Charlotte T. Lee

LONGEVITY CAN BUFFER PLANT AND ANIMAL POPULATIONS AGAINST CHANGING CLIMATIC VARIABILITY

Both means and year-to-year variances of climate variables such as temperature and precipitation are predicted to change. However, the potential impact of changing climatic variability on the fate of populations has been largely unexamined. We analyzed multiyear demographic data for 36 plant and animal species with a broad range of life histories and types of environment to ask how sensitive their long-term stochastic population growth rates are likely to be to changes in the means and standard deviations of vital rates (survival, reproduction, growth) in response to changing climate. We quantified responsiveness using elasticities of the long-term population growth rate predicted by stochastic projection matrix models. Short-lived species (insects and annual plants and algae) are predicted to be more strongly (and negatively) affected by increasing vital rate variability relative to longer-lived species (perennial plants, birds, ungulates). Taxonomic affiliation has little power to explain sensitivity to increasing variability once longevity has been taken into account. Our results highlight the potential vulnerability of short-lived species to an increasingly variable climate, but also suggest that problems associated with short-lived undesirable species (agricultural pests, disease vectors, invasive weedy plants) may be exacerbated in regions where climate variability decreases.

Population and prehistory I: Food-dependent population growth in constant environments.

We present a demographic model that describes the feedbacks between food supply, human mortality and fertility rates, and labor availability in expanding populations, where arable land area is not limiting. This model provides a quantitative framework to describe how environment, technology, and culture interact to influence the fates of preindustrial agricultural populations. We present equilibrium conditions and derive approximations for the equilibrium population growth rate, food availability, and other food-dependent measures of population well-being. We examine how the approximations respond to environmental changes and to human choices, and find that the impact of environmental quality depends upon whether it manifests through agricultural yield or maximum (food-independent) survival rates. Human choices can complement or offset environmental effects: greater labor investments increase both population growth and well-being, and therefore can counteract lower agricultural yield, while fertility control decreases the growth rate but can increase or decrease well-being. Finally we establish equilibrium stability criteria, and argue that the potential for loss of local stability at low population growth rates could have important consequences for populations that suffer significant environmental or demographic shocks.

Population and prehistory III: Food-dependent demography in variable environments

The population dynamics of preindustrial societies depend intimately on their surroundings, and food is a primary means through which environment influences population size and individual well-being. Food production requires labor; thus, dependence of survival and fertility on food involves dependence of a populations future on its current state. We use a perturbation approach to analyze the effects of random environmental variation on this nonlinear, age-structured system. We show that in expanding populations, direct environmental effects dominate induced population fluctuations, so environmental variability has little effect on mean hunger levels, although it does decrease population growth. The growth rate determines the time until population is limited by space. This limitation introduces a tradeoff between population density and well-being, so population effects become more important than the direct effects of the environment: environmental fluctuation increases mortality, releasing density dependence and raising average well-being for survivors. We discuss the social implications of these findings for the long-term fate of populations as they transition from expansion into limitation, given that conditions leading to high well-being during growth depress well-being during limitation.

Mutualism between Consumers and Their Shared Resource Can Promote Competitive Coexistence

Competitive coexistence depends on dynamic interactions between competitor and resource populations, including mutualism between the resource and each competitor. We add mutualism to a well‐known model of resource competition and show that it can powerfully stabilize competitive coexistence in the absence or presence of resource heterogeneity. We use a transition matrix approach to describe lottery competition, while allowing each of two competitors to affect the population dynamics of their shared resource. For example, two plant‐defending ant species may compete for nesting space within ant‐adapted (myrmecophytic) plants. We show that mutualism between consumers and a resource species can stabilize competitive coexistence of the consumers by allowing each competitor to influence resource dynamics in a way that benefits the other. The effect of this novel coexistence mechanism depends on a mutualism’s biological details: for example, altering myrmecophyte fecundity affects competing ant species differently than does altering plant survival. Finally, we consider a heterogeneous resource (e.g., two types of nest site) and show how niche partitioning can stabilize coexistence in the absence of resource dynamics. When resource heterogeneity is dynamic (e.g., small and large plants of the same species), niche partitioning also provides new routes for additional stabilization via mutualism.

Inherent demographic stability in mutualist-resource-exploiter interactions.

Core principles of ecological theory predict that, in the absence of other factors, mutualisms should experience destabilizing positive feedback and should be vulnerable to extinction through competitive exclusion by exploiter species. Many effective stabilizing mechanisms address one issue or the other, and many turn upon additional features. Using an explicitly demographic approach, I show that indirect, demography-mediated interactions between mutualists and exploiters can enable mutualist-exploiter coexistence, which in turn can stabilize the abundances of mutualists, exploiters, and their shared resources. This occurs because of the distinct resource demographic responses that are inherent to interaction with mutualistic and exploitative partners and can occur in long-lasting, exclusive interactions, such as protection mutualisms, as well as in apparently very different, short-lived mutualistic interactions, such as pollination. The key necessary factor—demographic response to interspecific interaction—is common in nature. Some demographic structure is also necessary and is generated through interspecific interaction in long-lasting associations; it is also very common in natural populations. Thus, the explicitly demographic and multispecies approach taken here constitutes a potentially promising single explanation for the apparent stability of mutualism in a wide range of natural systems.

Consumer Effects on the Vital Rates of Their Resource Can Determine the Outcome of Competition between Consumers

Current competition theory does not adequately address the fact that competitors may affect the survival, growth, and reproductive rates of their resources. Ecologically important interactions in which consumers affect resource vital rates range from parasitism and herbivory to mutualism. We present a general model of competition that explicitly includes consumer-dependent resource vital rates. We build on the classic MacArthur model of competition for multiple resources, allowing direct comparison with expectations from established concepts of resource-use overlap. Consumers share a stage-structured resource population but may use the different stages to different extents, as they do the different independent resources in the classic model. Here, however, the stages are dynamically linked via consumer-dependent vital rates. We show that consumers’ effects on resource vital rates result in two important departures from classic results. First, consumers can coexist despite identical use of resource stages, provided each competitor shifts the resource stage distribution toward stages that benefit other species. Second, consumers specializing on different resource stages can compete strongly, possibly resulting in competitive exclusion despite a lack of resource stage-use overlap. Our model framework demonstrates the critical role that consumer-dependent resource vital rates can play in competitive dynamics in a wide range of biological systems.

Dynamic preferential allocation to arbuscular mycorrhizal fungi explains fungal succession and coexistence

Evidence accumulates about the role of arbuscular mycorrhizal (AM) fungi in shaping plant communities, but little is known about the factors determining the biomass and coexistence of several types of AM fungi in a plant community. Here, using a consumer-resource framework that treats the relationship between plants and fungi as simultaneous, reciprocal exploitation, we investigated what patterns of dynamic preferential plant carbon allocation to empirically-defined fungal types (on-going partner choice) would be optimal for plants, and how these patterns depend on successional dynamics. We found that ruderal AM fungi can dominate under low steady-state nutrient availability, and competitor AM fungi can dominate at higher steady-state nutrient availability; these are conditions characteristic of early and late succession, respectively. We also found that dynamic preferential allocation alone can maintain a diversity of mutualists, suggesting that on-going partner choice is a new coexistence mechanism for mutualists. Our model can therefore explain both mutualist coexistence and successional strategy, providing a powerful tool to derive testable predictions.

Unifying mutualism diversity for interpretation and prediction

Coarse-grained rules are widely used in chemistry, physics and engineering. In biology, however, such rules are less common and under-appreciated. This gap can be attributed to the difficulty in establishing general rules to encompass the immense diversity and complexity of biological systems. Even when a rule is established, it is often challenging to map it to mechanistic details and to quantify these details. We here address these challenges on a study of mutualism, an essential type of ecological interaction in nature. Using an appropriate level of abstraction, we deduced a general rule that predicts the outcomes of mutualistic systems, including coexistence and productivity. We further developed a standardized calibration procedure to apply the rule to mutualistic systems without the need to fully elucidate or characterize their mechanistic underpinnings. Our approach consistently provides explanatory and predictive power with various simulated and experimental mutualistic systems. Our strategy can pave the way for establishing and implementing other simple rules for biological systems.

Elasticity of population growth with respect to the intensity of biotic or abiotic driving factors

Demographic analysis can elucidate how driving factors, such as climate or species interactions, affect populations. One important question is how growth would respond to future changes in the mean intensity of a driving factor or in its variability, such as might be expected in a fluctuating and shifting climate. Here I develop an approach to computing new stochastic elasticities to address this question. The linchpin of this novel approach is the multidimensional demographic difference that expresses how a population responds to change in the driving factor between two discrete levels of intensity. I use this difference to design a perturbation matrix that links data from common empirical sampling schemes with rigorous theory for stochastic elasticities. Although the starting point is a difference, the products of this synthesis are true derivatives: they are elasticity with respect to the mean intensity of a driving factor, and elasticity with respect to variability in a driving factor. Applying the methods to published data, I demonstrate how these new elasticities can shed light on growth rate response within and at the boundary of the previously observed range of the driving factor, thus helpfully indicating nonlinearity in the observed and in the potential future response. The stochastic approach simplifies in a fixed environment, yielding a compact formula for deterministic elasticity to a driving factor.