The role of economics in ecosystem based management: The case of the EU Marine Strategy Framework Directive, First lessons learnt and way forward

The EU Marine Strategy Framework Directive (MSFD) sets out a plan of action relating to marine environmental policy and in particular to achieving ‘good environmental status’ (GES) in European marine waters by 2020. Article 8.1 (c) of the Directive calls for ‘an economic and social analysis of the use of those waters and of the cost of degradation of the marine environment’. The MSFD is ‘informed’ by the Ecosystem Approach to management, with GES interpreted in terms of ecosystem functioning and services provision. Implementation of the Ecosystem Approach is expected to be by adaptive management policy and practice. The initial socio-economic assessment was made by maritime EU Member States between 2011 and 2012, with future updates to be made on a regular basis. For the majority of Member States, this assessment has led to an exercise combining an analysis of maritime activities both at national and coastal zone scales, and an analysis of the non-market value of marine waters. In this paper we examine the approaches taken in more detail, outline the main challenges facing the Member States in assessing the economic value of achieving GES as outlined in the Directive and make recommendations for the theoretically sound and practically useful completion of the required follow-up economic assessments specified in the MSFD.

Using life strategies to explore the vulnerability of ecosystem services to invasion by alien plants

Invasive plants can have different effects of ecosystem functioning and on the provision of ecosystem services, from strongly deleterious impacts to positive effects. The nature and intensity of such effects will depend on the service and ecosystem being considered, but also on features of life strategies of invaders that influence their invasiveness as well as their influence of key processes of receiving ecosystems. To address the combined effect of these various factors we developed a robust and efficient methodological framework that allows to identify areas of possible conflict between ecosystem services and alien invasive plants, considering interactions between landscape invasibility and species invasiveness. Our framework combines the statistical robustness of multi-model inference, efficient techniques to map ecosystem services, and life strategies as a functional link between invasion, functional changes and potential provision of services by invaded ecosystems. The framework was applied to a test region in Portugal, for which we could successfully predict the current patterns of plant invasion, of ecosystem service provision, and finally of probable conflict (expressing concern for negative impacts, and value for positive impacts on services) between alien species richness (total and per plant life strategy) and the potential provision of selected services. Potential conflicts were identified for all combinations of plant strategy and ecosystem service, with an emphasis for those concerning conflicts with carbon sequestration, water regulation and wood production. Lower levels of conflict were obtained between invasive plant strategies and the habitat for biodiversity supporting service. The added value of the proposed framework in the context of landscape management and planning is discussed in perspective of anticipation of conflicts, mitigation of negative impacts, and potentiation of positive effects of plant invasions on ecosystems and their services.

Soil management in relation to sustainable agriculture and ecosystem services

Requirements for research, practices and policies affecting soil management in relation to global food security are reviewed. Managing soil organic carbon (C) is central because soil organic matter influences numerous soil properties relevant to ecosystem functioning and crop growth. Even small changes in total C content can have disproportionately large impacts on key soil physical properties. Practices to encourage maintenance of soil C are important for ensuring sustainability of all soil functions. Soil is a major store of C within the biosphere – increases or decreases in this large stock can either mitigate or worsen climate change. Deforestation, conversion of grasslands to arable cropping and drainage of wetlands all cause emission of C; policies and international action to minimise these changes are urgently required. Sequestration of C in soil can contribute to climate change mitigation but the real impact of different options is often misunderstood. Some changes in management that are beneficial for soil C, increase emissions of nitrous oxide (a powerful greenhouse gas) thus cancelling the benefit. Research on soil physical processes and their interactions with roots can lead to improved and novel practices to improve crop access to water and nutrients. Increased understanding of root function has implications for selection and breeding of crops to maximise capture of water and nutrients. Roots are also a means of delivering natural plant-produced chemicals into soil with potentially beneficial impacts. These include biocontrol of soil-borne pests and diseases and inhibition of the nitrification process in soil (conversion of ammonium to nitrate) with possible benefits for improved nitrogen use efficiency and decreased nitrous oxide emission. The application of molecular methods to studies of soil organisms, and their interactions with roots, is providing new understanding of soil ecology and the basis for novel practical applications. Policy makers and those concerned with development of management approaches need to keep a watching brief on emerging possibilities from this fast-moving area of science. Nutrient management is a key challenge for global food production: there is an urgent need to increase nutrient availability to crops grown by smallholder farmers in developing countries. Many changes in practices including inter-cropping, inclusion of nitrogen-fixing crops, agroforestry and improved recycling have been clearly demonstrated to be beneficial: facilitating policies and practical strategies are needed to make these widely available, taking account of local economic and social conditions. In the longer term fertilizers will be essential for food security: policies and actions are needed to make these available and affordable to small farmers. In developed regions, and those developing rapidly such as China, strategies and policies to manage more precisely the necessarily large flows of nutrients in ways that minimise environmental damage are essential. A specific issue is to minimise emissions of nitrous oxide whilst ensuring sufficient nitrogen is available for adequate food production. Application of known strategies (through either regulation or education), technological developments, and continued research to improve understanding of basic processes will all play a part. Decreasing soil erosion is essential, both to maintain the soil resource and to minimise downstream damage such as sedimentation of rivers with adverse impacts on fisheries. Practical strategies are well known but often have financial implications for farmers. Examples of systems for paying one group of land users for ecosystem services affecting others exist in several parts of the world and serve as a model.

Food-web composition affects cross-ecosystem interactions and subsidies

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Effects of two traits of the ecological state equation on our understanding of species coexistence and ecosystem services

Species coexistence has been a fundamental issue to understand ecosystem functioning since the beginnings of ecology as a science. The search of a reliable and all-encompassing explanation for this issue has become a complex goal with several apparently opposing trends. On the other side, seemingly unconnected with species coexistence, an ecological state equation based on the inverse correlation between an indicator of dispersal that fits gamma distribution and species diversity has been recently developed. This article explores two factors, whose effects are inconspicuous in such an equation at the first sight, that are used to develop an alternative general theoretical background in order to provide a better understanding of species coexistence. Our main outcomes are: (i) the fit of dispersal and diversity values to gamma distribution is an important factor that promotes species coexistence mainly due to the right-skewed character of gamma distribution; (ii) the opposite correlation between species diversity and dispersal implies that any increase of diversity is equivalent to a route of “ecological cooling” whose maximum limit should be constrained by the influence of the third law of thermodynamics; this is in agreement with the well-known asymptotic trend of diversity values in space and time; (iii) there are plausible empirical and theoretical ways to apply physical principles to explain important ecological processes; (iv) the gap between theoretical and empirical ecology in those cases where species diversity is paradoxically high could be narrowed by a wave model of species coexistence based on the concurrency of local equilibrium states. In such a model, competitive exclusion has a limited but indispensable role in harmonious coexistence with functional redundancy. We analyze several literature references as well as ecological and evolutionary examples that support our approach, reinforcing the meaning equivalence between important physical and ecological principles.

Ecosystem functions are resistant to extreme changes to rainfall regimes in a mesotrophic grassland

Major changes to rainfall regimes are predicted for the future but the effect of such changes on terrestrial ecosystem function is largely unknown. We created a rainfall manipulation experiment to investigate the effects of extreme changes in rainfall regimes on ecosystem functioning in a grassland system. We applied two rainfall regimes; a prolonged drought treatment (30 % reduction over spring and summer) and drought/downpour treatment (long periods of no rainfall interspersed with downpours), with an ambient control. Both rainfall manipulations included increased winter rainfall. We measured plant community composition, CO2 fluxes and soil nutrient availability. Plant species richness and cover were lower in the drought/downpour treatment, and showed little recovery after the treatment ceased. Ecosystem processes were less affected, possibly due to winter rainfall additions buffering reduced summer rainfall, which saw relatively small soil moisture changes. However, soil extractable P and ecosystem respiration were significantly higher in rainfall change treatments than in the control. This grassland appears fairly resistant, in the short term, to even the more extreme rainfall changes that are predicted for the region, although prolonged study is needed to measure longer-term impacts. Differences in ecosystem responses between the two treatments emphasise the variety of ecosystem responses to changes in both the size and frequency of rainfall events. Given that model predictions are inconsistent there is therefore a need to assess ecosystem function under a range of potential climate change scenarios.

Plant diversity and ecosystem multifunctionality peak at intermediate levels of woody cover in global drylands

Aim The global spread of woody plants into grasslands is predicted to increase over the coming century. While there is general agreement regarding the anthropogenic causes of this phenomenon, its ecological consequences are less certain. We analysed how woody vegetation of differing cover affects plant diversity (richness and evenness) and the surrogates of multiple ecosystem processes (multifunctionality) in global drylands, and how these change with aridity. Location Two hundred and twenty-four dryland sites from all continents except Antarctica, widely differing in their environmental conditions (from arid to dry-subhumid sites) and relative woody cover (from 0 to 100). Methods Using a standardized field survey, we measured the cover, richness and evenness of perennial vegetation. At each site, we measured 14 soil variables related to fertility and the build-up of nutrient pools. These variables are critical for maintaining ecosystem functioning in drylands. Results Species richness and ecosystem multifunctionality were strongly related to woody vegetation, with both variables peaking at a relative woody cover (RWC) of 41–60. This relationship shifted with aridity. We observed linear positive effects of RWC in dry-subhumid sites. These positive trends shifted to hump-shaped RWC–diversity and multifunctionality relationships under semi-arid environments. Finally, hump-shaped (richness, evenness) or linear negative (multifunctionality) effects of RWC were found under the most arid conditions. Main conclusions Plant diversity and multifunctionality peaked at intermediate levels of woody cover, although this relationship became increasingly positive in wetter environments. This comprehensive study accounts for multiple ecosystem attributes across a range of levels of woody cover and environmental conditions. Our results help us to reconcile contrasting views of woody encroachment found in the current literature and can be used to improve predictions of the likely effects of encroachment on biodiversity and ecosystem services.

Aluminum toxicity in a tropical montane forest ecosystem in southern Ecuador

Aluminum phytotoxicity frequently occurs in acid soils (pH < 5.5) and was therefore discussed to affect ecosystem functioning of tropical montane forests. The susceptibility to Al toxicity depends on the sensitivity of the plant species and the Al speciation in soil solution, which can vary highly depending e.g., on pH, ionic strength, and dissolved organic matter. An acidification of the ecosystem and periodic base metal deposition from Saharan dust may control plant available Al concentrations in the soil solutions of tropical montane rainforests in south Ecuador. The overall objective of my study was to assess a potential Al phytotoxicity in the tropical montane forests in south Ecuador. For this purpose, I exposed three native Al non-accumulating tree species (Cedrela odorata L., Heliocarpus americanus L., and Tabebuia chrysantha (Jacq.) G. Nicholson) to increased Al concentrations (0 – 2400 μM Al) in a hydroponic experiment, I established dose-response curves to estimate the sensitivity of the tree species to increased Al concentrations, and I investigated the mechanisms behind the observed effects induced by elevated Al concentrations. Furthermore, the response of Al concentrations and the speciation in soil solution to Ca amendment in the study area were determined. In a final step, I assessed all major Al fluxes, drivers of Al concentrations in ecosystem solutions, and indicators of Al toxicity in the tropical montane rainforest in Ecuador in order to test for indications of Al toxicity. In the hydroponic experiment, a 10 % reduction in aboveground biomass production occurred at 126 to 376 μM Al (EC10 values), probably attributable to decreased Mg concentrations in leaves and reduced potosynthesis. At 300 μM Al, increased root biomass production of T. chrysantha was observed. Phosphorus concentrations in roots of C. odorata and T. chrysantha were significantly highest in the treatment with 300 μM Al and correlated significantly with root biomass, being a likely reason for stimulated root biomass production. The degree of organic complexation of Al in the organic layer leachate, which is central to plant nutrition because of the high root density, and soil solution from the study area was very high (mean > 99 %). The resulting low free Al concentrations are not likely to affect plant growth, although the concentrations of potentially toxic Al3+ increased with soil depth due to higher total Al and lower dissolved organic matter concentrations in soil solutions. The Ca additions caused an increase of Al in the organic layer leachate, probably because Al3+ was exchanged against the added Ca2+ ions while pH remained constant. The free ion molar ratios of Ca2+:Al3+ (mean ratio ca. 400) were far above the threshold (≤ 1) for Al toxicity, because of a much higher degree of organo-complexation of Al than Ca. High Al fluxes in litterfall (8.8 – 14.2 kg ha−1 yr−1) indicate a high Al circulation through the ecosystem. The Al concentrations in the organic layer leachate were driven by the acidification of the ecosystem and increased significantly between 1999 and 2008. However, the Ca:Al molar ratios in organic layer leachate and all aboveground ecosystem solutions were above the threshold for Al toxicity. Except for two Al accumulating and one non-accumulating tree species, the Ca:Al molar ratios in tree leaves from the study area were above the Al toxicity threshold of 12.5. I conclude that toxic effects in the hydroponic experiment occurred at Al concentrations far above those in native organic layer leachate, shoot biomass production was likely inhibited by reduced Mg uptake, impairing photosynthesis, and the stimulation of root growth at low Al concentrations can be possibly attributed to improved P uptake. Dissolved organic matter in soil solutions detoxifies Al in acidic tropical forest soils and a wide distribution of Al accumulating tree species and high Al fluxes in the ecosystem do not necessarily imply a general Al phytotoxicity.

Are shrubs really a sign of declining ecosystem function? Disentangling the myths and truths of woody encroachment in Australia

Since European settlement, there has been a dramatic increase in the density, cover and distribution of woody plants in former grassland and open woodland. There is a widespread belief that shrub encroachment is synonymous with declines in ecosystem functions, and often it is associated with landscape degradation or desertification. Indeed, this decline in ecosystem functioning is considered to be driven largely by the presence of the shrubs themselves. This prevailing paradigm has been the basis for an extensive program of shrub removal, based on the view that it is necessary to reinstate the original open woodland or grassland structure from which shrublands are thought to have been derived. We review existing scientific evidence, particularly focussed on eastern Australia, to question the notion that shrub encroachment leads to declines in ecosystem functions. We then summarise this scientific evidence into two conceptual models aimed at optimising landscape management to maximise the services provided by shrub-encroached areas. The first model seeks to reconcile the apparent conflicts between the patch- and landscape-level effects of shrubs. The second model identifies the ecosystem services derived from different stages of shrub encroachment. We also examined six ecosystem services provided by shrublands (biodiversity, soil C, hydrology, nutrient provision, grass growth and soil fertility) by using published and unpublished data. We demonstrated the following: (1) shrub effects on ecosystems are strongly scale-, species- and environment-dependent and, therefore, no standardised management should be applied to every case; (2) overgrazing dampens the generally positive effect of shrubs, leading to the misleading relationship between encroachment and degradation; (3) woody encroachment per se does not hinder any of the functions or services described above, rather it enhances many of them; (4) no single shrub-encroachment state (including grasslands without shrubs) will maximise all services; rather, the provision of ecosystem goods and services by shrublands requires a mixture of different states; and (5) there has been little rigorous assessment of the long-term effectiveness of removal and no evidence that this improves land condition in most cases. Our review provides the basis for an improved, scientifically based understanding and management of shrublands, so as to balance the competing goals of providing functional habitats, maintaining soil processes and sustaining pastoral livelihoods.

Direct and indirect effects of invasion by the alien tree Ailanthus altissima on riparian plant communities and ecosystem multifunctionality

Most existing studies addressing the effects of invasive species on biodiversity focus on species richness ignoring better indicators of biodiversity and better predictors of ecosystem functioning such as the diversity of evolutionary histories (phylodiversity). Moreover, no previous study has separated the direct effect of alien plants on multiple ecosystem functions simultaneously (multifunctionality) from those indirect ones mediated by the decrease on biodiversity caused by alien plants. We aimed to analyze direct and indirect effects, mediated or not by its effect on biodiversity, of the invasive tree Ailanthus altissima on ecosystem multifunctionality of riparian habitats under Mediterranean climate. We measured vegetation attributes (species richness and phylodiversity) and several surrogates of ecosystem functioning (understory plant biomass, soil enzyme activities, available phosphorous and organic matter) in plots infested by A. altissima and in control (non-invaded) ones. We used structural equation modelling to tease apart the direct and indirect effects of A. altissima on ecosystem multifunctionality. Our results suggest that lower plant species richness, phylodiversity and multifunctionality were associated to the presence of A. altissima. When analyzing each function separately, we found that biodiversity has the opposite effect of the alien plant on all the different functions measured, therefore reducing the strength of the effect (either positive or negative) of A. altissima on them. This is one of the few existing studies addressing the effect of invasive species on phylodiversity and also studying the effect of invasive species on multiple ecosystem functions simultaneously.

Shifts in microbial communities do not explain the response of grassland ecosystem function to plant functional composition and rainfall change

Ecosystem functioning in grasslands is regulated by a range of biotic and abiotic factors, and the role of microbial communities in regulating ecosystem function has been the subject of much recent scrutiny. However, there are still knowledge gaps regarding the impacts of rainfall and vegetation change upon microbial communities and the implications of these changes for ecosystem functioning. We investigated this issue using data from an experimental mesotrophic grassland study in south-east England, which had been subjected to four years of rainfall and plant functional composition manipulations. Soil respiration, nitrogen and phosphorus stocks were measured, and the abundance and community structure of soil microbes were characterised using quantitative PCR and multiplex-TRFLP analysis, respectively. Bacterial community structure was strongly related to the plant functional composition treatments, but not the rainfall treatment. However, there was a strong effect of both rainfall change and plant functional group upon bacterial abundance. There was also a weak interactive effect of the two treatments upon fungal community structure, although fungal abundance was not affected by either treatment. Next, we used a statistical approach to assess whether treatment effects on ecosystem function were regulated by the microbial community. Our results revealed that ecosystem function was influenced by the experimental treatments, but was not related to associated changes to the microbial community. Overall, these results indicate that changes in fungal and bacterial community structure and abundance play a relatively minor role in determining grassland ecosystem function responses to precipitation and plant functional composition change, and that direct effects on soil physical and chemical properties and upon plant and microbial physiology may play a more important role.

On the importance of shrub encroachment by sprouters, climate, species richness and anthropic factors for ecosystem multifunctionality in semi-arid Mediterranean ecosystems

One of the most important changes taking place in drylands worldwide is the increase of the cover and dominance of shrubs in areas formerly devoid of them (shrub encroachment). A large body of research has evaluated the causes and consequences of shrub encroachment for both ecosystem structure and functioning. However, there are virtually no studies evaluating how shrub encroachment affects the ability of ecosystems to maintain multiple functions and services simultaneously (multifunctionality). We aimed to do so by gathering data from ten ecosystem functions linked to the maintenance of primary production and nutrient cycling and storage (organic C, activity of β-glucosidase, pentoses, hexoses, total N, total available N, amino acids, proteins, available inorganic P, and phosphatase activity), and summarizing them in a multifunctionality index (M). We assessed how climate, species richness, anthropic factors (distance to the nearest town, sandy and asphalted road, and human population in the nearest town at several historical periods) and encroachment by sprouting shrubs impacted both the functions in isolation and M along a regional (ca. 350 km) gradient in Mediterranean grasslands and shrublands dominated by a non-sprouting shrub. Values of M were higher in those grasslands and shrublands containing sprouting shrubs (43 and 62%, respectively). A similar response was found when analyzing the different functions in isolation, as encroachment by sprouting shrubs increased functions by 2–80% compared to unencroached areas. Encroachment was the main driver of changes in M along the regional gradient evaluated, followed by anthropic factors and species richness. Climate had little effects on M in comparison to the other factors studied. Similar responses were observed when evaluating the functions in isolation. Overall, our results showed that M was higher at sites with higher sprouting shrub cover, longer distance to roads and higher perennial plant species richness. Our study is the first documenting that ecosystem multifunctionality in shrublands is enhanced by encroaching shrubs differing in size and leaf attributes. Our findings reinforce the idea that encroachment effects on ecosystem functioning cannot be generalized, and that are largely dependent on the traits of the encroaching shrub relative to those of the species being replaced.

Plant diversity influencing structure and functioning of Caatinga vegetation

O estudo dos efeitos que a diversidade de espécies pode causar nos processos ecossistêmicos tem crescido vertiginosamente nas últimas duas décadas. Diversos trabalhos experimentais realizados no mundo todo têm demonstrado que uma maior diversidade de plantas contribui para o aumento da produtividade de ecossistemas terrestres. Além disso, esse efeito pode influenciar processos em diversos níveis tróficos, contribuindo assim para a estabilidade dos processos ecossistêmicos a longo prazo. Paralelamente com os estudos do efeito da diversidade, muita atenção tem sido dada para desvendar o papel das características funcionais das espécies no funcionamento dos ecossistemas. Isto porque as características funcionais das espécies têm se mostrado importantes "peças" no entendimento dos efeitos que espécies individuais podem exercer nos ecossistemas e suas respostas ao ambiente. Nesta tese de doutorado eu explorei algumas lacunas de conhecimento dentro dessa área em crescente desenvolvimento conhecida na literatura ecológica como "biodiversidade e funcionamento dos ecossistemas". No primeiro capítulo, eu busquei evidências para mecanismos que podem explicar a relação positiva entre diversidade e funcionamento com foco em cinco mecanismos relacionados às interações entre plantas, tendo como parâmetro de funcionamento a produtividade primária. No segundo capítulo, eu utilizei técnicas para a estimativa de padrões de diversidade em escalas biogeográficas e bases de dados de satélites com longa duração para desvendar se a biodiversidade em escalas macroecológicas promove a estabilidade da produtividade dos ambientes terrestres no semiárido brasileiro. Por fim, o objetivo do terceiro capítulo foi entender como a perda da cobertura vegetal originária do uso da terra por comunidades tradicionais no semiárido brasileiro influenciaria os processos de interações entre plantas e o papel das características funcionais das espécies nessas interações. Acredito que a contribuição individual de cada capítulo preenche lacunas de conhecimento importantes dessa área da Ecologia que ainda se encontra em expansão.

The influence of balanced and imbalanced resource supply on biodiversity – functioning relationship across ecosystems

Numerous studies show that increasing species richness leads to higher ecosystem productivity. This effect is often attributed to more efficient portioning of multiple resources in communities with higher numbers of competing species, indicating the role of resource supply and stoichiometry for biodiversity-ecosystem functioning relationships. Here, we merged theory on ecological stoichiometry with a framework of biodiversity-ecosystem functioning to understand how resource use transfers into primary production. We applied a structural equation model to define patterns of diversity-productivity relationships with respect to available resources. Meta-analysis was used to summarize the findings across ecosystem types ranging from aquatic ecosystems to grasslands and forests. As hypothesized, resource supply increased realized productivity and richness, but we found significant differences between ecosystems and study types. Increased richness was associated with increased productivity, although this effect was not seen in experiments. More even communities had lower productivity, indicating that biomass production is often maintained by a few dominant species, and reduced dominance generally reduced ecosystem productivity. This synthesis, which integrates observational and experimental studies in a variety of ecosystems and geographical regions, exposes common patterns and differences in biodiversity-functioning relationships, and increases the mechanistic understanding of changes in ecosystems productivity.