause is increased levels of species turnover, or β diversity, in the tropics compared to temperate regions. Here, we leverage a large botanical dataset to map taxonomic and phylogenetic β diversity, as mean turnover between neighboring 100 × 100 km cells, across the Americas and determine key climatic drivers. We find taxonomic and tip‐weighted phylogenetic β diversity is higher in the tropics, but that basal‐weighted phylogenetic β diversity is highest in temperate regions. Supporting Janzen's ‘mountain passes’ hypothesis, tropical mountainous regions had higher β diversity than temperate regions for taxonomic and tip‐weighted metrics. The strongest climatic predictors of turnover were average temperature and temperature seasonality. Taken together, these results suggest β diversity is coupled to latitudinal richness gradients and that temperature is a major driver of plant community composition and change.

     
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