Interactions between temperature and nutrients determine the population dynamics of primary producers.

Global change is rapidly and fundamentally altering many of the processes regulating the flux of energy throughout ecosystems, and although researchers now understand the effect of temperature on key rates (such as aquatic primary productivity), the theoretical foundation needed to generate forecasts of biomass dynamics and extinction risk remains underdeveloped. We develop new theory that describes the interconnected effects of nutrients and temperature on phytoplankton populations and show that the thermal response of equilibrium biomass (i.e. carrying capacity) always peaks at a lower temperature than for productivity (i.e. growth rate). This mismatch is driven by differences in the thermal responses of growth, death, and per-capita impact on the nutrient pool, making our results highly general and applicable to widely used population models beyond phytoplankton. We further show that non-equilibrium dynamics depend on the pace of environmental change relative to underlying vital rates and that populations respond to variable environments differently at high versus low temperatures due to thermal asymmetries.

Stability of consumer-resource interactions in periodic environments.

Periodic fluctuations in abiotic conditions are ubiquitous across a range of temporal scales and regulate the structure and function of ecosystems through dynamic biotic responses that are adapted to these external forces. Research has suggested that certain environmental signatures may play a crucial role in the maintenance of biodiversity and the stability of food webs, while others argue that coupled oscillators ought to promote chaos. As such, numerous uncertainties remain regarding the intersection of temporal environmental patterns and biological responses, and we lack a general understanding of the implications for food web stability. Alarmingly, global change is altering the nature of both environmental rhythms and biological rates. Here, we develop a general theory for how continuous periodic variation in productivity, across temporal scales, influences the stability of consumer-resource interactions: a fundamental building block of food webs. Our results suggest that consumer-resource dynamics under environmental forcing are highly complex and depend on asymmetries in both the speed of forcing relative to underlying dynamics and in local stability properties. These asymmetries allow for environmentally driven stabilization under fast forcing, relative to underlying dynamics, as well as extremely complex and unstable dynamics at slower periodicities. Our results also suggest that changes in naturally occurring periodicities from climate change may lead to precipitous shifts in dynamics and stability.

Landscape modification and nutrient-driven instability at a distance.

Almost 50 years ago, Michael Rosenzweig pointed out that nutrient addition can destabilise food webs, leading to loss of species and reduced ecosystem function through the paradox of enrichment. Around the same time, David Tilman demonstrated that increased nutrient loading would also be expected to cause competitive exclusion leading to deleterious changes in food web diversity. While both concepts have greatly illuminated general diversity-stability theory, we currently lack a coherent framework to predict how nutrients influence food web stability across a landscape. This is a vitally important gap in our understanding, given mounting evidence of serious ecological disruption arising from anthropogenic displacement of resources and organisms. Here, we combine contemporary theory on food webs and meta-ecosystems to show that nutrient additions are indeed expected to drive loss in stability and function in human-impacted regions. Our models suggest that destabilisation is more likely to be caused by the complete loss of an equilibrium due to edible plant species being competitively excluded. In highly modified landscapes, spatial nutrient transport theory suggests that such instabilities can be amplified over vast distances from the sites of nutrient addition. Consistent with this theoretical synthesis, the empirical frequency of these distant propagating ecosystem imbalances appears to be growing. This synthesis of theory and empirical data suggests that human modification of the Earth is strongly connecting distantly separated ecosystems, causing rapid, expansive and costly nutrient-driven instabilities over vast areas of the planet. Similar to existing food web theory, the corollary to this spatial nutrient theory is that slowing down spatial nutrient pathways can be a potent means of stabilising degraded ecosystems.

Winter in water: differential responses and the maintenance of biodiversity.

The ecological consequences of winter in freshwater systems are an understudied but rapidly emerging research area. Here, we argue that winter periods of reduced temperature and light (and potentially oxygen and resources) could play an underappreciated role in mediating the coexistence of species. This may be especially true for temperate and subarctic lakes, where seasonal changes in the thermal environment might fundamentally structure species interactions. With climate change already shortening ice-covered periods on temperate and polar lakes, consideration of how winter conditions shape biotic interactions is urgently needed. Using freshwater fishes in northern temperate lakes as a case study, we demonstrate how physiological trait differences (e.g. thermal preference, light sensitivity) drive differential behavioural responses to winter among competing species. Specifically, some species have a higher capacity for winter activity than others. Existing and new theory is presented to argue that such differential responses to winter can promote species coexistence. Importantly, if winter is a driver of niche differences that weaken competition between, relative to within species, then shrinking winter periods could threaten coexistence by tipping the scales in favour of certain sets of species over others.

Supply and demand drive a critical transition to dysfunctional fisheries.

There is growing awareness of the need for fishery management policies that are robust to changing environmental, social, and economic pressures. Here we use conventional bioeconomic theory to demonstrate that inherent biological constraints combined with nonlinear supply-demand relationships can generate threshold effects due to harvesting. As a result, increases in overall demand due to human population growth or improvement in real income would be expected to induce critical transitions from high-yield/low-price fisheries to low-yield/high-price fisheries, generating severe strains on social and economic systems as well as compromising resource conservation goals. As a proof of concept, we show that key predictions of the critical transition hypothesis are borne out in oceanic fisheries (cod and pollock) that have experienced substantial increase in fishing pressure over the past 60 y. A hump-shaped relationship between price and historical harvest returns, well demonstrated in these empirical examples, is particularly diagnostic of fishery degradation. Fortunately, the same heuristic can also be used to identify reliable targets for fishery restoration yielding optimal bioeconomic returns while safely conserving resource abundance.

The productivity-stability trade-off in global food systems.

Historically, humans have managed food systems to maximize productivity. This pursuit has drastically modified terrestrial and aquatic ecosystems globally by reducing species diversity and body size while creating very productive, yet homogenized, environments. Such changes alter the structure and function of ecosystems in ways that ultimately erode their stability. This productivity-stability trade-off has largely been ignored in discussions around global food security. Here, we synthesize empirical and theoretical literature to demonstrate the existence of the productivity-stability trade-off and argue the need for its explicit incorporation in the sustainable management of food systems. We first explore the history of human management of food systems, its impacts on average body size within and across species and food web stability. We then demonstrate how reductions in body size are symptomatic of a broader biotic homogenization and rewiring of food webs. We show how this biotic homogenization decompartmentalizes interactions among energy channels and increases energy flux within the food web in ways that threaten their stability. We end by synthesizing large-scale ecological studies to demonstrate the prevalence of the productivity-stability trade-off. We conclude that management strategies promoting landscape heterogeneity and maintenance of key food web structures are critical to sustainable food production.

Food web rewiring in a changing world.

Climate change is asymmetrically altering environmental conditions in space, from local to global scales, creating novel heterogeneity. Here, we argue that this novel heterogeneity will drive mobile generalist consumer species to rapidly respond through their behaviour in ways that broadly and predictably reorganize - or rewire - food webs. We use existing theory and data from diverse ecosystems to show that the rapid behavioural responses of generalists to climate change rewire food webs in two distinct and critical ways. First, mobile generalist species are redistributing into systems where they were previously absent and foraging on new prey, resulting in topological rewiring - a change in the patterning of food webs due to the addition or loss of connections. Second, mobile generalist species, which navigate between habitats and ecosystems to forage, will shift their relative use of differentially altered habitats and ecosystems, causing interaction strength rewiring - changes that reroute energy and carbon flows through existing food web connections and alter the food web's interaction strengths. We then show that many species with shared traits can exhibit unified aggregate behavioural responses to climate change, which may allow us to understand the rewiring of whole food webs. We end by arguing that generalists' responses present a powerful and underutilized approach to understanding and predicting the consequences of climate change and may serve as much-needed early warning signals for monitoring the looming impacts of global climate change on entire ecosystems.