How nutrient retention and TN:TP ratios depend on ecosystem state in thousands of Chinese lakes.

Worldwide, anthropogenic activities threaten surface water quality by aggravating eutrophication and increasing total nitrogen to total phosphorus (TN:TP) ratios. In hydrologically connected systems, water quality management may benefit from in-ecosystem nutrient retention by preventing nutrient transport to downstream systems. However, nutrient retention may also alter TN:TP ratios with unforeseen consequences for downstream water quality. Here, we aim to increase understanding of how nutrient retention may influence nutrient transport to downstream systems to improve long-term water quality management. We analyzed lake ecosystem state, in-lake nutrient retention, and nutrient transport (ratios) for 3482 Chinese lakes using the lake process-based ecosystem model PCLake+. We compared a low climate change and sustainability-, and a high climate change and economy-focused scenario for 2050 against 2012. In both scenarios, the effect of nutrient input reduction outweighs that of temperature rise, resulting in more lakes with good ecological water quality (i.e., macrophyte-dominated) than in 2012. Generally, the sustainability-focused scenario shows a more promising future for water quality than the economy-focused scenario. Nevertheless, most lakes remain phytoplankton-dominated. The shift to more macrophyte-dominated lakes in 2050 is accompanied by higher nutrient retention fractions and less nutrient transport to downstream waterbodies. In-lake nutrient retention also alters the water's TN:TP ratio, depending on the inflow TN:TP ratio and the ecosystem state. In 2050 higher TN:TP ratios are expected in the outflows of lakes than in 2012, especially for the sustainability-focused scenario with strong TP loading reduction. However, the downstream impact of increased TN:TP ratios depends on actual nutrient loadings and the limiting nutrient in the receiving system. We conclude that nutrient input reductions, improved water quality, higher in-lake nutrient retention fractions, and lower nutrient transport to downstream waterbodies go hand in hand. Therefore, water quality management could benefit even more from nutrient pollution reduction than one would expect at first sight.

Regime shifts in shallow lakes explained by critical turbidity.

Worldwide, water quality managers target a clear, macrophyte-dominated state over a turbid, phytoplankton-dominated state in shallow lakes. The competition mechanisms underlying these ecological states were explored in the 1990s, but the concept of critical turbidity seems neglected in contemporary water quality models. In particular, a simple mechanistic model of alternative stable states in shallow lakes accounting for resource competition mechanisms and critical turbidity is lacking. To this end, we combined Scheffer's theory on critical turbidity with insights from nutrient and light competition theory founded by Tilman, Huisman and Weissing. This resulted in a novel graphical and mathematical model, GPLake-M, that is relatively simple and mechanistically understandable and yet captures the essential mechanisms leading to alternative stable states in shallow lakes. The process-based PCLake model was used to parameterize the model parameters and to test GPLake-M using a pattern-oriented strategy. GPLake-M's application range and position in the model spectrum are discussed. We believe that our results support the fundamental understanding of regime shifts in shallow lakes and provide a starting point for further mechanistic and management-focused explorations and model development. Furthermore, the concept of critical turbidity and the relation between light-limited submerged macrophytes and nutrient-limited phytoplankton might provide a new focus for empirical aquatic ecological research and water quality monitoring programs.

Bridging Theories for Ecosystem Stability Through Structural Sensitivity Analysis of Ecological Models in Equilibrium.

Ecologists are challenged by the need to bridge and synthesize different approaches and theories to obtain a coherent understanding of ecosystems in a changing world. Both food web theory and regime shift theory shine light on mechanisms that confer stability to ecosystems, but from different angles. Empirical food web models are developed to analyze how equilibria in real multi-trophic ecosystems are shaped by species interactions, and often include linear functional response terms for simple estimation of interaction strengths from observations. Models of regime shifts focus on qualitative changes of equilibrium points in a slowly changing environment, and typically include non-linear functional response terms. Currently, it is unclear how the stability of an empirical food web model, expressed as the rate of system recovery after a small perturbation, relates to the vulnerability of the ecosystem to collapse. Here, we conduct structural sensitivity analyses of classical consumer-resource models in equilibrium along an environmental gradient. Specifically, we change non-proportional interaction terms into proportional ones, while maintaining the equilibrium biomass densities and material flux rates, to analyze how alternative model formulations shape the stability properties of the equilibria. The results reveal no consistent relationship between the stability of the original models and the proportionalized versions, even though they describe the same biomass values and material flows. We use these findings to critically discuss whether stability analysis of observed equilibria by empirical food web models can provide insight into regime shift dynamics, and highlight the challenge of bridging alternative modelling approaches in ecology and beyond.

Will legal international rhino horn trade save wild rhino populations?

Wild vertebrate populations all over the globe are in decline, with poaching being the second-most-important cause. The high poaching rate of rhinoceros may drive these species into extinction within the coming decades. Some stakeholders argue to lift the ban on international rhino horn trade to potentially benefit rhino conservation, as current interventions appear to be insufficient. We reviewed scientific and grey literature to scrutinize the validity of reasoning behind the potential benefit of legal horn trade for wild rhino populations. We identified four mechanisms through which legal trade would impact wild rhino populations, of which only the increased revenue for rhino farmers could potentially benefit rhino conservation. Conversely, the global demand for rhino horn is likely to increase to a level that cannot be met solely by legal supply. Moreover, corruption is omnipresent in countries along the trade routes, which has the potential to negatively affect rhino conservation. Finally, programmes aimed at reducing rhino horn demand will be counteracted through trade legalization by removing the stigma on consuming rhino horn. Combining these insights and comparing them with criteria for sustainable wildlife farming, we conclude that legalizing rhino horn trade will likely negatively impact the remaining wild rhino populations. To preserve rhino species, we suggest to prioritize reducing corruption within rhino horn trade, increasing the rhino population within well-protected 'safe havens' and implementing educational programmes and law enforcement targeted at rhino horn consumers.

A Generically Parameterized model of Lake eutrophication (GPLake) that links field-, lab- and model-based knowledge.

Worldwide, eutrophication is threatening lake ecosystems. To support lake management numerous eutrophication models have been developed. Diverse research questions in a wide range of lake ecosystems are addressed by these models. The established models are based on three key approaches: the empirical approach that employs field surveys, the theoretical approach in which models based on first principles are tested against lab experiments, and the process-based approach that uses parameters and functions representing detailed biogeochemical processes. These approaches have led to an accumulation of field-, lab- and model-based knowledge, respectively. Linking these sources of knowledge would benefit lake management by exploiting complementary information; however, the development of a simple tool that links these approaches was hampered by their large differences in scale and complexity. Here we propose a Generically Parameterized Lake eutrophication model (GPLake) that links field-, lab- and model-based knowledge and can be used to make a first diagnosis of lake water quality. We derived GPLake from consumer-resource theory by the principle that lacustrine phytoplankton is typically limited by two resources: nutrients and light. These limitations are captured in two generic parameters that shape the nutrient to chlorophyll-a relations. Next, we parameterized GPLake, using knowledge from empirical, theoretical, and process-based approaches. GPLake generic parameters were found to scale in a comparable manner across data sources. Finally, we show that GPLake can be applied as a simple tool that provides lake managers with a first diagnosis of the limiting factor and lake water quality, using only the parameters for lake depth, residence time and current nutrient loading. With this first-order assessment, lake managers can easily assess measures such as reducing nutrient load, decreasing residence time or changing depth before spending money on field-, lab- or model- experiments to support lake management.

Success of lake restoration depends on spatial aspects of nutrient loading and hydrology.

Many aquatic ecosystems have deteriorated due to human activities and their restoration is often troublesome. It is proposed here that the restoration success of deteriorated lakes critically depends on hitherto largely neglected spatial heterogeneity in nutrient loading and hydrology. A modelling approach is used to study this hypothesis by considering four lake types with contrasting nutrient loading (point versus diffuse) and hydrology (seepage versus drainage). By comparing the longterm effect of common restoration measures (nutrient load reduction, lake flushing or biomanipulation) in these four lake types, we found that restoration through reduction of nutrient loading is effective in all cases. In contrast, biomanipulation only works in seepage lakes with diffuse nutrient inputs, while lake flushing will even be counterproductive in lakes with nutrient point sources. The main conclusion of the presented analysis is that a priori assessment of spatial heterogeneity caused by nutrient loading and hydrology is essential for successful restoration of lake ecosystems.

Modelling induced bank filtration effects on freshwater ecosystems to ensure sustainable drinking water production.

Induced bank filtration (IBF) is a water abstraction technology using different natural infiltration systems for groundwater recharge, such as river banks and lake shores. It is a cost-effective pre-treatment method for drinking water production used in many regions worldwide, predominantly in urban areas. Until now, research concerning IBF has almost exclusively focussed on the purification efficiency and infiltration capacity. Consequently, knowledge about the effects on source water bodies is lacking. Yet, IBF interrupts groundwater seepage and affects processes in the sediment potentially resulting in adverse effects on lake or river water quality. Securing sufficient source water quality, however, is important for a sustainable drinking water production by IBF. In this study, we analysed the effects of five predicted mechanisms of IBF on shallow lake ecosystems using the dynamic model PCLake: declining CO2 and nutrient availability, as well as increasing summer water temperatures, sedimentation rates and oxygen penetration into sediments. Shallow lake ecosystems are abundant worldwide and characterised by the occurrence of alternative stable states with either clear water and macrophyte dominance or turbid, phytoplankton-dominated conditions. Our results show that IBF in most scenarios increased phytoplankton abundance and thus had adverse effects on shallow lake water quality. Threshold levels for critical nutrient loading inducing regime shifts from clear to turbid conditions were up to 80% lower with IBF indicating a decreased resilience to eutrophication. The effects were strongest when IBF interrupted the seepage of CO2 rich groundwater resulting in lower macrophyte growth. IBF could also enhance water quality, but only when interrupting the seepage of groundwater with high nutrient concentrations. Higher summer water temperatures increased the share of cyanobacteria in the phytoplankton community and thus the risk of toxin production. In relative terms, the effects of changing sedimentation rates and oxygen penetration were small. Lake depth and size influenced the effect of IBF on critical nutrient loads, which was strongest in shallower and smaller lakes. Our model results stress the need of a more comprehensive ecosystem perspective including an assessment of IBF effects on threshold levels for regime shifts to prevent high phytoplankton abundance in the source water body and secure a sustainable drinking water supply.

Response of Submerged Macrophyte Communities to External and Internal Restoration Measures in North Temperate Shallow Lakes.

Submerged macrophytes play a key role in north temperate shallow lakes by stabilizing clear-water conditions. Eutrophication has resulted in macrophyte loss and shifts to turbid conditions in many lakes. Considerable efforts have been devoted to shallow lake restoration in many countries, but long-term success depends on a stable recovery of submerged macrophytes. However, recovery patterns vary widely and remain to be fully understood. We hypothesize that reduced external nutrient loading leads to an intermediate recovery state with clear spring and turbid summer conditions similar to the pattern described for eutrophication. In contrast, lake internal restoration measures can result in transient clear-water conditions both in spring and summer and reversals to turbid conditions. Furthermore, we hypothesize that these contrasting restoration measures result in different macrophyte species composition, with added implications for seasonal dynamics due to differences in plant traits. To test these hypotheses, we analyzed data on water quality and submerged macrophytes from 49 north temperate shallow lakes that were in a turbid state and subjected to restoration measures. To study the dynamics of macrophytes during nutrient load reduction, we adapted the ecosystem model PCLake. Our survey and model simulations revealed the existence of an intermediate recovery state upon reduced external nutrient loading, characterized by spring clear-water phases and turbid summers, whereas internal lake restoration measures often resulted in clear-water conditions in spring and summer with returns to turbid conditions after some years. External and internal lake restoration measures resulted in different macrophyte communities. The intermediate recovery state following reduced nutrient loading is characterized by a few macrophyte species (mainly pondweeds) that can resist wave action allowing survival in shallow areas, germinate early in spring, have energy-rich vegetative propagules facilitating rapid initial growth and that can complete their life cycle by early summer. Later in the growing season these plants are, according to our simulations, outcompeted by periphyton, leading to late-summer phytoplankton blooms. Internal lake restoration measures often coincide with a rapid but transient colonization by hornworts, waterweeds or charophytes. Stable clear-water conditions and a diverse macrophyte flora only occurred decades after external nutrient load reduction or when measures were combined.

Multimedia fate modeling of perfluorooctanoic acid (PFOA) and perfluorooctane sulphonate (PFOS) in the shallow lake Chaohu, China.

Freshwater shallow lake ecosystems provide valuable ecological services to human beings. However, these systems are subject to severe contamination from anthropogenic sources. Per- and polyfluoroalkyl substances (PFASs), including perfluorooctanoic acid (PFOA) and perfluorooctane sulphonate (PFOS), are among the contaminants that have received substantial attention, primarily due to abundant applications, environment persistence, and potential threats to ecological and human health. Understanding the environmental behavior of these contaminants in shallow freshwater lake environments using a modeling approach is therefore critical. Here, we characterize the fate, transport and transformation of both PFOA and PFOS in the fifth largest freshwater lake in China (Chaohu) during a two-year period (2013-2015) using a fugacity-based multimedia fate model. A reasonable agreement between the measured and modeled concentrations in various compartments confirms the model's reliability. The model successfully quantifies the environmental processes and identifies the major sources and input pathways of PFOA and PFOS to the Chaohu water body. Sensitivity analysis reveals the critical role of nonlinear Freundlich sorption, which contributes to a variable fraction of the model true uncertainty in different compartments (8.1%-93.6%). Through additional model scenario analyses, we further elucidate the importance of nonlinear Freundlich sorption that is essential for the reliable model performance. We also reveal the distinct composition of emission sources for the two contaminants, as the major sources are indirect soil volatilization and direct release from human activities for PFOA and PFOS, respectively. The present study is expected to provide implications for local management of PFASs pollution in Lake Chaohu and to contribute to developing a general model framework for the evaluation of PFASs in shallow lakes.

Spatial identification of critical nutrient loads of large shallow lakes: Implications for Lake Taihu (China).

Ongoing eutrophication frequently causes toxic phytoplankton blooms. This induces huge worldwide challenges for drinking water quality, food security and public health. Of crucial importance in avoiding and reducing blooms is to determine the maximum nutrient load ecosystems can absorb, while remaining in a good ecological state. These so called critical nutrient loads for lakes depend on the shape of the load-response curve. Due to spatial variation within lakes, load-response curves and therefore critical nutrient loads could vary throughout the lake. In this study we determine spatial patterns in critical nutrient loads for Lake Taihu (China) with a novel modelling approach called Spatial Ecosystem Bifurcation Analysis (SEBA). SEBA evaluates the impact of the lake's total external nutrient load on the local lake dynamics, resulting in a map of critical nutrient loads for different locations throughout the lake. Our analysis shows that the largest part of Lake Taihu follows a nonlinear load-response curve without hysteresis. The corresponding critical nutrient loads vary within the lake and depend on management goals, i.e. the maximum allowable chlorophyll concentration. According to our model, total nutrient loads need to be more than halved to reach chlorophyll-a concentrations of 30-40 µg L-1 in most sections of the lake. To prevent phytoplankton blooms with 20 µg L-1 chlorophyll-a throughout Lake Taihu, both phosphorus and nitrogen loads need a nearly 90% reduction. We conclude that our approach is of great value to determine critical nutrient loads of lake ecosystems such as Taihu and likely of spatially heterogeneous ecosystems in general.

Integrated ecological and chemical food web accumulation modeling explains PAH temporal trends during regime shifts in a shallow lake.

Shallow lakes can switch suddenly from a turbid situation with high concentrations of phytoplankton and other suspended solids to a vegetated state with clear water, and vice versa. These alternative stable states may have a substantial impact on the fate of hydrophobic organic compounds (HOCs). Models that are fit to simulate impacts from these complex interactions are scarce. We developed a contaminant fate model which is linked to an ecosystem model (PCLake) for shallow lakes. This integrated model was successful in simulating long-term dynamics (1953-2012) of representative polycyclic aromatic hydrocarbons (PAHs) in the main biotic and abiotic components in a large shallow lake (Chaohu in China), which has undergone regime shifts in this period. Historical records from sediment cores were used to evaluate the model. The model revealed that regime shifts in shallow lakes had a strong impact on the fate of less hydrophobic compounds due to the large storage capacity of macrophytes, which accumulated up to 55.6% of phenanthrene in the clear state. The abrupt disappearance of macrophytes after the regime shift resulted in a sudden change in phenanthrene distribution, as the sediment became the major sink. For more hydrophobic compounds such as benzo(a)pyrene, the modeled impact of the regime shift was negligible for the whole environment, yet large for biotic compartments. This study is the first to provide a full mechanistic analysis of the impact of regime shifts on the fate of PAHs in a real lake ecosystem.

Mowing Submerged Macrophytes in Shallow Lakes with Alternative Stable States: Battling the Good Guys?

Submerged macrophytes play an important role in maintaining good water quality in shallow lakes. Yet extensive stands easily interfere with various services provided by these lakes, and harvesting is increasingly applied as a management measure. Because shallow lakes may possess alternative stable states over a wide range of environmental conditions, designing a successful mowing strategy is challenging, given the important role of macrophytes in stabilizing the clear water state. In this study, the integrated ecosystem model PCLake is used to explore the consequences of mowing, in terms of reducing nuisance and ecosystem stability, for a wide range of external nutrient loadings, mowing intensities and timings. Elodea is used as a model species. Additionally, we use PCLake to estimate how much phosphorus is removed with the harvested biomass, and evaluate the long-term effect of harvesting. Our model indicates that mowing can temporarily reduce nuisance caused by submerged plants in the first weeks after cutting, particularly when external nutrient loading is fairly low. The risk of instigating a regime shift can be tempered by mowing halfway the growing season when the resilience of the system is highest, as our model showed. Up to half of the phosphorus entering the system can potentially be removed along with the harvested biomass. As a result, prolonged mowing can prevent an oligo-to mesotrophic lake from becoming eutrophic to a certain extent, as our model shows that the critical nutrient loading, where the lake shifts to the turbid phytoplankton-dominated state, can be slightly increased.

Hydrological regulation drives regime shifts: evidence from paleolimnology and ecosystem modeling of a large shallow Chinese lake.

Quantitative evidence of sudden shifts in ecological structure and function in large shallow lakes is rare, even though they provide essential benefits to society. Such 'regime shifts' can be driven by human activities which degrade ecological stability including water level control (WLC) and nutrient loading. Interactions between WLC and nutrient loading on the long-term dynamics of shallow lake ecosystems are, however, often overlooked and largely underestimated, which has hampered the effectiveness of lake management. Here, we focus on a large shallow lake (Lake Chaohu) located in one of the most densely populated areas in China, the lower Yangtze River floodplain, which has undergone both WLC and increasing nutrient loading over the last several decades. We applied a novel methodology that combines consistent evidence from both paleolimnological records and ecosystem modeling to overcome the hurdle of data insufficiency and to unravel the drivers and underlying mechanisms in ecosystem dynamics. We identified the occurrence of two regime shifts: one in 1963, characterized by the abrupt disappearance of submerged vegetation, and another around 1980, with strong algal blooms being observed thereafter. Using model scenarios, we further disentangled the roles of WLC and nutrient loading, showing that the 1963 shift was predominantly triggered by WLC, whereas the shift ca. 1980 was attributed to aggravated nutrient loading. Our analysis also shows interactions between these two stressors. Compared to the dynamics driven by nutrient loading alone, WLC reduced the critical P loading and resulted in earlier disappearance of submerged vegetation and emergence of algal blooms by approximately 26 and 10 years, respectively. Overall, our study reveals the significant role of hydrological regulation in driving shallow lake ecosystem dynamics, and it highlights the urgency of using multi-objective management criteria that includes ecological sustainability perspectives when implementing hydrological regulation for aquatic ecosystems around the globe.

An Integrated Coral Reef Ecosystem Model to Support Resource Management under a Changing Climate.

Millions of people rely on the ecosystem services provided by coral reefs, but sustaining these benefits requires an understanding of how reefs and their biotic communities are affected by local human-induced disturbances and global climate change. Ecosystem-based management that explicitly considers the indirect and cumulative effects of multiple disturbances has been recommended and adopted in policies in many places around the globe. Ecosystem models give insight into complex reef dynamics and their responses to multiple disturbances and are useful tools to support planning and implementation of ecosystem-based management. We adapted the Atlantis Ecosystem Model to incorporate key dynamics for a coral reef ecosystem around Guam in the tropical western Pacific. We used this model to quantify the effects of predicted climate and ocean changes and current levels of current land-based sources of pollution (LBSP) and fishing. We used the following six ecosystem metrics as indicators of ecosystem state, resilience and harvest potential: 1) ratio of calcifying to non-calcifying benthic groups, 2) trophic level of the community, 3) biomass of apex predators, 4) biomass of herbivorous fishes, 5) total biomass of living groups and 6) the end-to-start ratio of exploited fish groups. Simulation tests of the effects of each of the three drivers separately suggest that by mid-century climate change will have the largest overall effect on this suite of ecosystem metrics due to substantial negative effects on coral cover. The effects of fishing were also important, negatively influencing five out of the six metrics. Moreover, LBSP exacerbates this effect for all metrics but not quite as badly as would be expected under additive assumptions, although the magnitude of the effects of LBSP are sensitive to uncertainty associated with primary productivity. Over longer time spans (i.e., 65 year simulations), climate change impacts have a slight positive interaction with other drivers, generally meaning that declines in ecosystem metrics are not as steep as the sum of individual effects of the drivers. These analyses offer one way to quantify impacts and interactions of particular stressors in an ecosystem context and so provide guidance to managers. For example, the model showed that improving water quality, rather than prohibiting fishing, extended the timescales over which corals can maintain high abundance by at least 5-8 years. This result, in turn, provides more scope for corals to adapt or for resilient species to become established and for local and global management efforts to reduce or reverse stressors.

Food-web stability signals critical transitions in temperate shallow lakes.

A principal aim of ecologists is to identify critical levels of environmental change beyond which ecosystems undergo radical shifts in their functioning. Both food-web theory and alternative stable states theory provide fundamental clues to mechanisms conferring stability to natural systems. Yet, it is unclear how the concept of food-web stability is associated with the resilience of ecosystems susceptible to regime change. Here, we use a combination of food web and ecosystem modelling to show that impending catastrophic shifts in shallow lakes are preceded by a destabilizing reorganization of interaction strengths in the aquatic food web. Analysis of the intricate web of trophic interactions reveals that only few key interactions, involving zooplankton, diatoms and detritus, dictate the deterioration of food-web stability. Our study exposes a tight link between food-web dynamics and the dynamics of the whole ecosystem, implying that trophic organization may serve as an empirical indicator of ecosystem resilience.

Competition for light and nutrients in layered communities of aquatic plants.

Dominance of free-floating plants poses a threat to biodiversity in many freshwater ecosystems. Here we propose a theoretical framework to understand this dominance, by modeling the competition for light and nutrients in a layered community of floating and submerged plants. The model shows that at high supply of light and nutrients, floating plants always dominate due to their primacy for light, even when submerged plants have lower minimal resource requirements. The model also shows that floating-plant dominance cannot be an alternative stable state in light-limited environments but only in nutrient-limited environments, depending on the plants' resource consumption traits. Compared to unlayered communities, the asymmetry in competition for light-coincident with symmetry in competition for nutrients-leads to fundamentally different results: competition outcomes can no longer be predicted from species traits such as minimal resource requirements ([Formula: see text] rule) and resource consumption. Also, the same two species can, depending on the environment, coexist or be alternative stable states. When applied to two common plant species in temperate regions, both the model and field data suggest that floating-plant dominance is unlikely to be an alternative stable state.

Pattern formation at multiple spatial scales drives the resilience of mussel bed ecosystems.

Self-organized complexity at multiple spatial scales is a distinctive characteristic of biological systems. Yet, little is known about how different self-organizing processes operating at different spatial scales interact to determine ecosystem functioning. Here we show that the interplay between self-organizing processes at individual and ecosystem level is a key determinant of the functioning and resilience of mussel beds. In mussel beds, self-organization generates spatial patterns at two characteristic spatial scales: small-scale net-shaped patterns due to behavioural aggregation of individuals, and large-scale banded patterns due to the interplay of between-mussel facilitation and resource depletion. Model analysis reveals that the interaction between these behavioural and ecosystem-level mechanisms increases mussel bed resilience, enables persistence under deteriorating conditions and makes them less prone to catastrophic collapse. Our analysis highlights that interactions between different forms of self-organization at multiple spatial scales may enhance the intrinsic ability of ecosystems to withstand both natural and human-induced disturbances.

Community stoichiometry in a changing world: combined effects of warming and eutrophication on phytoplankton dynamics.

The current changes in our climate will likely have far-reaching consequences for aquatic ecosystems. These changes in the climate, however, do not act alone, and are often accompanied by additional stressors such as eutrophication. Both global warming and eutrophication have been shown to affect the timing and magnitude of phytoplankton blooms. Little is known about the combined effects of rising temperatures and eutrophication on the stoichiometry of entire phytoplankton communities. We exposed a natural phytoplankton spring community to different warming and phosphorus-loading scenarios using a full-factorial design. Our results demonstrate that rising temperatures promote the growth rate of an entire phytoplankton community. Furthermore, both rising temperatures and phosphorus loading stimulated the maximum biomass built up by the phytoplankton community. Rising temperatures led to higher carbon: nutrient stoichiometry of the phytoplankton community under phosphorus-limited conditions. Such a shift towards higher carbon: nutrient ratios, in combination with a higher biomass buildup, suggests a temperature-driven increase in nutrient use efficiency, the phytoplankton community. Importantly, with higher carbon: nutrient stoichiometry, phytoplankton is generally of poorer nutritional value for zooplankton. Thus, although warming may result in higher phytoplankton biomass, this may be accompanied by a stoichiometric mismatch between phytoplankton and their grazers, with possible consequences for the entire aquatic food web.

Alternative stable states and alternative endstates of community assembly through intra- and interspecific positive and negative interactions.

Positive and negative interactions within and between species may occur simultaneously, with the net effect depending on population densities. For instance, at low densities plants may ameliorate stress, while competition for resources dominates at higher densities. Here, we propose a simple two-species model in which con- and heterospecifics have a positive effect on per capita growth rate at low densities, while negative interactions dominate at high densities. The model thus includes both Allee effects (intraspecific positive effects) and mutualism (interspecific positive effects), as well as intra- and interspecific competition. Using graphical methods we derive conditions for alternative stable states and species coexistence. We show that mutual non-invasibility (i.e. the inability of each species to invade a population of the other) is more likely when species have a strong positive effect on the own species or a strong negative effect on the other species. Mutual non-invasibility implies alternative stable states, however, there may also be alternative stable states at which species coexist. In the case of species symmetry (i.e. when species are indistinguishable), such alternative coexistence states require that if the positive effect exerted at low densities at the own species is stronger than on the other species, the negative effect at higher densities is also stronger on the own species than on the other species, or, vice versa, if the interspecific positive effects at low densities are stronger than the intraspecific effects, the negative effects at higher densities are also stronger between species than within species. However, the reachability of alternative stable states is restricted by the frequency and density at which species are introduced during community assembly, so that alternative stable states do not always represent alternative endstates of community assembly.

Plant functional types define magnitude of drought response in peatland CO2 exchange.

Peatlands are important sinks for atmospheric carbon (C), yet the role of plant functional types (PFTs) for C sequestration under climatic perturbations is still unclear. A plant-removal experiment was used to study the importance of vascular PFTs for the net ecosystem CO2 exchange (NEE) during (i.e., resistance) and after (i.e., recovery) an experimental drought. The removal of PFTs caused a decrease of NEE, but the rate differed between microhabitats (i.e., hummocks and lawns) and the type of PFTs. Ericoid removal had a large effect on NEE in hummocks, while the graminoids played a major role in the lawns. The removal of PFTs did not affect the resistance or the recovery after the experimental drought. We argue that the response of Sphagnum mosses (the only PFT present in all treatments) to drought is dominant over that of coexisting PFTs. However, we observed that the moment in time when the system switched from C sink to C source during the drought was controlled by the vascular PFTs. In the light of climate change, the shifts in species composition or even the loss of certain PFTs are expected to strongly affect the future C dynamics in response to environmental stress.