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Open Access Linking key environmental stressors with the delivery of provisioning ecosystem services in the freshwaters of southern Africa Michelle C. Jackson1,2,3, Darragh J. Woodford2,4 and Olaf L. F. Weyl2,5 Societies’ growing global footprint is causing a rapid increase in the demand for natural resources (i.e. ecosystem services), while also reducing the capacity of ecosystems to provide them. Freshwater ecosystems contribute disproportionately to ecosystem services but are also particularly vulnerable to global environmental change. The provisioning of freshwater services, such as water and food production, is especially important in developing countries. Here, we review the evidence which demonstrates the impacts of key environmental stressors on these two important provisioning services in southern Africa. Land use change, species invasions and climate change can all be linked to a loss of the provisioning services provided by freshwater ecosystems in southern Africa. Water resources for drinking, agriculture, sanitation and power are expected to decline as a result of both climate and land use change. Fish production may be negatively or positively affected by the different stressors, highlighting the high context-dependency associated with their impacts. Evidence also suggests that these stressors can interact to alter one another’s impacts or promote the proliferation of further stressors. For instance, land use change can promote aquatic plant invasions and, subsequently, the stressors may interact synergistically to cause fish kills. Stressors may also interact to mitigate one another’s impact, for instance fish invasions may enhance total fish catch following a pollution event. Since stressors are unlikely to occur in isolation and multiple stressors frequently result in complex ‘ecological surprises’, it is urgent that we increase research effort on the links between multiple stressors and the loss of ecosystem services. Future research should, therefore, focus on the combined impacts of multiple environmental, social, and economic stressors on natural resources and provisioning ecosystem services in southern Africa. Key words climate change; fisheries; freshwater; invasive species; land use change; multiple stressors 1 Centre for Invasion Biology, Department of Zoology and Entomology, University of Pretoria, Private Bag X 20, Hatfield 0028, South Africa South African Institute for Aquatic Biodiversity, Grahamstown 6140, South Africa 3 Department of Life Sciences, Imperial College London, Silwood Park Campus, Buckhurst Road, Ascot, Berkshire SL5 7PY 4 Centre for Invasion Biology, School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Private Bag 3, Wits, Johannesburg 2050, South Africa 5 Centre for Invasion Biology, South African Institute for Aquatic Biodiversity, Grahamstown 6140, South Africa E-mail: [email protected] 2 Revised manuscript received 7 September 2016 Geo: Geography and Environment, 2016, 3 (2), e00026 Introduction Ecosystems provide people with numerous important ‘goods and services’ such as food, water and fuel, while also providing indirect benefits including flood mitigation and climate regulation (MA 2005). Societies’ growing global footprint is causing a rapid increase in the demand for these natural resources, while also reducing the capacity of ecosystems to provide them ( Schmidhuber and Tubiello 2007; Mooney et al. 2009). Despite covering less than 1% of the earth’s surface, freshwater ecosystems contribute disproportionately to ecosystem services (Dudgeon et al. 2006). Lakes, rivers and wetlands are vital for human survival and sanitation, for safe drinking water and through the provisioning of food and power. Freshwater systems are also especially vulnerable to environmental stressors and over exploitation, with water and fish protein growing in importance as commodities, and average species population declines since 1970 estimated at 76% (WWF 2014). This vulnerability means that freshwater ecosystems can act as local ‘ecosystem canaries’, or The information, practices and views in this article are those of the author(s) and do not necessarily reflect the opinion of the Royal Geographical Society (with IBG). ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. Page 1 | 2016 | Volume 3 | Issue 2 | e00026 Michelle C. Jackson et al. Page 2 | 2016 | Volume 3 | Issue 2 | e00026 sentinels to environmental perturbations occurring at larger scales (Woodward et al. 2010). The provisioning of ecosystem services is especially important in developing countries, where many rural and poor communities rely heavily on natural resources for survival and income (Egoh et al. 2012). Communities in southern Africa rely on freshwater ecosystems for critically important provisioning services, such as drinking water and food. Inland fisheries in Africa produce the second highest catch by continent (Figure 1a) and are particularly important in Uganda, Tanzania, The Democratic Republic of Congo and Kenya (Figure 1b). Furthermore, only 58% of the population in sub-Saharan Africa are thought to have access to safe drinking water (Banerjee and Morella 2011), and delivery of such services is becoming progressively more at risk due to stressors such as land use change, species invasions and climate change, as well as increasing demand (Schmidhuber and Tubiello 2007; Egoh et al. 2012). Southern Africa also has low food security and little legislation to control the use of natural resources (Egoh et al. 2012), while also experiencing high rates of human population growth (World Bank 2015). With the growing demand for resources and an increase in environmental perturbations, it is ever more important that we understand the impacts of Figure 1 Trends in inland freshwater fish catch by (a) continent, and (b) the 10 countries in Southern Africa with the highest catch Source: Data taken from The Food and Agriculture Organization of the United Nations’ global capture production database FishStatJ (http://www.fao.org/fishery/statistics/ software/fishstatj/en) environmental stressors on ecosystem services. Here we give an overview of the impact of three key stressors in freshwaters (land use change, species invasions and climate change) on provisioning ecosystem services in southern Africa; before discussing how these stressors may interact. We provide the first framework which considers how multiple stressors interact to alter provisioning ecosystem services and discuss management applications. We focus on access to water (for drinking, power, agriculture, sanitation) and food (i.e. fisheries) as provisioning ecosystem services because of their importance, particularly in developing countries, and consider any country in mainland Africa which is fully, or in part, in the southern hemisphere. Land use change The impacts of land use change, in the form of both physical habitat change and pollution, have been the largest drivers of losses in global biodiversity and ecosystem function in inland freshwater ecosystems over the last 50–100 years (MA 2005). Southern Africa, historically a region with relatively low population densities where food production was based on subsistence agriculture and pastoralism (Bloom et al. 1998; DeFries et al. 2004) has over the last century seen rapid shifts in economic activities and increased urbanisation (Bryceson 1996; Darkoh 2009). These altered economic activities have precipitated rapid changes in land use, often resulting in measurable degradation of both terrestrial and aquatic ecosystems (Reardon et al. 1999; Wasige et al. 2013). While land use change is rapid and severe in some places (Reid et al. 2000; Wasige et al. 2013), the economic activities underpinning urban development are stunted in many regions by unfavourable geography and other demographic drivers such as infectious diseases (Bloom et al. 1998). This means there remain many rural and peri-urban communities that directly depend on the provisioning ecosystem services provided by local aquatic ecosystems, which are in turn, threatened by regional developments. Land use change in rural areas is characterised by deforestation and expanding land under cultivation (Wasige et al. 2013). This form of land cover change can have several direct and indirect impacts on aquatic ecosystems. The most direct impacts of expanded or intensified agriculture are experienced by wetlands, which provide a wide range of ecosystem services that are directly affected by the removal or degradation of the wetlands (Rebelo et al. 2010). In East Africa, wetlands are a critical source of potable water to rural communities, in one documented case (the Yala Swamp in Kenya) providing 100% of drinking and washing water (Schuyt 2005). These ecosystems also provide raw materials for roofing, furniture and fish traps, as well as cropping and grazing land in the dry season and ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) Environmental stress and ecosystem services seasonal or permanent fisheries (Chapman et al. 2001; van Dam et al. 2011). The rapid encroachment of intensified agriculture, characterised by removal of wetland vegetation and hydrological alterations, is leading to a degradation of these resources (van Dam et al. 2011). The direct utilisation of water resources, by way of water abstraction or diversion, can play a substantial role in degrading downstream provisioning ecosystem services. Irrigation has historically been the primary recipient of such diverted supplies in Africa, comprising 80% of all water abstractions (Machibya and Mdemu 2005). While irrigated agriculture can drive significant socio-economic development in rural communities, downstream users can be negatively impacted, if they are disenfranchised from the upstream irrigation scheme, or depend on the system’s natural, preabstraction hydrology to receive the ecosystem services (Oosterbaan 1984). For example, a government scheme to upgrade traditional rice paddy irrigation infrastructure in the Usanga Basin, Tanzania resulted in an inequitable allocation of water resources favouring upstream users at the expense of downstream users (Machibya and Mdemu 2005). While many forms of water utilisation infrastructure exist, it is the construction of on-channel dams, a major consequence of agricultural intensification and urban expansion throughout southern Africa, which has had the most widely documented negative impact on downstream services. In the 1960s and 1970s, many countries invested in large dams to promote water security and produce hydropower (Schuyt 2005). While in some cases these projects have given rise to new provisioning ecosystem services above the dam wall, such as the commercial lift-net and small-scale light fisheries which have been established to harvest some 35 000 t year 1 kapenta Limnothrissa miodon (an introduced small pelagic fish species), from reservoirs in the Zambezi Basin (Ellender and Weyl 2014), the net change in ecosystem services below the wall appear to be consistently negative. The major impact on provisioning ecosystem services derives from reduced floods and sediment transport. In the Tana River delta in Kenya (Leauthaud et al. 2013), the Zambezi River in Mozambique (Davies et al. 1975) and the Kafue River in Zambia (Richter et al. 2010), for example, altered river hydrology has resulted in collapsed fisheries, degraded grazing land productivity and, in some cases, the need for local communities to shift from migrant livestock grazing to crop farming. Small, privately built dams can also have negative effects on downstream ecosystem services. A decrease in natural flooding caused by a proliferation of small upstream dams was shown to exacerbate bush encroachment in floodplain grazing land in semi-arid tributaries of the Limpopo River, South Africa, thus accelerating the depletion of this ecosystem service (O’Connor 2001). Page 3 | 2016 | Volume 3 | Issue 2 | e00026 While large dams can have the impact of decreasing the supply of sediments to downstream rivers, broader landscape changes such as deforestation, overgrazing, and poor farming practices that cause erosion of topsoil, all increase sediment loads to water bodies (Olago and Odada 2007). In the tributaries of Lake Victoria for example, navigability for fishing boats is disrupted directly by the sedimentation of stream channels (Olago and Odada 2007) while in Lake Tanganyika, sediment loading has depleted both fish diversity and densities in shallow shoreline areas (Cohen et al. 1993). In Lake Malawi, fish populations that are already under severe pressure of overfishing are increasingly pressured by increased sediment inputs resulting from poor land use practices (Weyl et al. 2010). Worst affected were the endemic potamodromous cyprinids, Labeo mesops and Opsaridium microlepis, whose decline since the 1930s has been linked primarily to alterations in flow regime and sedimentation of spawning areas resulting from catchment degradation (Weyl et al. 2010). Lake-dwelling fish species are negatively affected by increased turbidity which decreases light penetration and reduces overall productivity, as well as from the smothering of demersal algae and fauna by settling sediment (Weyl et al. 2010). Eutrophication, the oversupply of organic nutrients, is another major problem in Lake Victoria, caused by the inflow of fertile top soils eroding from the transformed farmland, as well as sewage effluents from urban developments along the shoreline (Olago and Odada 2007; Nyenje et al. 2010). It results in high biological oxygen demands from organisms such as blue-green algae, which can lead to hypoxia-driven fish kills, threatening local fisheries (Olago and Odada 2007). Similar fish kills have been reported in Lake Tanganyika (Odada et al. 2003), and are also recognised as a threat to Lake Malawi’s fisheries (Dudgeon et al. 2006). Additionally, eutrophication is recognised as a major threat to potable water supplies to communities along lake shorelines (e.g. Odada et al. 2003). The two major sources of this pollution are treated urban wastewater, which remains high in organic nutrients such as nitrogen and phosphorous; and untreated sewage emanating from damaged or overwhelmed wastewater treatment infrastructure (Nyenje et al. 2010). The latter brings an additional threat to the safety of drinking lake water by increasing microbial toxicity (Odada et al. 2003) and increasing the risk of exposure to infectious diseases (Momba et al. 2006). Despite the proportionately high impact of urban wastewater on water resources, certain industrial effluents also have significant negative impacts on water supply in certain regions. For instance, the capital of Uganda, Kampala, has several industries, including abattoirs and small-scale metal industries, which discharge untreated waste directly into local water ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) Page 4 | 2016 | Volume 3 | Issue 2 | e00026 bodies (Matagi 2002). Another significant industrial threat to water resources across southern Africa is that of mining, which has a long history of creating environmental problems (Boocock 2002). For example, the Kafue River in the Copper Belt of northern Zambia receives significant inputs of copper, cobalt and iron oxides from mine tailings, often exceeding international water quality standards (Sracek et al. 2012). Similarly, in the Witwatersrand gold-mining region in South Africa, water bodies are being contaminated by heavy metals, including uranium radio-isotopes, emanating from mine tailings deposits (Winde and van der Walt 2004), and acid mine drainage, whereby chemical reactions between water and exposed iron pyrite lower the pH of receiving streams to toxic levels (Akcil and Koldas 2006; McCarthy 2011). All of these chemical pollutants are likely to increase the burden on water treatment works in the future, depleting the ability of these aquatic ecosystems to safe drinking water to ever growing human populations. Invasive species As is the case elsewhere, alien invasive species are having increasing impacts on aquatic ecosystems in southern Africa. Most severe are the negative impacts of invasive terrestrial and aquatic plants. In South Africa, for example, colonisation of riparian zones by invasive trees, such as wattle Acacia mearnsii, gums Eucalyptus spp. and pines Pinus spp. are an ongoing problem (Richardson et al. 2010). Impacts on freshwater systems are linked to the higher evapo-transpiration rates of alien trees relative to those of the indigenous vegetation that they replace, resulting in reduced river flows and groundwater reserves (Malan and Day 2003). For river catchments receiving >800 mm rainfall per year, Cullis et al. (2007) estimated that the total loss of usable water resulting from alien plant invasions was equivalent to 4% of the total registered water use in South Africa. In a water-poor country such as South Africa, such reductions in flow can have severe consequences for water provisioning in both rural and urban areas. Aquatic plant invaders include water hyacinth Eichhornia crassipes, water lettuce Pistia stratiotes, Kariba weed Salvinia molesta and red water fern Azolla filiculoides, all of which increase rapidly in biomass resulting in the formation of dense mats in nutrient loaded environments (Chamier et al. 2012). While this can decrease nutrient loads, the resultant mats reduce light penetration and photosynthesis, reduce water circulation and the diffusion of air into water, and increase organic detritus loads which are broken down by anaerobic processes (Chamier et al. 2012). Collectively this often results in anoxic conditions that are uninhabitable by aquatic invertebrates Michelle C. Jackson et al. and fishes. On the Ugandan side of Lake Victoria, for example, stationary fringes of invasive plants covered much of the shoreline in the mid 1990s and resulted in significant socioeconomic and environmental impacts, including disruption of transport, tourism and fishing, negative impacts on water quality, and spread of waterborne diseases (Balirwa et al. 2003). This increased disease incidence is linked to improved habitat (and therefore increased abundance) for snails (Biomphalaria spp. and Bulinus spp.) which act as intermediate hosts for schistosomiasis (Masiwa et al. 2001), and mosquitoes (e.g. Anopheles spp.), which are vectors for diseases such as dengue fever, elephantiasis, encephalitis and malaria (Oliver 1993). Non-native fishes have been introduced into southern Africa for more than two centuries. In South Africa alone, a total of 55 fishes have been introduced into novel environments for angling, aquaculture, fisheries enhancements, as aquarium fish, and for the control of mosquito larvae (Ellender and Weyl 2014). While many introduced fishes now contribute significantly to local economies through fisheries and aquaculture, they also impact on native biota through direct predation, competition, hybridisation introduction of pathogens and environmental modification (Ellender and Weyl 2014). Nile perch Lates niloticus were introduced into Lake Victoria in 1954, and in the 1980s their populations increased explosively and their predation on native fishes resulted in the extinction of approximately 200 species which is considered to represent the largest extinction event among vertebrates during the twentieth century (Goldschmidt et al. 1993). The species has however had considerable economic impact and its introduction resulted in an increase in fisheries production from around 100 000 t year 1 in 1980 to about 1 000 000 t year 1 at present (Mkumbo and Marshall 2015). The fisheries as a whole now supports some 4 million people and Nile perch exports, mainly to Europe, are worth about US$350 million annually (Mkumbo and Marshall 2015). In South Africa, introduced common carp Cyprinus carpio are the main target species for an estimated 15 million freshwater anglers, who target this species for both recreation and subsistence (Ellender and Weyl 2014; Weyl and Cowley 2015). Carp therefore contribute not only to the local and national economy but are also a source of food security for the rural poor (Ellender and Weyl 2014). However, in South Africa, common carp is the primary vector for introduced parasites and diseases which affect native sport fishes such as the smallmouth yellowfish Labeobarbus aeneus which are also economically important (Ellender and Weyl 2014). Although the effects of C. carpio have not been quantified in southern Africa, the bottom-grubbing feeding strategy employed by this species alters invaded ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) Environmental stress and ecosystem services habitats through the resuspension of sediments which increases nutrient loading (i.e. eutrophication) and increases turbidity (Lougheed et al. 1998), which has resulted in additional costs associated with water treatment in South Africa (De Moor and Bruton 1988). Climate change The Earth’s surface has already experienced warming of 0.85 °C in the period from 1880 to 2012 (calculated from global combined land and ocean surface temperature; IPCC 2013), causing shifts in species geographic ranges, seasonal activities, migration patterns, abundances and interactions (Parmesan and Yohe 2003; Bellard et al. 2012; IPCC 2013). Future projections vary based on different socio-economic scenarios, but suggest warming of at least 2 °C by 2100 (IPCC 2013). Climate change will cause shifts in the seasonal patterns and amount of precipitation, while also increasing the number and severity of extreme weather events, such as drought and flooding (IPCC 2013). In southern Africa, these changes are expected to create stress on water resources, crop productivity and food security, and an increase in waterborne diseases (IPCC 2013). Moreover, Africa is considered one of the most vulnerable regions to global climate change (Magadza 1994; Schmidhuber and Tubiello 2007; Egoh et al. 2012; IPCC 2013) and communities are already experiencing its affects (e.g. Bunce et al. 2009). By the end of the twenty-first century most of the region is expected to experience a 10% decline in annual rainfall, with this being as high as 20% in Namibia (de Wit and Stankiewicz 2006). This will reduce access to fresh drinking water, impair sanitation and perhaps increase conflict as water becomes an increasingly valuable resource (de Wit and Stankiewicz 2006). However, projections for the north-eastern equatorial parts of southern Africa suggest that the region will experience benefits from climate change, through increased precipitation, river runoff and fresh water availability (de Wit and Stankiewicz 2006; Doherty et al. 2010). Nonetheless, this may also increase flood risk, soil erosion and pathogen loads (Doherty et al. 2010; IPCC 2013), and climate change is still likely to exacerbate the broader region’s overall water scarcity considerably (Milly et al. 2005; Schewe et al. 2014). Water shortages and accessibility to water are already ongoing problems in many parts of southern Africa (Banerjee and Morella 2011). For instance, in the Usangu plain in south-western Tanzania, only 13% of households have full-time access to safe drinking water while the national average is just 66% (Malley et al. 2009). Stressors associated with climate change are blamed for this scarcity (Malley et al. 2009), and the same water shortage scenario occurs in many communities throughout the arid regions of southern Africa. A global prioritisation scheme, which selected Page 5 | 2016 | Volume 3 | Issue 2 | e00026 the 10 most vulnerable watersheds for ecosystem service delivery, ranked 50% of them in Africa, including Rufiji in Tanzania (Luck et al. 2009). Like many watersheds in the region, this floodplain is vital for the food security and nutrition of local communities, but is expected to be impacted by global climate change. A change in the availability of ground and surface water, due to shifts in the amount or timing of seasonal rainfall, also negatively affects agriculture. Many rural communities in southern Africa rely on annual floods to increase soil productivity, and rain or local water bodies for irrigation (Cooper et al. 2008). In the past, severe droughts have caused a decline in commercial crop yields, impacting the economies of South Africa, Malawi and Zambia (Jury 2002; Thurlow et al. 2012). This is expected to be aggravated by increases in temperature, which may increase irrigation requirements and further water stress (Morton 2007). Future projections of important crop yields (including maize and millet) under warming scenarios suggest a considerable decline in productivity (Schlenker and Lobell 2010). Alternatively, climate change may cause extreme flooding events, resulting in widespread devastation to agricultural land and threatening food security (Conway et al. 2015). A decline in water resources may also have implications for electricity production from dams (Harrison and Whittington 2002; Conway et al. 2015). For example, Yamba et al. (2011) predicted a reduction in hydropower generation over the next 60 years for the Zambezi River Basin due to droughts and reduced reservoir storage capacity. Fish protein is important for both food and income in many parts of southern Africa (Figure 1), and globally, the countries with fisheries which are the most vulnerable to climate change are also the poorest, including many in this region (Allison et al. 2009). Although annual catch in inland fisheries has increased over time (Figure 1), this is due to increased effort and the growing population escalating demand. At a local scale, many fisheries have seen a decline in fish stocks due to this growing demand, which is exacerbated by climate change. Nine countries in southern Africa (Angola, Burundi, Democratic Republic of Congo, Mozambique, Malawi, Uganda, Tanzania, Zambia and Zimbabwe) have fisheries ranked as highly vulnerable to climate change in a global assessment (Allison et al. 2009). Given that six of these are land-locked countries with high inland fishery yields (Figure 1b), this implies that inland fisheries in southern Africa are especially vulnerable to climate stressors. A study in Tanzania, which used lake sediment cores to measure changes in productivity over time in Lake Tanganyika, suggested that climate change has caused a 20% decrease in primary productivity, implying around a 30% decrease in fish yields (O’Reilly et al. 2003). Similarly, Lake Chilwa in Malawi has seen fish ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) Michelle C. Jackson et al. Page 6 | 2016 | Volume 3 | Issue 2 | e00026 production decline dramatically since 1970 (Njaya et al. 2011). Inland fisheries in smaller lakes will be affected by declines in precipitation (and subsequently water levels) because there will be fewer opportunities for fishing and aquaculture (Allison et al. 2009). For instance, Lake Naivasha in Kenya has unpredictable lake level fluctuations which threaten the small fishery (Becht and Harper 2002). The lake level has been falling consistently since 1980 and there has been a linked decline in annual fish catch (Hickley et al. 2004). Other stressors associated with climate change, such as sea-level rise, increasing salinity and changes in weather patterns, will also have an impact on provisioning ecosystem services in southern Africa (Ngoran et al. 2015). Extreme weather events and unpredictable wet seasons may cause damage to assets and decrease the ability of communities to plan seasonal fishing activities (Allison et al. 2009). Changes in wind patterns will also have implications for inland fisheries, by altering lake mixing and stratification. Moderate mixing can promote fishery production by allowing nutrient influx into the water column. Stronger or weaker winds, however, can suspend anoxic sediments or cause low levels of dissolved oxygen, respectively (Kaufman et al. 1996; Ficke et al. 2007). For example, in Lake Victoria decreased mixing has led to low levels of dissolved oxygen and, subsequently, fish kills (Kaufman et al. 1996). Multiple stressors With accelerating change in freshwater systems (e.g. Aeschbach-Hertig and Gleeson 2012; Jackson and Grey 2013) it is becoming increasing unlikely that environmental stressors will occur in isolation (Jackson et al. 2016). Many freshwater ecosystems will simultaneously be affected by local, catchment-level, and global stressors, such as species invasions, land use change and global warming, respectively. However, if, how and when these stressors interact to impact ecosystems is still largely unknown (Piggott et al. 2015; Jackson et al. 2016). There are a variety of potential stressor interactions (Table I, Figure 2) which may alter ecosystem services in different ways depending on, for instance, stressor identity, community structure, the type of service, and the lag period between stressor events (Jackson et al. 2016). This high context dependency means it is extremely difficult to predict stressor interactions and the subsequent consequence for ecosystem services. Stressors can interact to create impacts which are equal to (additive interactions), smaller than (antagonistic interactions), or larger than (synergistic interactions) their single effects (Table I, Figure 2). Non-additive antagonistic interactions can occur when the impact of one stressor is so large, it overshadows that of the second stressor (Jackson et al. 2016). For instance, a severe drought might overshadow any impacts of invasive trees on water availability. Alternatively, antagonistic interactions can result from co-tolerance or acclimation of service delivery to stressors (Jackson et al. 2016). Synergistic interactions of stressors are often considered the most damaging because of the amplification of their independent impacts (Jackson et al. 2016). These can occur when the impact of two stressors are larger than predicted, perhaps because an ecosystem service is only vulnerable when both stressors are present (Table I, Figure 2). For example, flooding of cropland from extreme weather events might worsen with land cover change from deforestation because of reduced ability of the soil to hold water (Douglas et al. 2008; Wheater and Evans 2009). More complex stressor interactions may mitigate one another’s impacts on ecosystem services (Table I, Figure 2). For example, in Lake Victoria pollution was causing a decline in native fish stock and, although the introduction of tilapia and Nile Perch exacerbated this at a species level (Witte et al. 1992), their introduction boosted fishery catches overall (Mkumbo and Marshall 2015). However, a recent review by van Zwieten et al. (2016) reassessed Lake Victoria’s Nile perch takeover and concurrent reduction in native fishes and suggested that increased eutrophication (driven by catchment degradation) resulted in changes in algal and zooplankton composition, decreased water transparency, and widespread hypoxia, which most likely caused an initial native fish population crash. This then facilitated the rapid increase in Nile perch by reducing predation on larval Nile perch by native zooplanktivorous cichlids. Eutrophication in Lake Victoria has also promoted the proliferation of invasive water hyacinth, causing further anoxic fish kills in the lake (Figure 3; Odada et al. 2004; Olago and Odada 2007). This type of multiple stressor interaction, which we term successive interactions (Table I), involves one Table I Types of multi-stressor interactive impacts on ecosystem services Interaction type Additive Antagonistic Synergistic Mitigating Successive Description The impact of two stressors occurring simultaneously equals the sum of their single effects The impact of two stressors occurring simultaneously is less than the sum of their single effects The impact of two stressors occurring simultaneously is more than the sum of their single effects One stressor modulates the impact of a second stressor One stressor causes a second stressor to occur ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) Environmental stress and ecosystem services Page 7 | 2016 | Volume 3 | Issue 2 | e00026 Figure 2 A schematic highlighting some of the potential positive (‘Increased delivery’) and negative (‘Reduced delivery’) effects of three key stressors on freshwater ecosystem services in Southern Africa. The ecosystem services considered are (a) water resources, and (b) fisheries. Some of the possible consequences of interactions between the stressors are also given (‘Example multi-stress interaction’) ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) Page 8 | 2016 | Volume 3 | Issue 2 | e00026 Michelle C. Jackson et al. (3) monitoring of stressor events and ecosystem service delivery over time (e.g. Durance et al. 2016); and, modelling approaches which may provide theory which can later be tested for realism (e.g. Segner et al. 2014). Management of freshwater systems Figure 3 A case study of the interactions between multiple stressors and subsequent influence on fishery catch in Lake Victoria, East Africa. Eutrophication in the lake, resulting from high nutrient inputs in the surrounding catchment, causes anoxic fish kills while at the same time triggering the proliferation of invasive water hyacinth. This caused a successive and synergistic multi-stressor interaction whereby the increase in hyacinth biomass triggered further anoxic fish kills in the lake. In addition, the introduction of non-native Nile Perch caused a dramatic decline in native fish biodiversity but boosted the overall fishery catch in the lake, benefitting the surrounding populations stressor causing a second stressor to occur, and in this case, land use change promoting species invasions. However, since the two stressors also caused fish kills which were greater than the sum of their single effects, this is also a synergistic interaction (Figure 3). Another example of a successive interaction has been documented in Morogoro Tanzania, where climate change has forced households to extend or intensify agriculture and migrate to new areas, causing water resources to be depleted or degraded (i.e. climate change promoting land use change; Paavola 2008). These complex and non-additive impacts of multiple stressors have been termed ‘ecological surprises’ since they are hard to predict without context-dependent data (Jackson et al. 2016). Furthermore, nonenvironmental stressors (e.g. social and economic stressors) may also interact with the environment to exacerbate or mitigate impacts on ecosystem services. We suggest that future research goals should be directed towards understanding the multiple responses of ecosystem service delivery to these multiple, and potentially interacting, stressors. A multitude of approaches will be needed to achieve this goal including (1) controlled factorial experiments involving more than one stressor (Jackson et al. 2016); (2) space for time substitution field studies (Newbold et al. 2015); In southern Africa more than 50% of the population live below the poverty line (World Bank 2015). Issues of food security, livelihood provision, poverty alleviation, and economic development are therefore national policy objectives in many countries (Weyl et al. 2007). Adaptation to global change is therefore an important management issue, particularly with the multiple responses that may arise from multiple stressors. Aquatic resources provide considerable goods and services and inland fisheries in particular are an important safety net for the rural poor (Figure 1; Welcomme 2011; Béné et al. 2016). In some countries such as Malawi, the anthropogenic impacts are so profound that recovery of the catchment may be an unachievable management goal (Weyl et al. 2010). Nonetheless, in other countries there may still be opportunities to better manage catchments by involving communities in such a manner that the benefits of conservation outweigh the losses of their existing unsustainable resource use (Weyl et al. 2010). While the ecosystem services concept is rapidly establishing itself as a framework through which to implement the sustainable use of freshwater resources, many of its principles already exist within southern African water policy in the form of an older concept, integrated water resource management (IWRM). Both concepts approach the management of freshwater resources, by addressing competing interests such as socio-economic needs, environmental concerns, and human health considerations (Jewitt 2002; Malan and Day 2003; Cook and Spray 2012). There remains, however, a significant knowledge gap in understanding the fundamental interactions between environmental stressors and freshwater resources that, if not adequately addressed, could result in ineffective implementation of sustainable water resource management objectives (Jewitt 2002). Moreover, a disconnect between scientific knowledge, society and policy has hampered the implementation of IWRM in many southern African countries (van der Zaag 2005; Cook and Spray 2012). Scientific understanding of the complex interactions between multiple stressors thus cannot translate into improved management without strong co-operative governance between national agencies and local stakeholders, especially when the latter parties may both depend upon and negatively affect freshwater provisioning ecosystem services (van der Zaag 2005). An example of the need to understand both ecological and social requirements when addressing interacting ISSN 2054-4049 doi: 10.1002/geo2.26 © 2016 The Authors. Geo: Geography and Environment published by John Wiley & Sons Ltd and the Royal Geographical Society (with the Institute of British Geographers) Environmental stress and ecosystem services stressors on water resources can be found in the management of invasive riparian trees in South Africa. Research has demonstrated that these invasive species reduce surface flows through evapotranspiration, thus exacerbating the problems of equitable water allocation to multiple competing users downstream (Richardson et al. 2010; Forsyth et al. 2012). The government responded to this threat with the Working for Water Programme (www.dwa.gov.za/wfw/) which was launched in 1995 to alleviate the impacts of invasive species on water resources through multiple control measures (Richardson et al. 2010). This national-scale initiative aims to protect water supplies and restore productive land, primarily through labour-intensive invasive plant-clearing operations (Richardson et al. 2010). Between 1995 and the end of 2008 the programme had cleared invasive plants from more than 1.8 million ha of invaded land. Nonetheless, factors including the hiring of under-skilled contractors (in aid of poverty alleviation) with resulting ineffective clearing operations, and the emphasis on tackling as many species in as many regions as possible to maximise the social benefits of the programme, have led to significant inefficiencies in alleviating the stress on water resources in key catchments (van Wilgen et al. 2012). The Working for Water case study demonstrates that it is possible to prioritise efforts to manage interacting stressors (here, alien trees and water abstraction). In this case a multi-criteria approach called the analytic hierarchy process was used to gain consensus on which catchments should be invested in first for alien clearing based on current invasion levels, water demand from irrigators, and the local contractor capacity for ensuring successful clearing (Forsyth et al. 2012). In a broader water resource management context, prioritising which potentially interacting stressors to attack first will depend on understanding which threatened provisioning ecosystem services are most needed by particular communities. This presents a challenge, as different catchments will face a unique combination of interacting stressors, which will in turn affect provisioning ecosystem services that have greater significance to some communities than others (e.g. impact of siltation on floodplain farmers versus abstracting irrigators). Even once the prioritisation of interventions is made, the success of these interventions will still depend on strong, co-operative governance between government and local stakeholders to ensure the intended outcomes are achieved (van der Zaag 2005). Conclusions Land-use change, species invasions and climate change can all be linked to a loss of the ‘goods and services’ provided by freshwater systems in southern Africa. Evidence suggests that all three stressors can Page 9 | 2016 | Volume 3 | Issue 2 | e00026 independently reduce water availability and food security and may interact to alter these valuable provisioning ecosystem services. However, information on the impacts of stressors for this region is rarely quantitative, and there is an urgent need for more information in our changing world. This is particularly urgent given that stressors are unlikely to occur in isolation and multiple stressors frequently result in unpredictable and complex ‘ecological surprises’ (Jackson et al. 2016). It is also important to consider that the impacts of environmental stressors can be perpetuated by other concerns, such as war, economic policy, access to information and poor governance (Reid and Vogel 2006; Zimmerman et al. 2008; Bunce et al. 2009). Therefore, the large number of environmental, social and economic stressors, including the potential interactions among them, must all be considered in attempts to manage and conserve ecosystem services in southern Africa. The prioritisation of management efforts to mitigate against impacts of land-use change, invasive species, climate change, or the emergent effects of these stressors will first require a more explicit understanding and appreciation of the social-ecological context of these stressors at both local and regional scales. We therefore suggest there is an urgent requirement for research on the combined impacts of these environmental, social and economic stressors in southern Africa, and to effectively communicate the management implications of such research to policymakers across the region. Acknowledgements We thank Anson Mackay for inviting us to write this paper and two anonymous reviewers whose comments greatly improved the text. DJW (UID: 103581) and OLFW (UID: 77444) acknowledge the support of the National Research Foundation of South Africa. 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