Species richness, environmental fluctuations, and temporal change in total community biomass

Theory and empirical results suggest that high biodiversity should often cause lower temporal variability in aggregate community properties such as total community biomass. We assembled microbial communities containing 2 to 8 species of competitors in aquatic microcosms and found that the temporal change in total community biomass was positively but insignificantly associated with diversity in a constant temperature environment. There was no evidence of any trend in variable temperature environments. Three non-exclusive mechanisms might explain the lack of a net stabilising effect of species richness on temporal change. (1) A direct destabilising effect of diversity on population level variances caused some populations to vary more when embedded in more diverse communities. (2) Similar responses of the different species to environmental variability might have limited any insurance effect of increased species richness. (3) Large differences in the population level variability of different species (i.e., unevenness) could weaken the relation between species richness and community level stability. These three mechanisms may outweigh the stabilising effects of increases in total community biomass with diversity, statistical averaging, and slightly more negative covariance in more diverse communities. Our experiment and analyses advocate for further experimental investigations of diversity-variability relations.     

How enrichment,ecosystem size,and their effects on species richness co‐determine the stability of microcosm communities

Nutrient enrichment, ecosystem size, and richness each may directly affect the stability of both populations and communities. Alternatively, nutrient enrichment and ecosystem size each may directly affect richness, which in turn may affect stability. No previous studies, however, have tested empirically how these three factors interact and co‐determine stability. We manipulated nutrient input and ecosystem size in replicate microcosms containing a diverse bacterial flora, and a range of green algae and heterotrophic protozoa, and used these manipulations and the resulting variation in species richness to measure their combined effects on temporal stability of both populations and communities. Results showed that nutrient enrichment and ecosystem size controlled protist richness, and their effects on stability could be mediated by richness. In addition, both community‐level and population‐level stability increased with protist richness. Furthermore, mean species evenness and mean species richness was negatively related. Effects of statistical averaging, overyielding, and component population stability were identified as possible mechanisms involved explaini ng the stabilizing effects of richness on community stability. Their relative strength in influencing stability, however, is likely to change as mean evenness decreased with increasing richness. This decrease in evenness would tend to weaken the strength of the statistic averaging effect, but increase the strength of the other two mechanisms due to relatively lower population variability (component population stability) and higher mean biovolumes of dominant protists (overyielding).     

Ecological realism and mechanisms by which diversity begets stability

We investigated how ecological realism might impact the outcome of three experimental manipulations of species richness to determine whether the patterns and the mechanisms underlying richness–variability relationships differ as ecological communities are increasingly exposed to external forces that may drive richness–variability patterns in nature. To test for such an effect, we conducted experiments using rock pool meio‐invertebrate communities housed in three experimental venues: controlled laboratory microcosms, artificially constructed rock pools in the field, and naturally occurring rock pools in the field. Our results showed that experimental venue can have a strong effect on the outcome of richness manipulation experiments. As ecological realism increased, the strength of the relationship between species richness and community variability declined from 32.9% in the laboratory microcosms to 16.8% in the artificial pools to no effect of species richness on community variability in the natural rock pools. The determinants of community variability also differed as ecological realism increased. In laboratory microcosms, community variability was driven solely by mechanisms related to increasing species richness. In artificial rock pools, community variability was driven by a combination of direct and indirect environmental factors as well as mechanisms related to increasing species richness. In the natural rock pools community variability was independent of species richness and was only related to environmental factors. In summary, we found that stabilizing mechanisms associated with species interactions were influential in establishing species richness–variability relations only in the less realistic experimental venues (the laboratory microcosms and the artificial rock pools in the field), and that these mechanisms diminished in importance as ecological realism and complexity of the experimental venue increased. Our results suggest that the effects of diversity might be more difficult to detect in natural systems due to the combined effects of biotic and abiotic forcing, which can mask our ability to detect richness effects.     

Species richness–variability relationships in multi-trophic aquatic microcosms

While species loss may affect the temporal variability of populations and communities differently in multi- versus single-trophic level communities, the nature of these differences are poorly understood. Here, we report on an experiment where we manipulated species richness of multi-trophic rock pool invertebrate communities to determine the effects of species richness, S, on the temporal variability of communities, populations, and individual species. As in single-trophic level studies, temporal variability in community abundance decreased with increasing species richness. However, in contrast to most studies in single-trophic level systems, temporal variability of populations also decreased as species richness increased. Furthermore, the variability of the constituent populations strongly correlated with variability of community abundance suggesting that, in rock pools, S affects community variability through its stabilizing effect on component populations. Our results suggest that species loss may affect population and community variability differently in multi-trophic versus single trophic level communities. If this is so, then the mechanisms proposed to underlie the effects of S on community variability in single-trophic communities may have to be supplemented by those that describe contributions to population stability in order to fully describe the patterns observed in multi-trophic communities.     

Increasing synchrony opposes stabilizing effects of species richness on terrestrial communities

Aim

Ecological theory has predicted that species richness should stabilize communities, with mechanisms including species synchrony and population variability determining the net impacts. While these theories have been supported empirically, results can be sensitive to taxonomic bias as studies are often focussed on plants. Trophic differences between consumers and primary producers can lead to varying stabilizing effects of species richness. Here, we compared the impact of species richness on community variability in four taxonomic groups: terrestrial birds, mammals, invertebrates and plants.

Location

Global.

Method

Using data from 6763 time series globally (BioTIME) for four terrestrial taxa, we quantified community and population variability and species synchrony based on abundance fluctuations over time.

Results

Species richness destabilized communities through increasing synchrony and stabilized communities through reducing population variability in all taxa. Such opposing effects weakened the net impacts of species richness on communities. Population variability had higher importance relative to synchrony in plant communities. By contrast, synchrony had more comparable (or even higher) importance compared with population variability in animal communities. When synchrony and population variability were not controlled, stabilizing impacts of species richness were detected in plant communities only.

Main Conclusions

Our results highlight how species richness drives stabilizing and destabilizing mechanisms simultaneously across all taxa, with strong taxonomic variation in the relative importance of these mechanisms in regulating community variability. This questions the generality of previous findings on stabilizing impacts of species richness based on limited taxonomic coverage. Additionally, our results indicate the need to understand how the importance of stabilizing and destabilizing mechanisms differs in determining community variability across organisms and environments.     

Diversity-dependent stability under mowing and nutrient addition: evidence from a 7-year grassland experiment

Anthropogenic perturbations may affect biodiversity and ecological stability as well as their relationships. However, diversity-stability patterns and associated mechanisms under human disturbances have rarely been explored. We conducted a 7-year field experiment examining the effects of mowing and nutrient addition on the diversity and temporal stability of herbaceous plant communities in a temperate steppe in northern China. Mowing increased population and community stability, whereas nutrient addition had the opposite effects. Stability exhibited positive relationships with species richness at population, functional group and community levels. Treatments did not alter these positive diversity-stability relationships, which were associated with the stabilising effect of species richness on component populations, species asynchrony and portfolio effects. Despite the difficulty of pinpointing causal mechanisms of diversity-stability patterns observed in nature, our results suggest that diversity may still be a useful predictor of the stability of ecosystems confronted with anthropogenic disturbances.     

Biodiversity may regulate the temporal variability of ecological systems

The effect of biodiversity on natural communities has recently emerged as a topic of considerable ecological interest. We review studies that explicitly test whether the number of species in a community (species richness) regulates the temporal variability of aggregate community (total biomass, productivity, nutrient cycling) and population (density, biomass) properties. Theoretical studies predict that community variability should decline with increasing species richness, while population variability should increase. Many, but not all, empirical studies support these expectations. However, a closer look reveals that several empirical studies have either imperfect experimental designs or biased methods of calculating variability. Furthermore, most theoretical studies rely on highly unrealistic assumptions. We conclude that evidence to support the claim that biodiversity regulates temporal variability is accumulating, but not unequivocal. More research, in a broader array of ecosystem types and with careful attention to methodological considerations, is needed before we can make definitive statements regarding richness‐variability relationships.     

The impact of environmental variability and species composition on the stability of experimental microbial populations and communities

Understanding how environmental fluctuations affect the stability of populations and communities is complex, for example, because direct effects of environmental variability on populations may be modified and propagated across communities by species interactions. One way to explore and further understand these complexities is via a factorial manipulation of community composition and environmental conditions. Using laboratory based aquatic microcosms we manipulated environmental fluctuation by creating two environments; one with variable light and one with constant light. Within these environments, community composition was manipulated by constructing communities from all possible combinations of three species that vary in their reliance on light for growth (an autotroph: a diatom completely reliant on light, a heterotroph: a Paramecium species not reliant on light, and a mixotroph: a Paramecium species somewhat reliant on light). Community composition was predicted to affect populations and communities by introducing and altering competitive interactions between species and affecting the degree of niche differentiation between species. We found that population stability was predominantly influenced by an interaction between community composition and environmental variability, whereby the effect of environmental variability synergistically combined with effects of community composition to reduce population stability. Covariance of populations was determined by an interaction between community composition and environmental variability, though this did not result from the effect of niche differentiation between species. Species interactions drove correlations between population biomass and the environment which otherwise did not exist. Our results demonstrate the complex and interrelated effects of abiotic and biotic factors on population and community stability, and suggest the need to consider aspects of community composition when predicting the impact of environmental fluctuations.     

Predicting ecosystem stability from community composition and biodiversity

As biodiversity is declining at an unprecedented rate, an important current scientific challenge is to understand and predict the consequences of biodiversity loss. Here, we develop a theory that predicts the temporal variability of community biomass from the properties of individual component species in monoculture. Our theory shows that biodiversity stabilises ecosystems through three main mechanisms: (1) asynchrony in species’ responses to environmental fluctuations, (2) reduced demographic stochasticity due to overyielding in species mixtures and (3) reduced observation error (including spatial and sampling variability). Parameterised with empirical data from four long‐term grassland biodiversity experiments, our prediction explained 22–75% of the observed variability, and captured much of the effect of species richness. Richness stabilised communities mainly by increasing community biomass and reducing the strength of demographic stochasticity. Our approach calls for a re‐evaluation of the mechanisms explaining the effects of biodiversity on ecosystem stability.     

Biodiversity and ecosystem functioning in aquatic microbial systems: a new analysis of temporal variation and species richness-predictability relations

Studies of microbial communities from aquatic ecosystems provide important insights into relations between various aspects of ecosystem functioning and changes in biodiversity. Aquatic microbial systems provide a valuable counterpoint to studies of terrestrial systems, because patterns reflect consequences of interactions occurring over many generations of community development, and are unlikely to represent artifacts of the initial conditions established in experimental communities. In this paper we re-analyse our previously published data to separate the contributions of temporal and spatial variation to overall variation in ecosystem functioning. A new analysis based on re-sampling confirms a negative relationship between richness and the variability of one ecosystem process, carbon dioxide flux. The negative relationship reflects high variation among communities of low species richness, rather than high temporal variation within communities of low richness. We also review the various transformations and summary statistics proposed as alternate measures of variability in ecosystem functioning, to point out that different measures are often appropriate for different kinds of data. Finally, we conclude that arguments about the cosmopolitan distribution of microbes do not preclude the existence of important relations between microbial species richness and ecosystem functioning.     

Drought resistance increases with species richness in restored populations and communities

It is unknown to what extent or by what mechanisms introducing biodiversity influences stability of high-stress ecosystems undergoing restoration. Opportunity to investigate patterns of biodiversity and resistance to disturbance in a high-stress environment was presented when severe drought struck a restoration experiment underway on abandoned limestone quarry floors in Ontario, Canada. Experimental communities were previously established within small quarry-floor plots by sowing native grass and forb species considered to be characteristic of rare natural limestone pavements called alvars. Despite adding an identical 18-species seed-mixture to all plots, realized communities varied extensively with respect to the numbers of species established (species richness), the total number of individuals established (community abundance), and the number of individuals belonging to each species (population abundances). We investigated the relationship between species richness and resistance of community abundance to drought, while accounting for background richness–abundance correlation, by contrasting slopes and intercepts of the richness–abundance relationship immediately before vs. 6 weeks after the drought. This relationship was significantly positive prior to drought but 72% steeper in slope following drought, while the abundance intercept exhibited a 44% drop. Plots featuring richer, more abundant communities prior to drought thus suffered considerably less damage than species-poor, low-abundance plots. Population abundance was weakly related to richness prior to drought, but strongly and positively related to richness after the drought. At the individual species level, no species experienced greater losses of abundance with increased plot richness, but six species experienced reduced abundance losses where they co-occurred with more neighbour species. Facilitation or other mechanisms capable of increasing population resistance may thus underlie community resistance in high-stress environments. Though controlled experiments are required to establish causes of relationships reported here, the forms of these relationships suggest that managers may be able to promote resistance in high-stress ecosystems by establishing species-rich communities.     

Herbivores control effects of algal species richness on community biomass and stability in a laboratory microcosm experiment

Hundreds of studies that have explored how biodiversity affects the productivity and stability of ecosystems have produced a consensus that communities composed of more species tend to have higher biomass that is more stable through time. However, the majority of this work stems from studies performed using highly simplified food webs, often composed of just primary producers competing for inorganic resources in the absence of trophic interactions. When studies have incorporated trophic interactions, diversity‐function relationships have been more variable, leaving open the question of how biodiversity affects the functioning of ecosystems with more trophic levels. Here we report the results of a laboratory experiment that used freshwater microcosms to test for effects of algal diversity (one or four species) on community biomass and temporal variability in the presence and absence of two different herbivore species (cladocerans Ceriodaphnia dubia and Daphnia pulex). When no herbivores were present, we found the classic pattern observed in hundreds of other studies – as species richness of algae increased, algal biomass increased, and the temporal variation in biomass decreased. This pattern was retained when one of the herbivores (C. dubia) was present. Ceriodaphnia dubia exhibited weak and non‐selective grazing on the focal algae, leaving the effect of diversity on biomass and variability essentially intact. In contrast, D. pulex exhibited strong and selective grazing in algal polycultures that qualitatively altered both diversity–function relationships. As algal richness increased, total algal biomass decreased and variation through time increased. These changes were coupled with larger and less variable populations of D. pulex. Our results show that herbivory leads to a richer array of diversity–function relationships than often observed in studies focused on just one trophic level, and suggests trophic interactions should be given more attention in work that seeks to determine how biodiversity impacts the functioning of ecosystems.     

The Impact of Anthropogenic Disturbance on Assembly Patterns of Termite Communities

Intensification of land‐use threatens biodiversity, especially in tropical ecosystems that harbor the planet's highest species richness. This negative impact of anthropogenic disturbance on species numbers is well established, but the mechanisms underlying the community assembly processes are less well understood. Termites are of fundamental importance in tropical ecosystems where they are critical for nutrient recycling and species diversity. We tested the impact of anthropogenic disturbance on termite species diversity and assembly processes in a West African savanna applying the newest techniques of phylogenetic community analyses. Species richness dropped in areas of intensive land‐use and compositional similarity between intensive land‐use areas was high. This contrasted with a protected National Park where communities were characterized by high species richness and intermediate species turnover between sites. Slightly disturbed areas in the buffer zone surrounding the park were intermediate, they still had high species richness but similarity between sites increased. Strikingly, the assembly pattern also changed with disturbance from more phylogenetic overdispersion to more clustering (coexisting species became phylogenetically more similar), but only when the fungus‐growing termite Macrotermes bellicosus was absent. Our data suggest that the major forces structuring termite communities depend: (1) on the presence of this dominant mound‐building termite; and (2) that they change to more environmental filtering with disturbance. Anthropogenic disturbance seems to function as a filter that allows only a specific subset of species to occur. Such an effect might be widespread in ecology but it is difficult to document quantitatively. Phylogenetic community analyses can help to contribute such evidence.     

The relationship between species richness and evenness: a meta-analysis of studies across aquatic ecosystems

Biological diversity comprises both species richness, i.e., the number of species in a community, and evenness, measuring how similar species are in their abundances. The relationship between species richness and evenness (RRE) across communities remains, however, a controversial issue in ecology because no consistent pattern has been reported. We conducted a systematic meta-review of RRE in aquatic ecosystems along regional to continental gradients and across trophic groups, differing in body size by 13 orders of magnitude. Hypotheses that RRE responded to latitudinal and scale variability across trophic groups were tested by regression analyses. Significant correlations of species richness and evenness only existed in 71 out of 229 datasets. Among the RRE, 89 were negative and 140 were positive. RRE did not vary with latitude but showed a positive response to scale. In a meta-analysis with ecosystem type as a single explaining variable, RRE did not vary among ecosystem types, i.e. between marine and freshwater. Finally, autotrophs had more positive RRE than heterotrophs. The weak RRE in many aquatic datasets suggests that richness and evenness often reflect independent components of biodiversity, highlighting that richness alone may be an incomplete surrogate for biodiversity. Our results further elucidate that RRE is driven by organismal and environmental properties, both of which must be considered to gain a deeper understanding of large-scale patterns of biodiversity.     

Experimental design and the outcome and interpretation of diversity–stability relations

The nature of the relationship between diversity and stability has become the subject of intense research effort over the last few decades as the role of diversity as a major driver of ecosystem functioning and stability has come to the forefront of ecological interest. Here, we present a meta‐analysis of the impact of twelve experimental design factors on the strength and direction of relations between biotic richness and temporal variability at both the aggregate community‐ and population‐level. Based on 35 studies that report 59 community‐level and 36 population‐level relations, our results show that biotic richness has a highly general stabilizing effect on community properties that are only marginally affected by the nuances of experimental design. In contrast, experimental design factors have a highly significant effect on mean effect sizes and the resulting interpretation of relations between richness and population‐level variability. The strongest dichotomous effect was observed based on the method of calculating the response variable, such that when population variability was calculated as the mean variability of populations across all replicates, biotic richness showed a negative (stabilizing) mean effect size. In contrast, when population variability was calculated on a per replicate basis, biotic richness showed a positive (destabilizing) mean effect size. This latter result suggests that a renewed focus on the mechanisms by which populations can be stabilized (and destabilized) by diversity is needed.     

Species richness facilitates ecosystem resilience in aquatic food webs

1. Many studies indicate that biodiversity in ecosystems affects stability, either by promoting temporal stability of ecosystem attributes or by enhancing ecosystem resistance and resilience to perturbation. The effects on temporal stability are reasonably well understood and documented but effects on resistance and resilience are not. 2. Here, we report results from an aquatic mesocosm experiment in which we manipulated the species richness and composition of aquatic food webs (macrophytes, macro‐herbivores and invertebrate predators), imposed a pulse disturbance (acidification), and monitored the resistance (initial response) and resilience (recovery) of ecosystem productivity and respiration. 3. We found that species‐rich macroinvertebrate communities had higher resilience of whole‐ecosystem respiration, but were not more resistant to perturbations. We also found that resilience and resistance were unaffected by species composition, despite the strong role composition is known to play in determining mean levels of function in these communities. 4. Biodiversity’s effects on resilience were probably mediated through complex pathways affecting phytoplankton and microbial communities (e.g. via changes in nutrient regeneration, grazing or compositional changes) rather than through simpler effects (e.g. insurance effects, enhanced facilitation) although these simpler mechanisms probably played minor roles in enhancing respiration resilience. 5. Current mechanisms for understanding biodiversity’s effects on ecosystem stability have been developed primarily in the context of single‐trophic level communities. These mechanisms may be overly simplistic for understanding the consequences of species richness on ecosystem stability in complex, multi‐trophic food webs where additional factors such as indirect effects and highly variable life‐history traits of species may also be important.     

Restoration of plant–pollinator interaction networks via species translocation

The recent decline in pollinator biodiversity, notably in the case of wild bee populations, puts both wild and agricultural ecosystems at risk of ecological community collapse. This has triggered calls for further study of these mutualistic communities in order to more effectively inform restoration of disturbed plant–pollinator communities. Here, we use a dynamic network model to test a variety of translocation strategies for restoring a community after it experiences the loss of some of its species. We consider the reintroduction of extirpated species, both immediately after the original loss and after the community has reequilibrated, as well as the introduction of other native species that were originally absent from the community. We find that reintroducing multiple highly interacting generalist species best restores species richness for lightly disturbed communities. However, for communities that experience significant losses in biodiversity, introducing generalist species that are not originally present in the community may most effectively restore species richness, although in these cases the resultant community often shares few species with the original community. We also demonstrate that the translocation of a single species has a minimal impact on both species richness and the frequency of community collapse. These results have important implications for restoration practices in the face of varying degrees of community perturbations, the refinement of which is crucial for community management.     

Productivity, herbivory, and species traits rather than diversity influence invasibility of experimental phytoplankton communities

Biological invasions are a major threat to natural biodiversity; hence, understanding the mechanisms underlying invasibility (i.e., the susceptibility of a community to invasions by new species) is crucial. Invasibility of a resident community may be affected by a complex but hitherto hardly understood interplay of (1) productivity of the habitat, (2) diversity, (3) herbivory, and (4) the characteristics of both invasive and resident species. Using experimental phytoplankton microcosms, we investigated the effect of nutrient supply and species diversity on the invasibility of resident communities for two functionally different invaders in the presence or absence of an herbivore. With increasing nutrient supply, increased herbivore abundance indicated enhanced phytoplankton biomass production, and the invasion success of both invaders showed a unimodal pattern. At low nutrient supply (i.e., low influence of herbivory), the invasibility depended mainly on the competitive abilities of the invaders, whereas at high nutrient supply, the susceptibility to herbivory dominated. This resulted in different optimum nutrient levels for invasion success of the two species due to their individual functional traits. To test the effect of diversity on invasibility, a species richness gradient was generated by random selection from a resident species pool at an intermediate nutrient level. Invasibility was not affected by species richness; instead, it was driven by the functional traits of the resident and/or invasive species mediated by herbivore density. Overall, herbivory was the driving factor for invasibility of phytoplankton communities, which implies that other factors affecting the intensity of herbivory (e.g., productivity or edibility of primary producers) indirectly influence invasions.     

Conceptualising the interactive effects of climate change and biological invasions on subarctic freshwater fish

Climate change and species invasions represent key threats to global biodiversity. Subarctic freshwaters are sentinels for understanding both stressors because the effects of climate change are disproportionately strong at high latitudes and invasion of temperate species is prevalent. Here, we summarize the environmental effects of climate change and illustrate the ecological responses of freshwater fishes to these effects, spanning individual, population, community and ecosystem levels. Climate change is modifying hydrological cycles across atmospheric, terrestrial and aquatic components of subarctic ecosystems, causing increases in ambient water temperature and nutrient availability. These changes affect the individual behavior, habitat use, growth and metabolism, alter population spawning and recruitment dynamics, leading to changes in species abundance and distribution, modify food web structure, trophic interactions and energy flow within communities and change the sources, quantity and quality of energy and nutrients in ecosystems. Increases in temperature and its variability in aquatic environments underpin many ecological responses; however, altered hydrological regimes, increasing nutrient inputs and shortened ice cover are also important drivers of climate change effects and likely contribute to context‐dependent responses. Species invasions are a complex aspect of the ecology of climate change because the phenomena of invasion are both an effect and a driver of the ecological consequences of climate change. Using subarctic freshwaters as an example, we illustrate how climate change can alter three distinct aspects of species invasions: (1) the vulnerability of ecosystems to be invaded, (2) the potential for species to spread and invade new habitats, and (3) the subsequent ecological effects of invaders. We identify three fundamental knowledge gaps focused on the need to determine (1) how environmental and landscape characteristics influence the ecological impact of climate change, (2) the separate and combined effects of climate and non‐native invading species and (3) the underlying ecological processes or mechanisms responsible for changes in patterns of biodiversity.     

Species richness enhances both algal biomass and rates of oxygen production in aquatic microcosms

Accelerating rates of species extinction have generated much recent interest in understanding how biodiversity affects the functioning of ecosystems. Experiments to date have shown communities composed of fewer species generally capture a smaller fraction of available resources, and achieve lower standing stock biomass than more diverse communities. However, it is uncertain how changes in biodiversity and the resulting alterations in biomass affect the rates of important ecological processes like primary production, which regulates fluxes of CO₂ and O₂ between the biotic and abiotic components of the environment. Here we show that species richness influences not only the standing stock biomass of primary producers, but also rates of gross primary production measured by changes in O₂ concentrations in aquatic systems. We manipulated the richness of five widespread species of algae in laboratory microcosms and then quantified how richness impacts algal biomass, rates of gross primary production (GPP), and the ratio of production to respiration. Algal biomass increased by a factor of 1.82 for each level of species richness, and GPP by a factor of 1.20, for each additional species. Production to respiration ratios increased about 10% for each additional species, indicating that systems with more species were increasingly autotrophic – that is, they produced more O₂ than they consumed, and accumulated CO₂ faster than they released it. These trends were driven by two highly productive species that became co-dominant in species rich polycultures at the expense of other taxa. Our experiment suggests that changes in biodiversity may influence not only the rates at which O₂ and CO₂ are produced and released in ecosystems, but also the total amount of carbon that is sequestered and stored as biomass.