E.H. van Nes

Global resilience of tropical forest and savanna to critical transitions.

Tree distributions across continents indicate three distinct stable states in tree cover―forest, savanna, and treeless. It has been suggested that tropical forest and savanna could represent alternative stable states, implying critical transitions at tipping points in response to altered climate or other drivers. So far, evidence for this idea has remained elusive, and integrated climate models assume smooth vegetation responses. We analyzed data on the distribution of tree cover in Africa, Australia, and South America to reveal strong evidence for the existence of three distinct attractors: forest, savanna, and a treeless state. Empirical reconstruction of the basins of attraction indicates that the resilience of the states varies in a universal way with precipitation. These results allow the identification of regions where forest or savanna may most easily tip into an alternative state, and they pave the way to a new generation of coupled climate models.

Chaos in a long-term experiment with a plankton community

Mathematical models predict that species interactions such as competition and predation can generate chaos. However, experimental demonstrations of chaos in ecology are scarce, and have been limited to simple laboratory systems with a short duration and artificial species combinations. Here, we present the first experimental demonstration of chaos in a long-term experiment with a complex food web. Our food web was isolated from the Baltic Sea, and consisted of bacteria, several phytoplankton species, herbivorous and predatory zooplankton species, and detritivores. The food web was cultured in a laboratory mesocosm, and sampled twice a week for more than 2,300 days. Despite constant external conditions, the species abundances showed striking fluctuations over several orders of magnitude. These fluctuations displayed a variety of different periodicities, which could be attributed to different species interactions in the food web. The population dynamics were characterized by positive Lyapunov exponents of similar magnitude for each species. Predictability was limited to a time horizon of 15–30 days, only slightly longer than the local weather forecast. Hence, our results demonstrate that species interactions in food webs can generate chaos. This implies that stability is not required for the persistence of complex food webs, and that the long-term prediction of species abundances can be fundamentally impossible.

Slowing down in spatially patterned ecosystems at the brink of collapse.

Predicting the risk of critical transitions, such as the collapse of a population, is important in order to direct management efforts. In any system that is close to a critical transition, recovery upon small perturbations becomes slow, a phenomenon known as critical slowing down. It has been suggested that such slowing down may be detected indirectly through an increase in spatial and temporal correlation and variance. Here, we tested this idea in arid ecosystems, where vegetation may collapse to desert as a result of increasing water limitation. We used three models that describe desertification but differ in the spatial vegetation patterns they produce. In all models, recovery rate upon perturbation decreased before vegetation collapsed. However, in one of the models, slowing down failed to translate into rising variance and correlation. This is caused by the regular self-organized vegetation patterns produced by this model. This finding implies an important limitation of variance and correlation as indicators of critical transitions. However, changes in such self-organized patterns themselves are a reliable indicator of an upcoming transition. Our results illustrate that while critical slowing down may be a universal phenomenon at critical transitions, its detection through indirect indicators may have limitations in particular systems.

Resilience indicators: prospects and limitations for early warnings of regime shifts

In the vicinity of tipping points—or more precisely bifurcation points—ecosystems recover slowly from small perturbations. Such slowness may be interpreted as a sign of low resilience in the sense that the ecosystem could easily be tipped through a critical transition into a contrasting state. Indicators of this phenomenon of ‘critical slowing down (CSD)’ include a rise in temporal correlation and variance. Such indicators of CSD can provide an early warning signal of a nearby tipping point. Or, they may offer a possibility to rank reefs, lakes or other ecosystems according to their resilience. The fact that CSD may happen across a wide range of complex ecosystems close to tipping points implies a powerful generality. However, indicators of CSD are not manifested in all cases where regime shifts occur. This is because not all regime shifts are associated with tipping points. Here, we review the exploding literature about this issue to provide guidance on what to expect and what not to expect when it comes to the CSD-based early warning signals for critical transitions.

Early warning of climate tipping points from critical slowing down: comparing methods to improve robustness

We address whether robust early warning signals can, in principle, be provided before a climate tipping point is reached, focusing on methods that seek to detect critical slowing down as a precursor of bifurcation. As a test bed, six previously analysed datasets are reconsidered, three palaeoclimate records approaching abrupt transitions at the end of the last ice age and three models of varying complexity forced through a collapse of the Atlantic thermohaline circulation. Approaches based on examining the lag-1 autocorrelation function or on detrended fluctuation analysis are applied together and compared. The effects of aggregating the data, detrending method, sliding window length and filtering bandwidth are examined. Robust indicators of critical slowing down are found prior to the abrupt warming event at the end of the Younger Dryas, but the indicators are less clear prior to the Bølling-Allerød warming, or glacial termination in Antarctica. Early warnings of thermohaline circulation collapse can be masked by inter-annual variability driven by atmospheric dynamics. However, rapidly decaying modes can be successfully filtered out by using a long bandwidth or by aggregating data. The two methods have complementary strengths and weaknesses and we recommend applying them together to improve the robustness of early warnings.

Creating a safe operating space for iconic ecosystems

Manage local stressors to promote resilience to global change Although some ecosystem responses to climate change are gradual, many ecosystems react in highly nonlinear ways. They show little response until a threshold or tipping point is reached where even a small perturbation may trigger collapse into a state from which recovery is difficult (1). Increasing evidence shows that the critical climate level for such collapse may be altered by conditions that can be managed locally. These synergies between local stressors and climate change provide potential opportunities for proactive management. Although their clarity and scale make such local approaches more conducive to action than global greenhouse gas management, crises in iconic UNESCO World Heritage sites illustrate that such stewardship is at risk of failing.

Toxicity of reduced nitrogen in eelgrass (Zostera marina) is highly dependent on shoot density and pH

In sheltered, eutrophicated estuaries, reduced nitrogen (NHx), and pH levels in the water layer can be greatly enhanced. In laboratory experiments, we studied the interactive effects of NHx, pH, and shoot density on the physiology and survival of eelgrass (Zostera marina). We tested long-term tolerance to NHx at pH 8 in a 5-week experiment. Short-term tolerance was tested for two shoot densities at both pH 8 and 9 in a 5-day experiment. At pH 8, eelgrass accumulated nitrogen as free amino acids when exposed to high loads of NHx, but showed no signs of necrosis. Low shoot density treatments became necrotic within days when exposed to NHx at pH 9. Increased NH3 intrusion and carbon limitation seemed to be the cause of this, as intracellular NHx could no longer be assimilated. Remarkably, experiments with high shoot densities at pH 9 showed hardly any necrosis, as the plants seemed to be able to alleviate the toxic effects of high NHx loads through joint NHx uptake. Our results suggest that NHx toxicity can be important in worldwide observed seagrass mass mortalities. We argue that the mitigating effect of high seagrass biomass on NHx toxicity is a positive feedback mechanism, potentially leading to alternative stable states in field conditions.

Emergent neutrality leads to multimodal species abundance distributions

Recent analyses of data sampled in communities ranging from corals and fossil brachiopods to birds and phytoplankton suggest that their species abundance distributions have multiple modes, a pattern predicted by none of the existing theories. Here we show that the multimodal pattern is consistent with predictions from the theory of emergent neutrality. This adds to the observations, suggesting that natural communities may be shaped by the evolutionary emergence of groups of similar species that coexist in niches. Such self-organized similarity unifies niche and neutral theories of biodiversity.

Resonance of Plankton Communities with Temperature Fluctuations

The interplay between intrinsic population dynamics and environmental variation is still poorly understood. It is known, however, that even mild environmental noise may induce large fluctuations in population abundances. This is due to a resonance effect that occurs in communities on the edge of stability. Here, we use a simple predator-prey model to explore the sensitivity of plankton communities to stochastic environmental fluctuations. Our results show that the magnitude of resonance depends on the timescale of intrinsic population dynamics relative to the characteristic timescale of the environmental fluctuations. Predator-prey communities with an intrinsic tendency to oscillate at a period T are particularly responsive to red noise characterized by a timescale of . We compare these theoretical predictions with the timescales of temperature fluctuations measured in lakes and oceans. This reveals that plankton communities will be highly sensitive to natural temperature fluctuations. More specifically, we demonstrate that the relatively fast temperature fluctuations in shallow lakes fall largely within the range to which rotifers and cladocerans are most sensitive, while marine copepods and krill will tend to resonate more strongly with the slower temperature variability of the open ocean.

Local Facilitation May Cause Tipping Points on a Landscape Level Preceded by Early-Warning Indicators

Positive biotic interactions play a significant role in shaping ecological communities. We used an individual-based model to demonstrate that plant facilitation on a microscale may cause ecosystem shifts on a landscape scale that can be announced by generic early-warning indicators. Recruitment of woody plants in harsh environments such as drylands often depends on nurse plants that ameliorate stressful conditions and facilitate the establishment of seedlings under their canopy. We found that these facilitative interactions may cause a treeless and a woodland state to be alternative stable states on a landscape scale if nurse plant effects are strong and if the environment is harsh enough to make facilitation necessary for seedling survival. A corollary is that under such conditions environmental change can bring drylands to tipping points for woody plant encroachment or woodland collapse. We show that the proximity of tipping points may be indicated by slowness of recovery of woody vegetation cover from small perturbations as well as by elevated temporal and spatial autocorrelation and variance. These signs are known to be indicators of critical slowing down. This is the first demonstration that the systemic phenomena of tipping points, announced by critical slowing down as a warning signal, may plausibly arise from microscale individual interactions, such as plant facilitation.