N2o

reduced methane oxidation —2 tree loss fewer hikers reduced water retention higher discharge peak increased moisture increased temperature

abiotic routing

-1 N20 physical production degassing of N20

-1 less fishing aesthetic value declines physical — 1 degassing of N20

methano-genesis disease control limits invasive species migration up estuary increased freshwater decreased salinity increased fine sediment may fill interstices within stream substratum, decreasing hyporheic exchange (Morrice et al. 1997); this may reduce key biotic processes such as denitrification. However, increased fine inorganic sediment may increase uptake and storage of ions such as phosphate, which will bind to increased fine sediment (Meyer 1979). Increased sediment may reduce numbers of filter-feeding insects, which may lower downstream transport of particle carbon. In some cases, however, the reduction of filter feeders would not have as serious effects on transport of particle carbon as would the reduction of estuarine bivalves (Wallace & Merritt 1980; Caraco et al. 1997; Jassby et al. 2002; but see also Crowl et al. 2001).

The net effect of rivers and associated riparian zones is to lower the dissolved nutrient concentrations that are exported to estuaries (Alexander et al. 2000), with deforestation having to occur over a large portion of the watershed in order to substantially reverse this process. Given that nutrient concentrations decrease following regrowth of vegetation, and that large watersheds are cut sequentially, it is unlikely that increases in nutrients would rival agricultural or atmospheric inputs. Similarly, much of the sediment transported to streams may be stored in stream channels, thereby attenuating the delivery of sediment to downstream ecosystems. It is possible that temporary losses of organic matter from allochthonous (non-local) inputs could lower productivity in downstream ecosystems if there is substantial transport of this material (e.g., Vannote et al. 1980). It is more likely that loss of instream retention structures (e.g., woody debris) will decrease the ability of the stream to retain organic matter, with streams from logged catchments potentially having higher rates of organic matter turnover and higher export than non-logged catchments, at the expense of benthic storage (Webster et al. 1990). Despite small-scale catchment studies of organic matter export, there are few tests of large-scale impacts of land use on organic imports in downstream ecosystems. The provision of ecosystem goods and services within the freshwater domain have been discussed throughout this volume, and a brief summary of the impacts of deforestation within the terrestrial domain is provided below.

1. Food provision. The quantity and quality of fresh waters have particularly marked effects on salmon production, with decreased salmon spawning due primarily to increased sediment input to streams associated with felling. Whirling disease may exacerbate problems. Other biological changes are exerted via the shredders, which decline with the increased algal blooms resulting from water-quality changes. Water-quality changes will increase the numbers of deposit feeders, and abiotic factors, such as increasing temperatures, may also be affected.

2. Carbon sequestration. Streams are not considered to be primary sites for C sequestration because of their relatively small area and low speed of flow. However, after felling it is anticipated that the net balance between increased runoff and increased sediment load will lead to a net increase in river sediment.

3. Water quantity. For the reasons presented in the terrestrial domain section (above), the quantity of stream water will increase post-felling. This increased yield may be significant, and it is an important consideration when forested catchments are being managed largely for water provision.

4. Water quality. Most countries have recommended guidelines or legislation for forest-felling operations in areas where water from a catchment is important for human consumption. Specific limitations on stream-edge felling, road construction, pesticide usage, and so on are designed to protect water quality, with concentrations of NO3, Al, dissolved organic matter, and sediment being critical indexes of water quality. The impairment of this important ecosystem service can impact different sectors depending on the chemical changes that occur. For example, high concentrations of nitrate are considered a risk to human health, while aluminum may be toxic to fish and other stream biota; high DOC and associated dark water color may give rise to additional processing or mixing costs.

5. Trace gas production. During transfer in the stream, some nitrogen will be removed from the water course by algae and also by the process of denitrification, resulting in the production of N2O. Trace gases, including N2O and CO2 previously produced in the soil and dissolved in steam water, will be de-gassed from the stream into the atmosphere.

6. Recreation and transport. Deforestation is frequently associated with new roads, which may increase recreational access for fishing, canoeing, and so on. Major rivers are actually used to transport timber from logging sites. However, increased sediment loads and Al3+ may cause reductions in specific fish stock.

An overview of changes of ecosystem services and consequences for outputs toward freshwater systems is provided in Table 9.1 section B.

Marine Ecosystems: Deforestation Impacts

Key inputs into the coastal zone (estuaries and shelf) from upstream will include sediments, nutrients, pollutants, and, to a lesser extent, fresh water. They will have both positive and negative effects on the services provided by sediment habitats (salt marsh, mangrove, seagrass beds, and tidal flats) and by sediment-dwelling marine benthos. These services are discussed in more detail in Chapters 4 and 7, and are also reviewed by Snelgrove et al. (1997, 2000) and Levin et al. (2001). Briefly, they include ecological functions such as decomposition and nutrient regeneration; regulation of water fluxes, particles, and organisms; and habitat and nursery provision. They also provide more direct services: shoreline stabilization; water purification; the production of har-vestable fish, shellfish, and plants; and trophic support for these fisheries. Key functional groups involved in providing these services include bacteria and fungi, shredders, suspension feeders, and bioturbators (see Wall et al. 2001b). Below we review a range of scenarios regarding altered sediment function in the coastal zone that might result from increased inputs of nutrients in sediments of freshwater associated with upland deforestation.

The life histories and health of many estuarine organisms are finely tuned to the quantity and timing of freshwater input to estuaries, and to the resulting salinity variations. Attrill (2002) has demonstrated that estuarine diversity in the Thames River Estuary is lowest where the variation is greatest (often in the zone where salinity averages 5 — 15 ppt). Drought usually increases estuarine salinity and may reduce habitabil-ity for upper estuary oligohaline organisms. Invasions by marine species and increased incidence of disease have been attributed to reduced freshwater inputs associated with drought, as the timing of reproduction or growth is finely tuned to winter or spring flooding. However, increases in freshwater input associated with inland deforestation are likely to be minor and these effects will be minimal in comparison with those of sediment and nutrient input.

Increased nutrient inputs to the estuary from freshwater sources (rivers, streams, and groundwater) may impair or enhance key ecosystem services, depending on the level of nutrient loading. Because nitrogen availability often limits primary production by phy-toplankton, algae, and rooted vegetation, low-level increases in nitrate concentration could increase primary production. The sediment biota may respond with increased abundance of all functional groups, leading to greater trophic support for, and production of, fish and shellfish. Increased nutrient availability may yield enhanced growth and peat deposition by vascular plants (salt marsh plants, mangroves, and sea grasses). This would increase C sequestration, stabilize sediments, and improve water quality through slowed flows and particle removal. The spread of wetlands may also lead to elevated organic matter accumulation, greater microbial activity, and greater production of methane, a potent greenhouse gas. However, concurrent increase in turbidity may negate the positive effects of nutrients on primary production. High nutrient loading, in contrast, will lead to levels of phytoplankton production that quickly overload the system. Decay of unconsumed primary production will increase biological oxygen demand and generate hypoxic waters, a process termed eutrophication. A sequence of changes in the sediment biota may be expected, with large epibiota (e.g., crabs, shrimp, bottom fish) and subsurface dwelling forms (bivalves, echiurans, annelids) declining first, with loss of bioturbation and nutrient regeneration capacity. Small, opportunistic taxa that tolerate hypoxia construct tubes or shallow burrows in dense assemblages near the sediment-water interface (Pearson & Rosenberg 1978), and they have limited influence on the cycling or burial of organic matter (Rhoads & Boyer 1982). Diversity of species and functional groups will decline with increasing hypoxia. With severe eutrophication, the sea floor may be carpeted with dead plankton, fish, and other organisms. These will become covered by sulfur bacteria, and underlying sediments will be anoxic and azoic. Such conditions have been noted in the Baltic and North Sea in the past, and occur in summer in the coastal Gulf of Mexico (Rabalais et al. 2001). It is unlikely, however, that the release of nutrients and organic debris by deforestation alone would be of a magnitude sufficient to induce such severe eutrophication in coastal estuaries. Typically, nitrogen fertilizer and phosphate runoff from domestic and industrial use are contributing factors.

Steep rivers that cut through mountainous regions may deposit large amounts of terrestrial debris and fine organic matter on narrow continental shelves and slopes during flood events (Leithold & Hope 1999; Wheatcroft 2000). On the Eel River margin of northern California, deposits of woody debris coincide with the occurrence of methane seepage. On the slope at 500 m, distinctive infaunal assemblages are formed (Levin et al. 2003) that have carbon isotopic signatures consistent with a carbon source derived from terrestrial organic matter (Levin & Michener 2002). It is possible that deforestation effects on organic inputs could be felt on the shelf and slope where steep mountainous terrain abuts the coastal zone.

Suspended Sediment

Total suspended particulate matter (TSM) contains both organic particulate organic matter (POM) and inorganic particles transported with riverine systems to estuaries and the sea. Its content varies seasonally and geographically, ranging from 10 to 1000 mg TSM per cubic decimeter. As they reach the marine zone, suspended sediments undergo rapid transformation and are largely removed from the water column in the process of physicochemical flocculation, aggregation, and finally sedimentation. In open river mouths discharging the freshwaters directly to the full saline sea, as much as 95 percent of POM is sedimented in the immediate vicinity of river mouths, and only a small fraction of the original suspended load is transported to the shelf (Lisitzin 1999). Sedimentation rates in river mouths are extremely variable, but may range over 1500 g/m2/day; similar values have been reported from vegetated salt wetlands and tidal flats. TSM loads may enter the estuaries both as bedload transported and surface-layer transported particles, depending on the hydrology, season, speed of the flow, and so on. In the first instance, the surface estuarine waters are relatively transparent (euphotic) and local primary production may be high. In other cases, the water transparency drops and local primary production is completely inhibited. The increase of suspended load and consequent increase of sedimentation have profound effects on the estuarine sedimentary biota.

The overall reaction of benthic assemblages follows the pattern described by Pearson and Rosenberg (1978) in disturbed or enriched marine sediments. Large, sessile, structure-forming species disappear and are replaced by small, mobile, surface-dwelling taxa. The increase of TSM load influences the services provided by estuarine-shelf biota. Effects include a drop in biomass of harvestable species or enhanced carbon sequestration by bacterial development on mass aggregates of organic matter. Other services, such as recreation and aesthetic values, are likely to be affected negatively due to the turbid water, decrease of fish production, and organic matter deposits.

The effects of terrestrial sediment deposition on New Zealand estuaries, as mentioned earlier, have been particularly well studied and are considered here in more detail. Extensive deforestation and other development-related activities have led to sediment inputs via upland landslides, sediment runoff, and river-carried material in New Zealand. Historical sediment deposition records indicate that New Zealand experienced massively increased sediment input to the rivers and estuaries with the arrival of humans on the island, and again with the arrival of Europeans, who were responsible for much of the deforestation (Hume & McGlone 1986). The most deadly sediment inputs occur in the form of clay-dominated subsoils, and flooding and landslides may result in layers of this material millimeters to centimeters thick being deposited into estuaries; single deposition events of up to 10 cm have been recorded (Thrush 2004). The clays are associated with low pH and often with refractory organic matter. Few animals are able to burrow through clay layers, and sediments beneath these layers become highly anoxic. Experimental deposition of clay-dominated sediments reveal significant negative impacts and slow rates of macrobenthic recovery (Norkko et al. 2002; Cummings et al. 2003; Hewitt et al. 2003; Thrush et al. in press). Even more chronic deposition events (1, 3, 5, 7 mm thick) can produce changes in the density of macrofaunal density of up to 50 percent (Thrush 2004). These effects include bivalve and polychaete mortality, as well as changes in feeding behavior, declines in growth density, and slowed recruitment, with effects lasting up to 18 months or more (Norkko et al. 2002). Only large burrowing crustaceans (crabs and shrimp) are able to cope with the high sediment loads. Microphytobenthos (benthic microalgae) that can migrate up through the layers of clay will survive, but those that cannot will die, diminishing the food supply for many grazers in sediments. Experiments reveal that species-level identification is necessary to evaluate consequences of sediment input for estuarine sediment communities. Within a single functional group (Spi-onidae, a family of surface-feeding polychaetes), sensitivity to clay deposition may differ greatly among species (Thrush et al. 2003, in press).

In New Zealand, sediment fecal bacteria often accompany sediment runoff from agricultural land. Together the sediments and bacteria have negative impacts on harvests of filter-feeding oysters (Crassostreagigas) and mussels (Perna canaliculus; Hawkins et al. 1999) from marine farms as well as affecting recreational and traditional shellfish harvesting by the Maori population. Increased sedimentation and associated nutrient input may also account for expansion of mangroves in many of the harbors and estuaries in northern New Zealand (Dingwall 1984).

Below, in summary, we list some of the key impacts of deforestation across a range of marine and coastal ecosystems services:

1. Food provision. Water quality, including inorganic and organic solutes and, particularly in this case, sediment load from rivers, can have major effects on food provision by coastal and estuarine systems. These effects may be both positive and negative, ranging from potentially increasing the build-up of fish nursery grounds through to declines in the number of filter-feeding bivalves. As outlined above, the sediment input via fresh water entering marine systems has particularly marked effects on salmon production.

2. Carbon sequestration. The continental margins play a major, but poorly quantified, role as burial sites of organic carbon derived from both oceanic and terrestrial sources. Perturbations in freshwater discharge alter the intensity and depth of penetration of thermohaline circulation, with consequences for the fate of carbon, the transfer of dissolved organic carbon and sediment, and the export of dissolved organic carbon and particulate organic matter.

3. Water quantity. For the reasons presented above, the quantity of stream water will increase post-felling.

An overview of changes of ecosystem services and consequences for outputs toward marine systems is provided in Table 9.1 section C.

Discussion and Conclusions

In this analysis we have necessarily restricted our considerations to the cross-domain impacts of one specific disturbance in one ecosystem type; however, some generic conclusions have emerged (Table 9.2). First, it is very clear that significant impacts can be readily passed on through a chain of domains, with important consequences for the downstream systems. For example, the logging of large forest areas can affect remote coastal marine domains, as seen in the clear example from New Zealand discussed above. The impacts on the sediment biota are central to the sensitivity of the system, with attendant consequences for ecosystem services of importance to humans. Again, we have restricted discussion to the felling of temperate forests but similar, and equally dramatic, effects have also been reported for nontemperate systems. For example, impacts of increased sediment loads on sensitive systems such as coral reefs (e.g., Dubinsky & Stambler 1996) are particularly noteworthy because of their large conser-vational, and increasingly recreational, value.

Second, it is also clear that a single type of disturbance may have differing impacts on the different receiving domains, and that these effects may not be simply proportionate to distance from the disturbance. For example, deforestation may lead locally to increased stream nitrate and aluminum concentrations with, respectively, consequences for local water potability and fish stocks. However, by the time these streams have merged with others and then entered the oceans, the effects may have become insignificant. The sensitivity of the organisms and processes in the receiving system, amounts of dilution, and existing load from other sources are all key in controlling the extent of the effects attributable to a specific land management change. It should be emphasized that the case for sediment load is very different from the case for solutes, with the coastal system effectively acting as an accumulation point for sedi-

Table 9.2. Net effects of deforestation on soil, freshwater, and marine domains.

Terrestrial

Freshwater

Marine

Loss of sediment Loss of nutrients Change in soil biota

Loss of carbon

Increased temperature

High sediment load Increased NO3 + other nutrients More water

Increased light and temperature

Switch from autotrophy to heterotrophy

Alteration of sediment biota

More wetland Sediment load Nitrogen stimulation of production Altered benthic production ment, while also containing organisms and processes sensitive to changes in these inputs. In the case of nitrate loading, nutrients in the coastal zone are normally dominated by fertilizer and effluent inputs, which largely dwarf any solute contributions arising from deforestation.

In their evaluation of the spatial appropriation of both marine and terrestrial ecosystem resources for the Baltic Sea drainage basin, Jansson et al. (1999) emphasized the critical interdependence between freshwater flows and the capacity of ecosystems to generate services, emphasizing the need for a holistic approach to watershed landscape management to avoid any unintentional effects and loss of services. We would argue that improved anticipation of unintentional effects across domains can be achieved using the framework described here, with a systematic assessment of likely cross-domain effects. Management options that lead to the lowest impairment of ecosystem services, balanced across all domains, should be selected.

The decision to limit the current evaluation to deforestation and temperate systems, while making the process tractable, has also limited the extent to which any resulting conclusions can be generalized. However, a brief consideration of studies in nontem-perate regions actually lends support to the general conclusions. For example, the role of deforestation in increasing sediment loads and, more specifically, mercury contamination in lacustrine sediments in the central Amazon has been reported by Roulet et al. (2000), indicating a major impact of anthropogenic disturbance on mineral and organic cycling across the entire region. Studies examining the causes for the substantial quantitative and qualitative changes in deposited sedimentary organic matter in major tributaries of the Amazon also strongly implicate increased deforestation as a major cause (Farella et al. 2001), although the consequences are less fully described. As mentioned above, increased sediment load from deforestation and topsoil erosion have been linked to substantial marine changes in specific nontemperate marine systems, including coral death and eutrophication (Dubinsky & Stambler 1996). Downing et al.

(1999) argued that alterations in the N cycle, resulting from forest disturbance in tropical systems, will have much greater impacts on tropical aquatic ecosystems than on temperate equivalents, largely because of the greater importance of nutrient limitations in these systems.

Thus, it appears to be the case that the general conclusions arising from the specific case study will extend to other biome types; however, we have not addressed the question of the extent to which the concepts developed here can be applied to other environmental changes. The evidence from the current evaluation would suggest that changes resulting from perturbing a single domain will frequently be seen as impacts in other domains, and that different domains may respond surprisingly different to the same environmental change—it is clear that one domain's meat may be another domain's poison.

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