Helmut Hillebrand

• Carl von Ossietzky University of Oldenburg

The elemental content of life is a key trait shaping ecology and evolution, yet organismal stoichiometry has largely been studied on a case-by-case basis. This limitation has hindered our ability to identify broad patterns and mechanisms across taxa and ecosystems. To address this, we present StoichLife, a global dataset of 28,049 records from 5,87...

Nutrients and light are major resources controlling growth, biomass, and community structure of phytoplankton. When looking at those resources individually, resource uptake and biochemical transformation, and thereby also the demand for resources, have been shown to be temperature‐dependent. However, there is still a lack of understanding of how te...

Turnover in species composition through time is a dominant form of biodiversity change, which has profound effects on the functioning of ecological communities1, 2, 3–4. Turnover rates differ markedly among communities⁴, but the drivers of this variation across taxa and realms remain unknown. Here we analyse 42,255 time series of species compositio...

Ecosystems worldwide are experiencing a range of natural and anthropogenic disturbances, many of which are intensifying as global change accelerates. Ecological responses to those disturbances are determined by both the vulnerabilities of species and their interspecific interactions. Understanding how individual species contribute to the (in‐)stabi...

Ecological stability is a vital component of natural ecosystems that can inform effective conservation and ecosystem management. Furthermore, there is increasing interest in making comparisons of stability values across sites, systems and taxonomic groups, often using comparative synthetic approaches, such as meta‐analysis. However, these synthetic...

Temperature and resource availability are pivotal factors influencing phytoplankton community structures. Numerous prior studies demonstrated their significant influence on phytoplankton stoichiometry, cell size, and growth rates. The growth rate, serving as a reflection of an organism's success within its environment, is linked to stoichiometry an...

Biodiversity is temporally dynamic, reflecting historical environmental conditions and influencing ecosystem stability. Colonisation and extinction dynamics frequently exhibit asynchronous patterns, resulting in net imbalances and thus to long‐lasting richness trends. If these trends are not functionally random, functional net imbalances between co...

Biodiversity generally increases productivity in ecosystems; however, this is mediated by the specific functional traits that come with biodiversity loss or gain and how these traits interact with environmental conditions. Most biodiversity studies evaluate the effects of species richness alone, despite our increasing understanding that intraspecif...

Zusammenfassung Der Biodiversitätsverlust schreitet in bedrohlichem Ausmaß voran. Mit dem Global Biodiversity Framework und voraussichtlich dem Nature Restoration Law bestehen nun auf internationaler und europäischer Ebene vielversprechende Ansätze, ihm Herr zu werden. Jetzt ist der Bundesgesetzgeber – nicht zuletzt aus verfassungsrechtlichen Erwäg...

Animal stoichiometry affects fundamental processes ranging from organismal physiology to global element cycles. However, it is unknown whether animal stoichiometry follows predictable scaling relationships with body mass and whether adaptation to life on land or water constrains patterns of elemental allocation. To test both interspecific and intra...

Many of the global challenges that confront humanity are interlinked in a dynamic complex network, with multiple feedback loops, nonlinear interactions and interdependencies that make it difficult, if not impossible, to consider individual threats in isolation. These challenges are mainly dealt with, however, by considering individual threats in is...

One of the key challenges in managing eutrophication in coastal marine ecosystems is the harmonized cross-border assessment of phytoplankton. Some general understanding of the consequences of shifting nutrient regimes can be derived from the detailed investigation of the phytoplankton community and its biodiversity. Here, we combined long-term moni...

Compositional measurements from species assemblages define a high dimensional dataspace in which the data can form complex structures, termed manifolds. Comparing assemblages in this dataspace is difficult because the data is often sparse relative to its dimensionality and the complex structure of the manifold introduces bias and error in measureme...

Thresholds and tipping points are frequently used concepts to address the risks of global change pressures and their mitigation. It is tempting to also consider them to understand biodiversity change and design measures to ensure biotic integrity. Here, we argue that thresholds and tipping points do not work well in the context of biodiversity chan...

The discrepancy between global loss and local constant species richness has led to debates over data quality, systematic biases in monitoring programmes and the adequacy of species richness to capture changes in biodiversity. We show that, more fundamentally, null expectations of stable richness can be wrong, despite independent yet equal colonizat...

One of the key challenges in managing eutrophication in coastal marine ecosystems is the harmonized cross-border assessment of phytoplankton. Some general understanding of the consequences of shifting nutrient regimes can be derived from the detailed investigation of the phytoplankton community and its biodiversity. Here, we combined long-term moni...

Introduction Any measure of ecological stability scales with the spatial and temporal extent of the data on which it is based. The magnitude of stabilization effects at increasing spatial scale is determined by the degree of synchrony between local and regional species populations. Methods We applied two recently developed approaches to quantify t...

The availability of underwater light, as primary energy source for all aquatic photoautotrophs, is (and will further be) altered by changing precipitation, water turbidity, mixing depth, and terrestrial input of chromophoric dissolved organic matter (CDOM). While experimental manipulations of CDOM input and turbidity are frequent, they often involv...

Due to the strong interconnectedness between the ocean and our societies worldwide, improved ocean governance is essential for sustainable development in the context of the UN Ocean Decade. However, a multitude of different perspectives—ecological, societal, political, economic—and relations between these have to be understood and taken into consid...

The twin crises of climate change and biodiversity loss define a strong need for functional diversity monitoring. While the availability of high-quality ecological monitoring data is increasing, the quantification of functional diversity so far requires the identification of species traits, for which data are harder to obtain. However, the traits t...

Primary consumers in aquatic ecosystems are frequently limited by the quality of their food, often expressed as phytoplankton elemental and biochemical composition. However, the effects of these food quality indicators vary across studies, and we lack an integrated understanding of how elemental (e.g. nitrogen, phosphorus) and biochemical (e.g. fat...

Biodiversity is expected to change in response to future global warming. However, it is difficult to predict how species will track the ongoing climate change. Here we use the fossil record of planktonic foraminifera to assess how biodiversity responded to climate change with a magnitude comparable to future anthropogenic warming. We compiled time...

Microphytobenthos (MPBs) are the main primary producers in shallow marine ecosystems, such as the Wadden Sea. We investigated the spatial and temporal dynamics of MPB communities across the marine-terrestrial boundary over three seasons (spring, summer, and fall) on three East Frisian Islands (Norderney, Spiekeroog, and Wangerooge) in the German Wa...

The current policy and goals aimed to conserve biodiversity and manage biodiversity change are often formulated at the global scale. At smaller scales however, biodiversity change is more nuanced leading to a plethora of trends in different metrics of alpha diversity and temporal turnover. Therefore, large‐scale policy targets do not translate easi...

Primary consumers in aquatic ecosystems are frequently limited by the quality of their food, often expressed as phytoplankton elemental and biochemical composition. Effects of these food quality indicators vary across studies, and the relative importance of elemental (nitrogen and phosphorus) versus biochemical (fatty acid and sterol) limitation in...

Primary consumers in aquatic ecosystems are frequently limited by the quality of their food, often expressed as phytoplankton elemental and biochemical composition. Effects of these food quality indicators vary across studies, and the relative importance of elemental (nitrogen and phosphorus) versus biochemical (fatty acid and sterol) limitation in...

While environmental science, and ecology in particular, is working to provide better understanding to base sustainable decisions on, the way scientific understanding is developed can at times be detrimental to this cause. Locked‐in debates are often unnecessarily polarised and can compromise any common goals of the opposing camps. The present paper...

Cell size is a master trait in the functional ecology of phytoplankton correlating with numerous morphological, physiological, and life‐cycle characteristics of species that constrain their nutrient use, growth, and edibility. In contrast to well‐known spatial patterns in cell size at macroecological scales or temporal changes in experimental conte...

Body size is a decisive functional trait in many organisms, especially for phytoplankton, which span several orders of magnitude in cell volume. Therefore, the analysis of size as a functional trait driving species’ performance has received wide attention in aquatic ecology, amended in recent decades by studies documenting changes in phytoplankton...

Anthropogenic climate change is altering global biogeographical patterns. However, it remains difficult to quantify how bioregions are changing because pre‐industrial records of species distributions are rare. Marine microfossils, such as planktonic foraminifera, are preserved in seafloor sediments and allow the quantification of bioregions in the...

Ecological stability encompasses multiple dimensions of functional and compositional responses to environmental change. Though no single stability dimension used in isolation can fully reflect the overall response to environmental change, a common vulnerability assessment that integrates simultaneously across multiple stability components is highly...

This paper provides a critical contribution to the UN Decade of Ocean Science for Sustainable Development 2021-2030, outlining some of the core challenges of sustainable ocean governance and management by reflecting on modes and logics of interactions between ocean-related sciences, policy-makers and practitioners. In this regard, we give an overvi...

Positive biodiversity–ecosystem functioning (BEF) relationships are predicted to increase in strength when high environmental variability allows for complementarity between resource use strategies in diverse communities. This environmental variability can be represented by spatial or temporal variation in nutrient ratios, but resource use efficienc...

Metacommunity ecology currently lacks a consistent functional trait perspective across trophic levels. To foster new cross‐taxa experiments and field studies, we present hypotheses on how three trait dimensions change along gradients of density of individuals, resource supply and habitat isolation. The movement dimension refers to the ability to mo...

The twin crises of climate change and biodiversity loss define a strong need for functional diversity monitoring. While the availability of high-quality ecological monitoring data is increasing, the quantification of functional diversity so far requires the identification of species traits, for which data is harder to obtain. However, the traits th...

For aquatic scientists mesocosm experiments are important tools for hypothesis testing as they offer a compromise between experimental control and realism. Here we present a new mesocosm infrastructure—SITES AquaNET—located in five lakes connected to field stations in Sweden that cover a ~760 km latitudinal gradient. SITES AquaNet overcomes major h...

Aim The aim was to evaluate the effects of climate warming on biodiversity across spatial scales (i.e., α‐, β‐ and γ‐diversity) and the effects of patch openness and experimental context on diversity responses. Location Global. Time period 1995–2017. Major taxa studied Fungi, invertebrates, phytoplankton, plants, seaweed, soil microbes and zoopl...

Increasing human impact on the environment is causing drastic changes in disturbance regimes and how they prevail over time. Of increasing relevance is to further our understanding on biological responses to pulse disturbances (short duration) and how they interact with other ongoing press disturbances (constantly present). Because the temporal and...

While environmental science, and ecology in particular, is working to provide better understanding to base sustainable decisions on, the way scientific understanding is developed can at times be detrimental to this cause. Locked-in debates are often unnecessarily polarized and can compromise any common goals of the opposing camps. The present paper...

To understand ecosystem responses to anthropogenic global change, a prevailing framework is the definition of threshold levels of pressure, above which response magnitudes and their variances increase disproportionately. However, we lack systematic quantitative evidence as to whether empirical data allow definition of such thresholds. Here, we summ...

Estimates of biodiversity change are essential for the management and conservation of ecosystems. Accurate estimates rely on selecting representative sites, but monitoring often focuses on sites of special interest. How such site‐selection biases influence estimates of biodiversity change is largely unknown. Site‐selection bias potentially occurs a...

The sea surface microlayer (SML) is the boundary layer between the ocean and the atmosphere and plays a unique role in marine biogeochemistry. Phytoplankton communities in this uppermost surface layer are exposed to extreme ultraviolet (UV) radiation and potentially high nutrient supplies. In order to understand the response of SML communities to s...

Pronounced atmospheric and oceanic warming along the West Antarctic Peninsula (WAP) has resulted in abundance shifts in populations of Antarctic krill and Salpa thompsoni determined by changes in the timing of sea-ice advance, the duration of sea-ice cover and food availability. Krill and salps represent the most important macrozooplankton grazers...

Aim Biodiversity dynamics comprise evolutionary and ecological changes on multiple temporal scales from millions of years to decades, but they are often interpreted within a single time frame. Planktonic foraminifera communities offer a unique opportunity for analysing the dynamics of marine biodiversity over different temporal scales. Our study ai...

A central concept for understanding the mechanisms linking diversity and primary production or more general, ecosystem functioning, is resource use efficiency (RUE). It quantifies the amount of biomass production over time relative to unit resource supplied, that is, represents a quota of matter use efficiency. Given anthropogenic alterations of bi...

Unfortunately, in the print version of this article the below appendix

Most ecosystems are affected by anthropogenic or natural pulse disturbances, which alter the community composition and functioning for a limited period of time. Whether and how quickly communities recover from such pulses is central to our understanding of biodiversity dynamics and ecosystem organisation, but also to nature conservation and managem...

Tidal flats are subject to dynamic processes, characterized by sedimentation and erosion, which become more important in times of climate change. To investigate impacts of elevation levels on ecosystem functioning, we focused on macrozoobenthos as an essential ecosystem component that links trophic levels and determines sediment structure. Therefor...

Planktonic primary consumers have been shown to strongly influence phytoplankton communities via top-down effects such as grazing and nutrient recycling. However, it remains unclear how changes in consumer richness may alter the stoichiometric constrains between producer and consumer assemblages. Here we test whether the stoichiometry of producer–c...

Marine systems outpace terrestrial habitats in biodiversity erosion

Nutrient availability and temperature are important drivers of phytoplankton growth and stoichiometry. However, the interactive effects of nutrients and temperature on phytoplankton have been analyzed mostly by addressing changes in average temperature, whereas recent evidence suggests an important role of temperature fluctuations. In a laboratory...

Identifying stable coexistence in empirical systems is notoriously difficult. Here, we show how spatiotemporal structure and complex system dynamics can confound two commonly used stability metrics in empirical contexts: response to perturbation and invasion rate when rare. We use these metrics to characterize stable coexistence across a range of s...

The relationship between biodiversity and ecosystem functioning (BEF) is a topic of considerable interest to scientists and managers because a better understanding of its underlying mechanisms may help us mitigate the consequences of biodiversity loss on ecosystems. Our current knowledge of BEF relies heavily on theoretical and experimental studies...

The ocean—the Earth’s largest ecosystem—is increasingly affected by anthropogenic climate change1,2. Large and globally consistent shifts have been detected in species phenology, range extension and community composition in marine ecosystems3–5. However, despite evidence for ongoing change, it remains unknown whether marine ecosystems have entered...

Changes in salinity are known to alter the morphology of protists, and we hypothesized that these changes subsequently alter also the predatory behavior of the dinoflagellate Oxyrrhis marina. Oxyrrhis was grown in media of 33, 25, 20, and 10% of the regular salinity of f/2 medium (31–32‰). In all cases, the cells discharged trichocysts and swelled....

Food-chain length (FCL) is a fundamental ecosystem attribute, integrating information on both food web composition and ecosystem processes. It remains untested whether FCL also reflects the history of community assembly known to affect community composition and ecosystem functioning. Here, we performed microcosm experiments with a copepod (top pred...

Resource use efficiency (RUE) is an ecological concept that measures the proportion of supplied resources, which is converted into new biomass, i.e., it relates realized to potential productivity. It is also commonly perceived as one of the main mechanisms linking biodiversity to ecosystem functioning based on the assumption that higher species num...

Concern about the functional consequences of unprecedented loss in biodiversity has prompted biodiversity-ecosystem functioning (BEF) research to become one of the most active fields of ecological research in the past 25 years. Hundreds of experiments have manipulated biodiversity as an independent variable and found compelling support that the fun...

Metaecosystem theory addresses the link between local (within habitats) and regional (between habitats) dynamics by simultaneously analyzing spatial community ecology and abiotic matter flow. Here we experimentally address how spatial resource gradients and connectivity affect resource use efficiency (RUE) and stoichiometry in marine phytoplankton...

Field experiments investigating biodiversity and ecosystem functioning require the observation of abiotic parameters, especially when carried out in the intertidal zone. An experiment for biodiversity–ecosystem functioning was set up in the intertidal zone of the back-barrier salt marsh of Spiekeroog Island in the German Bight. Here, we report the...

Reduced nitrogen species are key nutrients for biological productivity in the oceans. Ammonium is often present in low and growth-limiting concentrations, albeit peaks occur during collapse of algal blooms or via input from deep sea upwelling and riverine inflow. Autotrophic phytoplankton exploit ammonium peaks by storing nitrogen intracellularly....

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta dive...

While there is a lot of data on interactive effects of eutrophication and warming, to date, we lack data to generate reliable predictions concerning possible effects of nutrient decrease and temperature increase on community composition and functional responses. In recent years, a wide‐ranging trend of nutrient decrease (re‐oligotrophication) was r...

Field experiments investigating biodiversity and ecosystem functioning require observation of abiotic parameters, especially when carried out in the intertidal zone. An experiment for biodiversity-ecosystem functioning at the intersection of land and sea was set up in the intertidal zone of the back-barrier salt marsh of Spiekeroog Island in the Ge...

• A resampling of 38 small farmland ponds in Belgium after 10 years revealed a high temporal species turnover for both phytoplankton and zooplankton communities, associated with substantial changes in abiotic factors, especially a reduction in total phosphorus concentration. • Across ponds, phytoplankton biomass decreased while evenness and richnes...

Ecological stability is the central framework to understand an ecosystem's ability to absorb or recover from environmental change. Recent modelling and conceptual work suggests that stability is a multidimensional construct comprising different response aspects. Using two freshwater mesocosm experiments as case studies, we show how the response to...

Biodiversity ensures ecosystem functioning and provisioning of ecosystem services, but it remains unclear how biodiversity-ecosystem multifunctionality relationships depend on the identity and number of functions considered. Here, we demonstrate that ecosystem multifunctionality, based on 82 indicator variables of ecosystem functions in a grassland...

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining e...

The research of a generation of ecologists was catalysed by the recognition that the number and identity of species in communities influences the functioning of ecosystems. The relationship between biodiversity and ecosystem functioning (BEF) is most often examined by controlling species richness and randomising community composition. In natural sy...

Recent events and trends in international relations are making it necessary for scientists to design their projects in ways that can integrate disciplinary perspectives and learn how to communicate their results in governance processes. Some examples of settings in which such skills would be needed are the debates about the political and legal rele...

The feeding behavior of the dinoflagellate Oxyrrhis marina with the prasinophyte Pyramimonas grossii as prey was studied in order i) to determine the ingestion rate for this armed microalga, ii) to survey the impact and fate of ejectisomes discharged within predatory cells, and iii) to check if birefringent bodies occur in both, starved and fed Oxy...

The extensive use of traits in ecological studies over the last few decades to predict community functions has revealed that plant traits are plastic and respond to various environmental factors. These plant traits are assumed to predict how plants compete and capture resources. Variation in stoichiometric ratios both within and across species refl...

According to climate models, coastal ecosystems will face an increased frequency of heat waves and increased turbidity due to terrestrial sediment run-off induced by increasing precipitation. Several studies have examined the effects of heat waves and turbidity separately, whereas this study analysed the individual effects of both stressors as well...

There is growing interest in the integration of macroecology and palaeoecology towards a better understanding of past, present, and anticipated future biodiversity dynamics. However, the empirical basis for this integration has thus far been limited. Here we review prospects for a macroecology-palaeoecology integration in biodiversity analyses with...