Rationalising Biodiversity Conservation in Dynamic Ecosystems (RUBICODE) Research Overview For further information contact Paula Harrison (email: paharriso@aol.com) Funded under the European Commission Sixth Framework Programme Contract Number: 036890

RUBICODE research themes Frameworks and concepts for the assessment of ecosystem services in terrestrial and freshwater ecosystems. Approaches for linking ecosystem service provision to functional traits. Socio-economic and environmental drivers of biodiversity change. Indicators for monitoring ecosystem services.   Strategies for conserving and managing biodiversity and the services it provides that take account of drivers of biodiversity change. Identification of current gaps in knowledge and future research needs.

Importance of ecosystem services Key: A: Agro-ecosystems; F: Forests; G: Grasslands; H: Heathlands; M: Mountains; S: Soil ecosystems; W: Freshwater ecosystems. Service Key contribution Some contribution No contribution Poorly known Provisioning services: Food, fibre, fuel/energy A, F, G, M, S,W H Fresh water F, M, W G, H A(-), S Regulatory services: Pollination A, F, G, H M, S W A, F, M Pest regulation A, S G, M H, W Water purification S, W F, G, H, M A(-) A, M Cultural services: Recreation/ecotourism A, F, G, H, M, W S Religious values A, F, M, S Supporting services: Provision of habitat A, F, G, H, M, S, W RUBICODE collected evidence on the variety and relative magnitude of the services provided by the main terrestrial and freshwater ecosystems in Europe. This table shows just a few examples of services from the four classes of the Millennium Ecosystem Assessment: provisioning, regulating, cultural and supporting. The relative importance of different services within an ecosystem has been ranked into four categories: significant contribution, some contribution, no contribution and contribution poorly known. This latter category helps distinguish where the ranking is based on expert opinion and requires further study to provide quantitative evidence. Harrison et al. (in press) Biodiversity & Conservation

Status of ecosystem services Status of selected ecosystem services in rivers and lakes in 2 time periods RUBICODE collected evidence on past trends in the status and human use of the services provided by the main terrestrial and freshwater ecosystems in Europe. This slide shows results for selected services in river and lake ecosystems. Aquaculture in Europe increased between 1950 and 1990, but then slightly decreased after 1990, along with freshwater capture fisheries, due to a declining supply of natural and aquacultural freshwater fish. The status of virtually all other services has become degraded over both time periods due to intensified land use causing increased water abstraction, physical modification of river courses, drainage and devastation of floodplains, and eutrophication. The loss of extended floodplain forests has also had negative implications for regional climate regulation services. Harrison et al. (in press) Biodiversity & Conservation

Service-providing unit (SPU) “The collection of organisms and their characteristics necessary to deliver a given ecosystem service at the level required by service beneficiaries.” Need to know: The sections of society that need/use the service (Ecosystem Service Beneficiaries - ESBs). The level at which it is required. The organisms that provide the service (Ecosystem Service Providers – ESPs). The characteristics of these organisms required to provide the service at the desired level (SPU). Information on how ecosystem services are provided is fairly limited. This includes approaches for understanding the interactions between biodiversity, ecosystem functioning and ecosystem services. One approach we have evaluated is the Service Providing Unit concept, which is defined as “the collection of organisms and their characteristics necessary to deliver a given ecosystem service at the level required by service beneficiaries”. The latter part of this definition “at the level required by service beneficiaries” is very important and highlights that we need to know which sections of the human community use the service (the Ecosystem Service Beneficiaries) and at what level is it required, in addition to what components of the ecosystem provide the service (the Ecosystem Service Providers), and what characteristics of these components are required to provide the service at the desired level (the Service Providing Unit). Luck et al. (2009) Bioscience, 59: 223-235

{ { 1. IDENTIFICATION 2. QUANTIFICATION 3. APPRAISAL Quantify the ecosystem service demand: determine the net level of demand/need for the service Quantify the service-providing unit (SPU): determine the characteristics of organisms necessary for service provision quantify the relationships between SPUs and service supply quantify the components of biodiversity that support the SPU Identify and value potential alternatives for providing the service Evaluate options: compare valuations and examine trade-offs determine implications for biodiversity conservation determine implications for policy and sustainable livelihoods Define the ecosystem service: identify the ecosystem service beneficiaries identify the spatio-temporal scale of service delivery identify the ecosystem service providers (ESPs) 1. IDENTIFICATION 2. QUANTIFICATION { Value the service as provided by the SPU 3. APPRAISAL RUBICODE has developed a conceptual framework showing the steps that need to be undertaken to identify and quantify an ecosystem service. The first step involves the identification of the human beneficiaries of the service, the spatial and temporal scale of service delivery and the species or ecological communities that provide the service. The second stage involves the quantification of the demand and supply of the service. Ecosystem service supply is quantified in terms of the service providing unit. The relevant SPU characteristics which need quantifying will depend on the service in question and the organism(s) that supply it, but could include population size, phenology, distribution, or functional traits. Once the relevant SPU characteristics have been defined, it is important to understand how incremental changes in these particular characteristics impact on service provision. The final appraisal stage then involves the valuation of the service as provided by the SPU and potential man-made alternatives and assessment of the implications for management and policy.   In RUBICODE, we undertook an extensive search of the literature to locate case studies to test this conceptual framework. We found about 60 studies which contained some information on the identification stage, but far fewer undertook any quantification of the service providing unit and detailed quantification of the level of need is rarely done.

Seed dispersal in an urban park Need: Cultural, recreational and biodiversity ‘value’ of park (15M visits/year). Eurasian Jay primary facilitator of acorn germination (85% of the oaks in the NUP are estimated to result from natural regeneration by the jay). Quantification (SPU): A minimum of 12 pairs of Eurasian Jay present each year for 14 years. Valuation: Estimate replacement cost of seed dispersal service = 16,880 €/pair. ctmsu.sytes.net One example where quantifying need is partly addressed is by Hougner et al. (2006) who describe the seed dispersal service provided by Eurasian jays in the oak forest in the National Urban Park of Stockholm, Sweden. First, they present general arguments of the cultural, recreational (e.g., 15 million visits/year) and biodiversity value of the park. They argue that the oak forest makes a substantial contribution to these values in addition to oaks being recognised as keystone species in the region. Second, they show that the foraging and dispersal behaviour of the Eurasian jay facilitates acorn germination to an extent much greater than any other animal species in the park. So, how many pairs of jays are required to maintain the oak forest? This defines the Service Providing Unit which is estimated to be a minimum of 12 resident jay pairs present each year for 14 years. Finally, the authors estimate the replacement cost of the seed dispersal service provided by jays (i.e., the cost of seeding or planting oak trees by humans), resulting in a replacement cost of nearly 17,000 Euros for each jay pair. Hougner et al. (2006) Ecol. Econ. 59, 364-74.

Buffering nutrient and sediment pollution of rivers Need: Riparian vegetation regulates the flow of nutrients and sediment from uplands to the stream. No practical alternatives. wider zone of riparian woody vegetation assimilates nitrate from the groundwater grass strip filters sediment particles and assimilates phosphorus ctmsu.sytes.net A more complex example whereby a service is provided by a multi-species plant community is shown in this slide for the service of regulating nutrient and sediment pollution of rivers. The service is provided by the multi-species-multi-zone riparian plant community (that is different trees, shrubs, herbs and grasses), but the service rate depends on the number, width and density of each zone. For example, 30 m of mixed riparian buffer has been found to remove 92-100 % of ground water nitrate and 5-20 m grass strips retain 40-100 % of sediments. Replacement costs if the service were to be provided by conventional waste water treatment systems have also been calculated for this example at between 61 and 108 Euros per person equivalent per year, but alternatives for the removal of nutrients are generally impractical and the removal of sediments is impossible. Hence, restoration of riparian buffers is unavoidable to meet the demands of the Water Framework Directive and improve river water and habitat quality. SPU = riparian plant community (different trees, shrubs, herbs and grasses). But: the service rate depends on the number of constituent zones density and width of buffer multi-zone riparian buffers are most effective!

Social-Ecological System Drivers e.g. Economy Demography Society Technology (exogenous) Pressures e.g. Climate change Land use change Air pollution (endogenous) States Ecosystem service beneficiaries (ESB) Supporting system service providers (ESP) Impact on service provision Responses Policy, strategic decisions and management strategies Baseline/Futures Service Providing Units (SPUs) Valuation of services and alternatives Trade-offs Mitigation Social-Ecological System Adaptation The conceptual framework which assesses how ecosystem services are provided using the Service Providing Unit concept has been integrated within a drivers framework to investigate what will happen to services in the future. The integrated framework is known as FESP (Framework for Ecosystem Service Provision). Drivers and pressures, such as climate change, land use change and air pollution, are represented by multiple scenarios which explore different plausible futures. The Drivers are exogenous to the system. This means they are influenced primarily by factors, processes and interactions that occur outside of the ecosystem under consideration. The Pressures represent the variables that act directly upon the ecosystem State. This is represented by the variables that are important for service provision, that is, the human beneficiaries of the service, the species providing the service and the system that supports these species (often their habitat). As the state of these 3 elements change it may reach a certain threshold which defines the Service Providing Unit, above which service provision is at a level demanded by the service beneficiary, below which it is not. The impact is assessed using valuation techniques, including for example, trade-offs with alternative approaches to service provision. Responses, such as policy measures and conservation management, are then implemented in accordance with the measured costs of the impact. Rounsevell et al. (in press) Biodiversity & Conservation

The Stockholm Urban Park (Socio-ecological system) States Oaks & Coniferous forest (Supporting) Drivers Macroeconomics, EU regulations/policies Global climate change Consumer trends Technology (exogenous) SPU threshold (12 breeding pairs) Scenarios Urban Population (ESB) Jays (ESP) Pressures Land cover changes Local climate, Local air, water, soil pollution Alien species, Increases/decreases in visitors (endogenous) Provision of cultural & aesthetic services Storylines This framework is briefly illustrated in this slide using the example shown in slide 7 for the seed dispersal service provided by Eurasian jays in Stockholm urban park. The exogeneous drivers include macroeconomics, global climate change, consumer trends and EU policies. These alter one or more of the pressures, such as local climate and changes in visitor numbers to the park, which have a direct influence on the ecosystem service. The ecosystem service provider is defined as the jays themselves, which require the oaks and coniferous forest (for nesting) as supporting habitats, and the ecosystem service beneficiaries are the urban population of Stockholm supplemented by tourists. The Service Providing Unit which defines the threshold for providing the service at the level required by the beneficiaries is 12 jay pairs, so if the number of jays falls below this threshold it will lead to an impact on the provision of the cultural and aesthetic services provided by the urban park. The cost of alternatives, such as seeding or planting by humans, was calculated as nearly 17,000 Euros per jay pair. This informs the evaluation of possible responses, including continued investment in management that safeguards the jay population at a level suitable for the continued and successfully regeneration of oak forest. Planting or seeding by humans = 16,800 €/jay pair Adaptation (application/ implementation) Valuation of alternatives Responses Protection policies Seeding/planting regimes Mitigation policy Trade-offs

A functional approach to biodiversity A tool to reduce complexity Trait: a measurable character of an organism that has demonstrable links to the organism’s function Groups species or populations with a similar RESPONSE to environmental change and/or similar EFFECTS on ecosystem function as a result of shared characters = traits Traits may be morphological, ecophysiological, demographic, biochemical, behavioural… Ecosystem services are rarely dependent on single species. This presents something of a problem when trying to predict the impacts of environmental changes on service provision, as even quite closely related species often respond very differently to particular changes. A functional approach is one possible way in which to tackle this challenge. This involves grouping ecosystem service providers according to their relevant characteristics or traits in two ways: Response traits determine how organisms respond to change whilst Effect traits define those characteristics responsible for providing the service of interest. Relevant plant traits may be morphological (e.g. seed size), ecophysiological (e.g. cold hardiness), demographic (e.g. life cycle type) or biochemical (e.g. photosynthetic pathway type). In the case of animals, behavioural traits are also often important.

Linking traits to ecosystem services RUBICODE collected around 250 published studies showing effects of functional traits on various ecosystem services, and the underlying ecosystem processes. Analyses of the data within these studies showed predictable clusters of effect traits and services. Here you can see the analysis discriminated services mediated by plant and soil organisms (top left cluster), from those mediated by aquatic organisms (bottom left) and by other insects (centre right). De Bello et al. (in press) Ecosystems

Environmental / management pressure Trophic Effect Traits Response Functional Ecosystem service TROPHIC LEVEL 1 TROPHIC LEVEL 2 Environmental / management pressure Pressure Pressure Response Traits The provision of services is also often affected by species from different trophic levels. RUBICODE has developed a framework for this purpose, which considers the traits affecting the interactions between trophic levels as well as those providing the service (functional effect traits) and those affecting response to change (pressure response traits). Thus, trophic effect traits are introduced as those that influence the next trophic level up, and trophic response traits determine how that trophic level reacts. For example, the trophic effect traits might be plant defence mechanisms and the trophic response traits might be the traits that allow insects to overcome those defences. Lavorel et al. (in review) Functional Ecology

Indicators of ecosystem services Nutrient cycling: 78% Water retention: 36% Fresh water: 81% RUBICODE has also reviewed over 600 papers on indicators which could be used to monitor the condition of ecosystems (and related biodiversity and ecosystem services). Results show that most biodiversity indicators are based on measures of species and community richness across ecosystems, for example number of taxa or diversity indices, whilst other components are much less studied, particularly genetic, structural and functional indicators. The potential application of traits as indicators of ecosystem services also warrants further investigation. Ecosystem service indicators were mainly restricted to a few ecosystems, particularly nutrient cycling in soils and the provision of fresh water. Very few indicators were found for cultural services, and those that were available tended to relate to education and knowledge systems, and recreation. Feld et al. (2009) Oikos

Indicators of ecosystem services The relative proportion of indicators of provisioning, regulating and supporting services for different ecosystems type was analysed using a Principal Components Analysis. Results showed that most indicators refer to regulating and supporting services across ecosystems. Among the provisioning services, only the provision of fuel wood in forests and the provision of fresh water in lakes and rivers are frequently addressed. The predominant regulating service is water retention which is frequently addressed by studies on forest and grassland/shrubland ecosystems. The majority of soil and wetland indicators refer to the supporting services of nutrient cycling and decomposition.

Framework for integrating ecosystem services into conservation (part 1) Human aesthetic, cultural and moral values Species/habitat protection Conservation policy and management strategy RUBICODE has undertaken two reviews on European habitat management strategies and the effectiveness of existing conservation policy. We also held a stakeholder workshop in Slovenia in April which discussed strategies for a new dynamic approach for conservation focused on ecosystem service provision. The information from all these sources has been summarised within a draft conceptual framework for integrating ecosystem services into conservation strategies. This slide illustrates the conventional approach to conservation. It is our aesthetic, cultural and moral values that provide the stimulus for conserving nature, which requires policies and appropriate management strategies. This has led to the present situation of Protected Areas in ecological networks for the conservation of species and habitats. Visiting these Protected Areas or seeing photographs of them reinforces our aesthetic appreciation and the value of feeling somehow “close to nature”. Static site-based PAs & networks

ES & conservation framework (part 2) Ecosystem service provision Societal needs Human aesthetic, cultural and moral values Species/habitat protection Conservation policy and management strategy Sectoral policy and management Societal needs from nature are much broader than just aesthetic and cultural values. Supply of provisioning, regulatory and supporting services at levels required by the service beneficiaries, while also protecting biodiversity, requires that sectoral policy and management for ecosystem services be integrated with conservation within entire socio-ecological systems and these systems managed to provide all societal needs sustainably. This is seen in the outer loop. However, there may also be services whose provision will be antagonistic to biodiversity conservation interests or to other services. If left to run in isolation, or unsustainably, this loop may have severe detrimental effects on biodiversity that is not required to provide non-cultural services. Thus it is of utmost importance that both loops of the framework are maintained and equally important, that the loops not be considered in isolation of each other, but must be closely linked in all appropriate places and at all scales of organisation.   All this implies an acute awareness of the dynamic nature of ecosystems and our societal interactions with them – change to any part of the system, biological or socio-economic, from within or external, is likely to have profound consequences for the other components and their relationships. This re-emphasises that it would be naïve to continue to consider biodiversity conservation as something on its own; and that entire Socio-Ecological Systems (SES) are the appropriate level for responding to future conservation needs. Ecosystem sustainability and integrity Static site-based PAs & networks Management for sustainable ecosystem services Conservation within socio-ecological systems Haslett et al. (in press) Biodiversity & Conservation

Thank you from the RUBICODE team!