Review TRENDS in Ecology and Evolution Vol.21 No.2 February 2006 Regional and global concerns over wetlands and water quality Jos T.A. Verhoeven1, Berit Arheimer2, Chengqing Yin3 and Mariet M. Hefting1 1 Landscape Ecology, Institute of Environmental Biology, Utrecht University, PO Box 800.84, 3508 TB Utrecht, the Netherlands Swedish Meteorological and Hydrological Institute (SMHI), SE-601 76, Norrköping, Sweden 3 Research Center for Eco-Environmental Science, Chinese Academy of Sciences, Beijing, P.R. China 2 Water quality in many stream catchments and river basins is severely impacted by nutrient enrichment as a result of agriculture. Water-resource managers worldwide are considering the potential role of riparian zones and floodplain wetlands in improving stream-water quality, as there is evidence at the site scale that such wetlands are efficient at removing nutrients from through-flowing water. However, recent studies have highlighted disadvantages of such use of wetlands, including emissions of greenhouse gases and losses of biodiversity that result from prolonged nutrient loading. Here, we discuss the water purification function of wetlands at the site and catchment scale and suggest ways in which these disadvantages could be overcome. Introduction Nutrient loading of terrestrial, freshwater and coastal ecosystems occurs as a result of human waste disposal and agriculture at a global scale. Although technological purification plants are the best option to reduce the nutrient fluxes to the environment, nutrient loading owing to intensive agricultural practices typically occurs through diffuse or ‘non-point’ sources (Box 1), which are hard to tackle technologically. Measures to reduce diffuse sources of nutrient loading are: (i) the reduction of fertilizer application; (ii) the use of nitrogen-fixing crops; and (iii) the restoration or creation (hydrological connections to) of wetlands in the landscape [1]. Here, we critically evaluate the practice of using wetlands to manage water quality in catchments dominated by agriculture. We first consider this practice from a functional perspective and evaluate under what conditions of loading intensity and relative wetland surface area in the catchment wetlands are effective. We pay special attention to undesired effects such as enhanced greenhouse gas production and breakthrough of nutrients stored earlier in the system as a result of overloading. Then, we evaluate the ecological consequences of nutrient loading on the species composition and structure of wetlands. This issue has been ignored in most catchment water-quality enhancement initiatives, where wetlands are considered simply as systems with a high potential for nutrient retention. We review evidence that prolonged Corresponding author: Verhoeven, J.T.A. ([email protected]). Available online 15 December 2005 nutrient loading can have profound effects on the nutrient dynamics of the wetland, leading to shifts from one stable state to the next, often involving structural changes in the vegetation and losses of plant species diversity. We discuss the functionality and biodiversity of wetlands from the perspective of sustainable land and water resource-use management in agricultural catchments to demonstrate that certain conditions must be fulfilled for wetlands to have a substantial effect on water quality without losing their ecological integrity or providing additional risks for global warming. The water purification function of wetlands Wetlands are in use worldwide to reduce concentrations of nutrients in through-flowing water. Many studies at the site scale have demonstrated that wetlands have a high and long-term capacity to improve water quality and this evidence has resulted in many initiatives to restore or even create wetlands for this particular purpose. The most ‘human-controlled’ examples are the so-called ‘treatment wetlands’, which are constructed, planted and hydraulically controlled for the purpose of removing pollutants from wastewater [2,3]. Apart from these constructed wetlands, (semi)natural wetlands in landscapes, such as riparian (stream-side) wetlands, also reduce the nutrient load of through-flowing water [4–8] by removing nitrate and phosphorus from surface and subsurface runoff. With water quality in streams severely deteriorating in densely populated areas and intensively farmed regions worldwide, the interest from natural resource managers in the purifying functionality of wetlands in river catchments is increasing. In Europe, future water-quality standards will become stricter as a result of the implementation of the EU Water Framework Directive (http://europa.eu.int/comm/ environment/water/water-framework/index_en.html). In Canada and the USA, there is also pressure to tighten water-quality standards. Manuals to restore riparian wetlands have been produced (e.g. http://www.chesapeakebay.net/info/forestbuff.cfm) and there have been studies to determine the most optimal location or spatial arrangement of wetlands in agricultural catchments [9]. The use of riparian zones and other types of wetlands for water-quality enhancement has been advocated as a mitigation procedure for ever-more intensive land-use practices, involving denser livestock rearing and increased fertilizer application www.sciencedirect.com 0169-5347/$ - see front matter Q 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.tree.2005.11.015 Review TRENDS in Ecology and Evolution Vol.21 No.2 February 2006 97 Box 1. Diffuse nutrient sources in agricultural areas Figure I indicates the areas of the world that are intensively farmed. In North America, most of the Mississippi basin is used for intensive agriculture, and most parts of Europe, except for latitudes above 558 N, are also used intensively for agriculture, resulting in nutrient loading in the basins of the major river catchments all across the continent (e.g. the Rhine and Danube). In Asia, the major areas with intensive agriculture are southeast China, India and Indonesia, involving the catchments of major rivers such as the Yellow River, the Yangtze and the Ganges. In South America, large parts of the eastern Amazon basin are cultivated, as well as a large area draining towards the Pantanal in Brazil and Argentina. In Africa, a system of subsistence agriculture is common, which is less intense and often does not apply artificial fertilizer. This uneven distribution of agriculture is reflected in the location of areas with diffuse nutrient loading and eutrophication. A study of nutrient loads in some of the major rivers of the world has shown that the European rivers and the Mississippi have increased their nutrient loads ten- to 20-fold from 1960 to 1990 [1]. Severe eutrophication problems involving blooms of cyanobacteria and fish kills have occurred in shallow and deep freshwater lakes [67], in the southern part of the North Sea [68] and in the coastal areas of the Baltic [64] and the Mediterranean [69]. Repeated, prolonged hypoxia has occurred in parts of the Gulf of Mexico, with major fish kills and adverse effects on fisheries [70]. Severe eutrophication problems involving algal blooms, loss of seagrass beds and major damage to blue crab production have also occurred along the east coast of North America, particularly in the Chesapeake Bay estuary [71]. More recently, increasing fertilizer use and diffuse pollution problems have been reported for southeast China [72]. Expectations are that, with the current rapid economic development, agricultural practices will become much less sustainable in this area and will involve severe fertilizer overuse, with detrimental effects on freshwater systems and coastal waters. Here, we focus on the three areas of the world where diffuse nutrient pollution is most severe and where the need for finding solutions including wetland nutrient sinks in the landscape is most pressing (i.e. Europe, eastern North America and southeast China). That does not imply that eutrophication is insignificant in other areas. For instance, the Amazon is the river carrying the highest load of nutrients globally, but its load has, as a result of agriculture, merely doubled [1]. In Africa, there are eutrophication problems in Lake Victoria [73] and Lake Malawi [74] as a result of diffuse nutrient inputs. Figure I. Cultivated areas of the world. Brown regions indicate areas in which at least 30% of the landscape is cultivated. Reproduced with permission from the Millennium Ecosystem Assessment 2005 (http://www.MAweb.org), UNEP. and manure dumping. In many cases, the conservation or restoration of habitat for plant and animal species is seen as an additional benefit of such initiatives [10]. Using wetlands to improve water quality: how does it work? River catchments in which the dominant land-use type is agricultural often have lower-order stream subcatchments that are strongly influenced by runoff from fields or grasslands. In intensively farmed areas, nutrient loading is often so high that large quantities of nitrate leach into the groundwater, which discharges into streams www.sciencedirect.com as seepage or subsurface runoff (Box 2). In intensively farmed catchments, phosphorus (P) and nitrogen (N) are also transported to streams in surface runoff [11]. Between agricultural fields and streams, one often finds riparian areas that can influence surface and subsurface runoff before it reaches streams. There is a large body of literature based on studies at individual sites that indicates that riparian habitats remove nutrients from the water flowing through them on its way from the agricultural land to the stream. The most frequently documented function is the removal of nitrate from subsurface run-off in wetland zones with Review 98 TRENDS in Ecology and Evolution Vol.21 No.2 February 2006 Box 2. Nutrient loading in an agricultural landscape: pathways, processes and effects Heavily fertilized agricultural fields and grasslands show substantial leaching of the very mobile nitrogen species NOK 3 . These nitrates leach downward to the groundwater and are transported laterally to streams, mainly through subsurface runoff and deeper groundwater fluxes (Figure I). Simultaneously, phosphates are transported from the agricultural field in particulate form through overland flow (surface runoff). In the absence of riparian buffer zones in the landscape, these high nutrient loadings flow into the stream water and cause severe eutrophication of the streams, rivers, lakes and coastal areas downstream. In landscapes with riparian zones, the superficial groundwater layer, which is rich in nitrates, comes into contact with the wetland soil, which is highly organic and covered by wetland vegetation. Nitrogen and phosphorus are taken up by the wetland vegetation and are cycled in the ecosystem, which increases their residence time. Thus, that peaks in their concentrations are levelled out before the water enters the stream. If the vegetation is herbaceous, nutrients can be exported although harvesting [17]. Nitrates entering anaerobic soil zones are easily subject to denitrification. The nitrate is first reduced to N2O, and then further to dinitrogen gas. The denitrification process removes nitrogen permanently from the through-flowing water and contributes highly to the sink function of the wetland. If the bacterial process is in some way hampered, the major end product might be N2O, which is a strong greenhouse gas. This might occur when buffer zones are overloaded with nitrates. Greenhouse gas emission Riparian buffer zone Agricultural field off Water table Denitrification Leaching NO3– NO3– (Su b)s PO4 urf ac er un Plant uptake N2, N2O Eutrophication Lakes, ocean Stream NO3– PO4 Water saturated zone Confined layer Loss of biodiversity TRENDS in Ecology & Evolution Figure I. Cross-section of a riparian wetland showing hydrological fluxes, nutrient processes and environmental impacts of nutrient loading. Thicker arrows with warmer colours indicate a higher nutrient loading rate. anaerobic soil conditions [12–17]. Denitrification is generally the most important process for nitrate removal, whereby dead organic matter is decomposed by bacteria in the absence of oxygen, using nitrate as an electron acceptor. Nitrate is converted to nitrous oxide (N2O) and, subsequently, to atmospheric nitrogen (N2), which is emitted by the wetland [18–23]. Nutrient uptake in vegetation as water passes through the riparian zone is also important and results in long-term nitrogen storage [16,24–26]. However, its removal from the system only occurs if the vegetation is harvested as part of the management of that system [18]. Phosphorous removal in riparian habitats has also been reported, with sedimentation, soil adsorption and plant uptake being the most important mechanisms [6,27,28]. www.sciencedirect.com Research on the nutrient removal capacity of wetlands in the temperate zone has revealed that the maximum potential rate of nitrogen and phosphorous removal typically ranges from 1000 to 3000 kg N haK1 yK1 and from 60 to 100 kg P haK1 yK1 [2,20,29]. These are high values, if one considers that they are an order of magnitude higher than fertilizer applications in intensively farmed areas. The capacity of riparian wetlands to remove nitrogen and phosphorous is, however, only important if it can be demonstrated that there has been a significant reduction of the load of nitrogen and phosphorous that reaches stream ecosystems. What is a ‘significant reduction’? In some cases, it has been argued that riparian wetlands contribute in a ‘significant’ way if they remove at least 30% of the total Review TRENDS in Ecology and Evolution Sw ed en Se a Söderköpingsån (882 km2) Ba l ti c Rönneå (1900 km2) (b) N 0 5 99 the USA (http://www.epa.gov/seahome/wqs.html), and have requirements in terms of the presence of indicator species rather than strict limits for nitrogen and phosphorous concentrations in Europe (‘ecological quality’, EU Water Framework Directive). For practical reasons, therefore, we use the first approach by adopting a standard of 30% removal of nitrogen and phosphorous loading as the boundary between ‘significant’ and ‘insignificant’ reductions. (a) Genevadsån (224 km2) Vol.21 No.2 February 2006 10 km TRENDS in Ecology & Evolution Figure 1. Agricultural catchments in Sweden (a) Location of the catchments Söderköpingsån, Genevadsån and Rönneå. These catchments have heavy agricultural use and cause substantial nitrogen enrichment of the Baltic Sea. (b) The 148 riparian wetlands under construction (black dots) in the Rönneå catchment. nitrogen and phosphorous load [30,31]. Another approach is to investigate whether nitrogen and phosphorous removal by riparian wetlands contributes to meeting the water-quality standards in the receiving surface water. These standards are, however, often pragmatic compromises and subject to change. For example, water-quality standards differ regionally in How much land area is required? Studies evaluating the removal of nitrogen and phosphorous by wetlands at the catchment level have been carried out at different scales. For the whole of the Mississippi basin, Mitsch et al. [32] calculated that 20–50% of the total nitrogen load carried by the river into the Gulf of Mexico could be removed by restoring a major proportion of all riparian zones and wetlands associated with the small, lower-order streams, together covering 1–2% of the total catchment area. An additional 20–50% of this total nitrogen load to the Gulf of Mexico could be removed if bottomland hardwood forests associated with the river floodplains were restored to the point that they would cover 3–7% of the total Mississippi basin. Similar estimates were made in catchments in southern Sweden (Figure 1; Table 1), which are a major source of nitrate enrichment of the Baltic Sea. In the Rönneå catchment, restoration of 148 wetlands covering 0.6% of the catchment area failed to improve river nitrate concentrations [33]. A modeling study revealed that 40% nitrogen removal would require a wetland area covering 5% of the total catchment [34]. The traditional Chinese ‘multipond systems’ (Box 3) indicate that effective nitrogen and phosphorous retention occurs when the water in a catchment is directed through a system of ditches and created wetlands covering around 5% of the catchment area, preventing high nutrient loading of the streams and rivers and associated eutrophication problems [35–37]. These examples from the USA, Sweden and China all suggest a global rule that wetlands can contribute significantly to water-quality improvement at the catchment level if they account for at least 2–7% of the catchment area. Effects of nutrient loading on wetland ecosystems Given that many wetlands in agricultural catchments are continually enriched with nutrients, it is relevant to consider the consequences for the species composition and functioning of the wetland ecosystem. Enriched wetland systems can lose species and can show drastic changes in nutrient cycling rates. Table 1. Modelled nitrogen removal by wetland constructions on arable land in three catchments in Swedena,b Catchment Söderköpingsån Genevadsån Rönneå a Catchment size (km2) 882 224 1900 Present situation (mt N yK1) 234 267 1990 With potential wetlands (mt N yK1) 132 251 1753 Removal by wetlands (mt N yK1) 102 16 137 (%) 43% 6% 7% Wetland area (% of total) 5.0 0.4 0.6 See also Figure 1, main text. These studies imply that 5% of the total catchment must be considered for wetland restoration to have a significant impact on nitrogen reduction. b www.sciencedirect.com Refs [65] [33] [34,66] 100 Review TRENDS in Ecology and Evolution Vol.21 No.2 February 2006 Box 3. Multipond systems in China: ancient water management in agricultural catchments Multipond systems, which are large artificial irrigation networks dating back 20 centuries, are common in southeast China. They consist of many tiny ponds, scattered in agricultural catchments and connected by ditches and streams [36,37]. Typically, the pond area is 6–10% of the total catchment and each pond has an area of 0.05– 1 ha, serving an area of 0.5–10 ha. The ponds collect surface runoff and agricultural drainage water, which is used to irrigate rice and other crops. Each pond is connected via ditches to three–five other ponds and the water cascades from one pond to the next. The multipond systems have a large retention capacity for nutrients. In the Liuchahe Watershed (Figure I), there is a high variation in nutrient concentrations exported from different land-use types. Runoff nutrient concentrations are highest in water discharging from villages, croplands and rice paddies. After capture of the runoff in the ditches and ponds, the nutrient concentrations were reduced significantly and nutrient concentrations at the watershed outlet were consistently lower than in the runoff water [35]. The retention and removal mechanisms of nutrients in the multipond systems include sedimentation, adsorption, recycling through irrigation and uptake by aquatic plants. Nutrient budgets compiled for phosphorus during five years in Liuchahe catchment showed reductions of nutrients in runoff by the multipond systems of O90% [35]. In some regions of China in recent years, there has been a tendency to ignore the importance of the multipond systems owing to the development of modern reservoirs and irrigation channels. The number of ponds throughout the country was 8.3 million during the 1950s, and the irrigation area was 133 million ha, which accounted for 39% of whole irrigation area. The number of ponds is now only 6.3 million, and the corresponding irrigation area is 80 million ha [75]. Some ponds were destroyed to increase arable land and housing, whereas others became filled with sediments that were no longer excavated to fertilize arable land. The distribution area has been reduced significantly and this is already having negative impacts upon water quality in the rural areas of the country. (a) (b) N 0 1000 km Liuchahe river Figure I. ‘Multipond’ landscape in southern China. (a) Typical landscape showing ponds, rice fields, woodlands and villages near Chaohu Lake, P.R China. (b) Water system with connected ponds (in blue) and ditches (blue lines) in the Liuchahe catchment area at the same location. Critical loads Nutrient inputs to ecosystems have increased over the past century in many parts of the world. The resulting nutrient enrichment often has significant effects, including increased productivity, higher rates of nutrient leaching and shifts in the dominance, and composition, of species [38–40]. Most ecosystems can incorporate higher loading rates, which have only minor effects as long as they do not surpass a certain limit. However, when nutrient loading rates surpass this critical level, species composition and ecosystem functioning change dramatically over short periods of time and the systems often move to a different stable state. Among the best-known examples are shallow lakes that shift from water with low turbidity and abundant submerged macrophyte vegetation to a turbid state with prolonged phytoplankton dominance [41]. Wetlands that are characterized by low productivity and high plant diversity dominated by slowgrowing, nutrient-conserving species shift to systems dominated by large, fast-growing helophytes following a strong increase in nutrient-loading rates. The degree to which the species composition changes depends on the natural nutrient richness of the system. www.sciencedirect.com Naturally nutrient-poor (i.e. oligotrophic and mesotrophic) systems react more drastically than do naturally nutrient-rich (eutrophic) systems. Nutrient-poor systems show a complete shift in plant species composition as well as a drastic change in nutrient dynamics, whereas nutrient-rich systems might show only further increased productivity. All systems, however, show a characteristic breakdown of the nutrient retention function after prolonged high nutrient loading [42]. An example of a drastic shift in plant species composition and nutrient dynamics is the effect of agricultural runoff on a large mesotrophic wetland system in the Everglades. Species-rich communities dominated by sawgrass Cladium jamaicense have been replaced by tall species-poor cattail Typha domingensis stands in areas enriched with nutrients. In the same areas, the rate of phosphorus cycling has increased as a result of higher decomposition rates [38,43]. This reinforces the nutrient richness of the system and creates a situation that is difficult to reverse to mesotrophic conditions. Such shifts in structure (i.e. species composition) and functioning (nutrient cycling and retention) have spurred scientists to propose critical loads of nitrogen and Review TRENDS in Ecology and Evolution phosphorous for ecosystems. A critical load of a nutrient is defined as the loading rate below which the system remains all but unchanged, but beyond which it exhibits sudden, drastic changes, including a shift in species dominance and species composition and a major change in ecosystem functioning, in terms of carbon (C) and nutrient outputs, trophic interactions and/or nutrient cycling rates. Such situations of nutrient overloads cannot only be detected as a major shift in species composition or structure, but also as a distinct increase in the nutrient concentrations of water that is exported from the ecosystem (e.g. in wetland outflow). A drastic increase in output nutrient concentrations is another indicator that can be used to establish critical loads for ecosystems. For wetlands, a critical loading rate of 10 kg P haK1 yK1 has been proposed, based on the analysis of a large database of wetlands enriched with nutrients [44,45]. For nitrogen, much research has been carried out along gradients of atmospheric nitrogen deposition in Europe [46–48]. Increased levels of atmospheric nitrogen deposition occur as a result of air pollution owing to fossil fuel combustion (NOx) and agriculture (NH3). In Western Europe during the 1980s, levels were higher than 45 kg N haK1 yK1, which is ten times the background value [48]. Metadata sets on the effects of increased deposition have proposed a critical nitrogen load of w25 kg N haK1 yK1 for wetlands, but there is little published information available about differences among different wetland types in this respect [46–48]. It is striking that the proposed critical loading rates of nitrogen and phosphorous for wetlands are several orders of magnitude lower than the typical loading rates in natural or constructed wetlands used for improving water quality (Table 2). Only in a few cases, for example in wetlands used for water treatment in the Mississippi [49], loading rates of nitrogen and phosphorous have been close to these critical values. This implies that critical nutrient loads are easily surpassed in many natural wetlands and that, depending on their original trophic status, shifts in species composition and/or increases in nutrient concentrations in the outflow will occur, in spite of rates of nutrient retention remaining high. As an example, forested riparian zones in the Netherlands typically receive nitrate-rich runoff from heavily fertilized fields [50,51]. Many of them lost their original plant diversity decades ago [52], but continue to retain high quantities of nitrate, even though the surface and subsurface water discharged from them still have high concentrations of Vol.21 No.2 February 2006 101 nitrate [13]. Hence, wetlands in agricultural catchments can contribute significantly to water-quality improvement, but their loading rates often surpass critical values. Enhanced nitrous oxide emissions The major process responsible for nitrate removal in wetlands is denitrification. However, in situations where the reduction of the nitrate to N2 is incomplete, the denitrification process can also be a major source of the greenhouse gas N2O, which has a global warming potential 310 times that of CO2 [53]. N2O accounts for w6% of the total greenhouse effect and also has an important role in the destruction of the stratospheric ozone. N2O emission in wetlands is generally promoted under conditions that are suboptimal for denitrification, such as low pH or soil moisture. However, N2O production is also promoted by high nitrate availability, because it is energetically favorable for denitrifyers to reduce nitrate instead of N2O. There is much recent information about the increase in N2O emission after nitrogen addition to agricultural soils [54–56]. Much less data exist on the indirect effect of nitrogen addition by agriculture on N2O emissions from riparian wetlands [57–61]. Recently, Barnard et al. [62] reviewed studies of nitrogen addition to natural ecosystems and concluded that nitrogen addition significantly increased N2O emissions in both laboratory and field experiments. Recent publications [57–60,62], strongly suggest that nitrogen transformations in buffer zones receiving high levels of nitrate result in a significant increase in N2O emissions. For example, riparian buffer zones in Twente, the Netherlands, showed high N2 O emissions (20 kg N haK1 yK1) at sites with high nitrate loads [63]. This implies that the nutrient retention benefit of riparian zones comes at an environmental cost, in particular when nitrate-loading rates are also high. This potential negative consequence of loading wetlands with nitrate is often ignored or downplayed. Thus, there is a great need for additional information about the risk of N2O emissions from nitrogen-loaded wetlands and about management options to balance that risk against the environmental benefit of water-quality improvement. Wetlands in the landscape for water-quality improvement: finding the right balance Wetlands in nutrient-loaded agricultural catchments have a significant role in improving water quality. Our analysis Table 2. Characteristic nitrogen and phosphorous loading rates of wetlands in agricultural catchments in relation to relevant loading thresholds Catchment Liuchahe Regge, Twente Everglades Mississippi Various Treatment wetlands Maximum loadb Critical loadc a Location Wetland type P.R. China Multipond Netherlands Riparian USA Marsh USA Forested USA Riparian in USA and Europe Origin Constructed Natural Natural Natural Natural Constructed N load (kg haK1 yK1) O500 200–1140 N/Aa 19–39 20–155 500–9000 1000 25 P load (kg haK1 yK1) O50 2–40 2–9 N/A 100–2000 60 10 No data available. Beyond this limit, the wetland will show substantial leaching and associated high concentrations in the outflow. c Beyond this limit, wetlands with a species-rich vegetation will show a dramatic increase of productivity and associated change in species composition. b www.sciencedirect.com Refs [35–37] [13,63] [38,43] [49] [32,49] [2,32] [2,20,32] [44,45,47,48] 102 Review TRENDS in Ecology and Evolution shows that this potential is only realized in catchments with a minimum area of wetlands relative to total catchment size. Measurements from different regions around the world indicate that at least 2–7% of the total catchment needs to be in wetland habitat to see a significant increase of water quality at the catchment scale, a remarkably narrow range. This minimum value has already raised much dispute among policy makers as to the practicality of restoring or creating such a large area of wetlands as a management tool, particularly in many European catchments, where the proportion of wetlands has often become close to zero. In Sweden, resource managers advocate other measures such as less drainage of fields and grasslands, or lower fertilizer applications as less costly and easier to realize than wetland restoration [34,64]. In areas with intensive land use and very heavy use of fertilizer, such as the Netherlands, nitrate concentrations in the groundwater are so high that high N2O emissions in the riparian wetlands are to be expected. Again, restoration of riparian zones should be accompanied by lower fertilizer applications to avoid adverse effects on the environment. The implications for land-use management in areas under intensive agricultural use (e.g. Europe, South Asia or the Mississippi basin) are that only a combination of measures will result in acceptable environmental quality, that is, (i) fertilizer levels have to be reduced significantly; (ii) riparian zones and floodplain wetlands should be rehabilitated; and (iii) their location in the landscape should be carefully selected on the basis of hydrological studies. This will only be possible at great economic cost, which will delay implementation. Another clear outcome of our discussion is that many wetlands in agricultural catchments are loaded beyond the ‘critical’ limits, and some even beyond the ‘maximum’ limits (Table 2). This means that many riparian zones will lose plant species or have already done so. Riparian wetlands in such catchments will all converge towards a narrow set of ecosystem types characterized by high levels of nutrient richness, a high primary productivity and low species density. The example of wetlands used for water treatment in the Mississippi Delta [52] shows that the nutrient retention function can be combined successfully with conservation of the original status of the ecosystem and its biodiversity under the condition that loading rates are low and not surpassing critical limits. Conclusions Land-use planners and environmental resource managers are facing many challenges to ensure good surface-water quality in agricultural catchments. In their development of sustainable water-quality management options, they must give consideration to the existence of loading limits as discussed here. Because of their high potential for nutrient retention, it is still a good idea to use wetlands in catchment water resources management for water-quality improvement. Restoring wetlands and their hydrological connections to the upland and the stream can be a rewarding activity in this respect. Most importantly, however, the combination of waterquality improvement with wetland biodiversity www.sciencedirect.com Vol.21 No.2 February 2006 conservation requires loading rates of wetlands to remain below critical thresholds. In many agricultural catchments, these limits have been surpassed and biodiversity restoration would require a decrease in the loading through decreased fertilizer applications or the use of larger areas of wetlands for nutrient retention. In catchments with intensive land use, loading rates could surpass another, much higher critical limit, beyond which the wetland ecosystem no longer performs its retention function properly but releases nutrients or emits the greenhouse gas N2O. In such catchments, the only feasible measure is to decrease fertilizer levels. 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