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Soft systems methodology and the ecosystem approach: a system study of the Cooum River and environs in Chennai, India.

This paper discusses the integration of soft systems methodology (SSM) within an ecosystem approach in research to support rehabilitation and management of the Cooum River and environs in Chennai, India. The Cooum is an extremely polluted urban stream. Its management is complicated by high rates of population growth, poverty, uncontrolled urban development, jurisdictional conflicts, institutional culture, flat topography, tidal action, blockage of the river mouth, and monsoon flooding. The situation is characterized by basic uncertainty about main processes and activities, and the nature of relationships among actors and elements in the system.SSM is an approach for dealing with messy or ill-structured problematic situations involving human activity. In this work SSM contributed techniques (such as "rich picture" and "CATWOE" tools) to description of the Cooum situation as a socioecological system and informed the approach itself at a theoretical level. Application of three general phases in SSM is discussed in the context of the Cooum River research: (1) problem definition and exploration of the problem situation, (2) development of conceptual models of relevant systems, and (3) the use of these to generate insight and stimulate debate about desirable and feasible change. Its use here gives weight to the statement by others that SSM would be a particularly appropriate methodology to operate the ecosystem approach. As well as informing efforts at management of the Cooum system, this work led the way to explore an adaptive ecosystem approach more broadly to management of the urban environment for human health in Chennai.

Conservation of Natural Resources↗

Legitimizing fluvial ecosystems as users of water: an overview.

We suggest that fluvial ecosystems are legitimate users of water and that there are basic ecological principles guiding the maintenance of long-term ecological vitality. This article articulates some fundamental relationships between physical and ecological processes, presents basic principles for maintaining the vitality of fluvial ecosystems, identifies several major scientific challenges and opportunities for effective implementation of the basic ecological principles, and acts as an introduction to three specific articles to follow on biodiversity, biogeochemistry, and riparian communities. All the objectives, by necessity, link climate, land, and fresh water. The basic principles proposed are: (1) the natural flow regime shapes the evolution of aquatic biota and ecological processes, (2) every river has a characteristic flow regime and an associated biotic community, and (3) aquatic ecosystems are topographically unique in occupying the lowest position in the landscape, thereby integrating catchment-scale processes. Scientific challenges for the immediate future relate to quantifying cumulative effects, linking multidisciplinary knowledge and models, and formulating effective monitoring and assessment procedures. Additionally, forecasting the ecological consequences of changing water regimes is a fundamental challenge for science, especially as environmental issues related to fresh waters escalate in the next two to three decades.

Climate↗

Economic valuation of a mangrove ecosystem threatened by shrimp aquaculture in Sri Lanka.

Mangrove ecosystems in Sri Lanka are increasingly under threat from development projects, especially aquaculture. An economic assessment is presented for a relatively large (42 ha) shrimp culture development proposed for the Rekawa Lagoon system in the south of Sri Lanka, which involved an extended cost-benefit analysis of the proposal and an estimate of the "total economic value" (TEV) of a mangrove ecosystem. The analysis revealed that the internal benefits of developing the shrimp farm are higher than the internal costs in the ratio of 1.5:1. However, when the wider environmental impacts are more comprehensively evaluated, the external benefits are much lower than the external costs in a ratio that ranges between 1:6 and 1:11. In areas like Rekawa, where agriculture and fisheries are widely practiced at subsistence levels, shrimp aquaculture developments have disproportionately large impacts on traditional livelihoods and social welfare. Thus, although the analysis retains considerable uncertainties, more explicit costing of the environmental services provided by mangrove ecosystems demonstrates that low intensity, but sustainable, harvesting has far greater long-term value to local stakeholders and the wider community than large shrimp aquaculture developments.

Animals↗

Effects of fish stocking on ecosystem services: an overview and case study using the Stockholm Archipelago.

In this article, we focus on documented and possible effects of fish stocking in terms of ecosystem services. The increasing use of fish stocking between 1970 and 2000 in the semiurban setting of Stockholm archipelago, Sweden, is used as case study. The objective is to analyze this management practice from an ecosystem perspective, accounting for both the ecological and social context of releasing fish. The results show that enhancements of four native species (Salmo S. trutta, Salmo salar, Stizostedion lucioperca, and Anguilla anguilla) have dominated over new introductions of one nonnative species. (Oncorhynchus mykiss). The major objective has been to increase fish catches for local resource users. Involved stakeholders include three management agencies, one hydropower company, and several local sport fishing associations. Documented effects focus on recapture and production rates. However, our analysis suggests that additional positive or negative effects on biodiversity, food web dynamics, mobile links, or ecological information may also result, with possible consequences for the long-term provision of food, game, and aesthetic values. We conclude that a more adaptive and cooperative management approach could benefit from a deeper analysis of where, when, and what species is released, by whom, which stakeholders that use the fish and those ecosystem services the fish generate, and of the role of formal and informal institutions for monitoring and evaluating the success of releasing fish.

Animal Husbandry↗

Terrestrial and aquatic invertebrates as bioindicators for environmental monitoring, with particular reference to mountain ecosystems.

The use of terrestrial and aquatic invertebrates as a management tool for monitoring change in ecosystems is reviewed and critically evaluated. Their suitability and value for assessing a range of environmental problems from pollution impacts, through habitat evaluation for conservation to the long-term degradation and recovery of ecosystems, is critically discussed. Guidelines are provided for the choice of appropriate bioindicators. Examples of the use of a broad spectrum of invertebrates to assess a variety of environmental problems are summarized. The particular potential of invertebrates for monitoring montane ecosystems is highlighted.

Animals↗

Conceptual design of monitoring and evaluation plans for fish and wildlife in the Columbia River ecosystem.

A logical sequence of seven steps is proposed as a generic template to design plans for monitoring and evaluating fish and wildlife in the Columbia River ecosystem. Management programs for these resources fail to include coordinated monitoring and evaluation plans. This short-coming is indicative of pervasive management conflicts detected from regional to local geographic scales. In the absence of a cohesive ecological management framework, monitoring and evaluation activities proceed without a clear understanding of what uncertainty they are intended to address, nor is there a clear description of the process to utilize the information gained. As a result, the accountability for the investment of public funds for fish and wildlife restoration is poor, information collected from the environment is not included in decision-making, and the ability to gain knowledge while taking management actions is compromised. The sequence of steps discussed here does not identify or describe distinct monitoring activities or methodologies at any particular location or listed under any specific monitoring plan. Instead, it concentrates on the generic elements necessary for the design and implementation of coordinated fish and wildlife monitoring plans. It is proposed that at least four major issues demand considerable attention in order to improve regional monitoring and evaluation capabilities: The first is adoption of an ecological framework for the management of fish and wildlife at relevant geographic scales within the ecosystem. Such a framework must include an explicit identification of goals, objectives, and actions to steer coordinated decisions across the boundaries of technical disciplines, management jurisdictions, and institutional responsibilities. The second is that the identification of these management goals for the geographic location of interest must precede the design of monitoring and evaluation plans from the top down. Third, the evaluation component must be considered early on in the planning process, so that it blends smoothly with monitoring at the time of implementation. Fourth, decision-makers and scientists engaged in the planning of fish and wildlife monitoring and evaluation efforts in the region must have a close collaborative relationship. Monitoring and evaluation plans designed under these premises may enhance our collective observational capabilities, promote cost-effectiveness and adequate evaluation, and provide a useful tool to adjust our management practices to the challenges of complex ecosystems.

Animals↗

The concept of habitat diversity between and within ecosystems applied to river side-arm restoration.

Since returning an ecosystem to its pristine state may not be realistic in every situation, the concept of habitat diversity is proposed to help decision-makers in defining realistic restoration objectives. In order to maintain habitat diversity and enhance the long-term success of restoration, process-oriented projects should be preferred to species-oriented ones. Because the hydrogeomorphological processes that influence biodiversity operate at different spatiotemporal scales, three scales are considered: river sectors, floodplain waterbodies, and mesohabitats within each waterbody. Based on a bibliographical review, three major driving forces are proposed for incorporation into the design of restoration projects: (1) flow velocity and flood disturbances, (2) hydrological connectivity, and (3) water supply. On the sector scale, increased habitat diversity between waterbodies can be achieved by combining various intensities of these driving forces. On the waterbody scale, increased habitat diversity within the ecosystem can be achieved by varying water depth, velocity, and substrate. The concept is applied to a Rhĵne River sector (France) where three terrestrialized side arms will be restored. Two were designed to be flood scoured, one having an additional supply of groundwater, the other being connected to the river at both ends. The third cannot be scoured by floods because of upstream construction and would be supplied by river backflow through a downstream connection. Habitat diversity within the ecosystem is exemplified on one side arm through the design of a sinuous pathway combined with variation of water depth, wetted width, and substrate grain size. Self-colonization of the side arms is expected owing to the restoration of connectivity to upstream sources of potential colonizers.

Animals↗

Temporal variation in delta(13)C of ecosystem respiration in the Pacific Northwest: links to moisture stress.

We measured seasonal and interannual variations in delta(13)C values within the carbon reservoirs (leaves and soil) and CO(2) fluxes (soil and ecosystem respired CO(2)) of an old growth coniferous forest in the Pacific Northwest USA with relation to local meteorological conditions. There were significant intra-annual and interannual differences in the carbon isotope ratios of CO(2) respired at both the ecosystem (delta(13)C(R)) and the soil levels (delta(13)C(R-soil)), but only limited variations in the carbon isotope ratios of carbon stocks. The delta(13)C(R) values varied by as much as 4.4 per thousand over a growing season, while delta(13)C(R-soil )values changed as much as 6.2 per thousand. The delta(13)C of soil organic carbon (delta(13)C(SOC)) and needle organic carbon (delta(13)C(P)) exhibited little or no significant changes over the course of this study. Carbon isotope discrimination within leaves (Delta(p)) showed systematic decreases with increased canopy height, but remained fairly constant throughout the year (Delta(p)=17.9 per thousand -19.2 per thousand at the top of the canopy, Delta(p)=19.6 per thousand -20.9 per thousand at mid-canopy, Delta(p)=23.3 per thousand -25.1 per thousand at the canopy base). The temporal variation in the delta(13)C of soil and ecosystem respired CO(2) was correlated ( r=0.93, P<0.001) with soil moisture levels, with dry summer months having the most (13)C-enriched values. The dynamic seasonal changes in delta(13)C of respired CO(2) are hypothesized to be the result of fast cycling of recently fixed carbon back to the atmosphere. One scaling consequence of the seasonal and interannual variations in delta(13)C(R) is that inversion-based carbon-cycle models dependent on observed atmospheric CO(2) concentration and isotope values may be improved by incorporating dynamic delta(13)C(R) values to interpret regional carbon sink strength.

Carbon Dioxide↗

Experimental evidence that terrestrial carbon subsidies increase CO2 flux from lake ecosystems.

Subsidies are donor-controlled inputs of nutrients and energy that can affect ecosystem-level processes in a recipient environment. Lake ecosystems receive large inputs of terrestrial carbon (C) in the form of dissolved organic matter (DOM). DOM inputs may energetically subsidize heterotrophic bacteria and determine whether lakes function as sources or sinks of atmospheric CO(2). I experimentally tested this hypothesis using a series of mesocosm experiments in New England lakes. In the first experiment, I observed that CO(2) flux increased by 160% 4 days following a 1,000 microM C addition in the form of DOM. However, this response was relatively short lived, as there was no effect of DOM enrichment on CO(2) flux beyond 8 days. In a second experiment, I demonstrated that peak CO(2) flux from mesocosms in two lakes increased linearly over a broad DOM gradient (slope for both lakes=0.02+/-0.001 mM CO(2).m(-2) day(-1) per microM DOC, mean+/-SE). Concomitant changes in bacterial productivity and dissolved oxygen strengthen the inference that increasing CO(2) flux resulted from the metabolism of DOM. I conducted two additional studies to test whether DOM-correlated attributes were responsible for the observed change in plankton metabolism along the subsidy gradient. First, terrestrial DOM reduced light transmittance, but experimental shading revealed that this was not responsible for the observed patterns of CO(2) flux. Second, organically bound nitrogen (N) and phosphorus (P) accompanied DOM inputs, but experimental nutrient additions (without organic C) caused mesocosms to be saturated with CO(2). Together, these results suggest that C content of terrestrial DOM may be an important subsidy for freshwater bacteria that can influence whether recipient aquatic ecosystems are sources or sinks of atmospheric CO(2).

Carbon↗

Hierarchy of responses to resource pulses in arid and semi-arid ecosystems.

In arid/semi-arid ecosystems, biological resources, such as water, soil nutrients, and plant biomass, typically go through periods of high and low abundance. Short periods of high resource abundance are usually triggered by rainfall events, which, despite of the overall scarcity of rain, can saturate the resource demand of some biological processes for a time. This review develops the idea that there exists a hierarchy of soil moisture pulse events with a corresponding hierarchy of ecological responses, such that small pulses only trigger a small number of relatively minor ecological events, and larger pulses trigger a more inclusive set and some larger ecological events. This framework hinges on the observation that many biological state changes, where organisms transition from a state of lower to higher physiological activity, require a minimal triggering event size. Response thresholds are often determined by the ability of organisms to utilize soil moisture pulses of different infiltration depth or duration. For example, brief, shallow pulses can only affect surface dwelling organisms with fast response times and high tolerance for low resource levels, such as some species of the soil micro-fauna and -flora, while it takes more water and deeper infiltration to affect the physiology, growth or reproduction of higher plants. This review first discusses how precipitation, climate and site factors translate into soil moisture pulses of varying magnitude and duration. Next, the idea of the response hierarchy for ecosystem processes is developed, followed by an exploration of the possible evolutionary background for the existence of response thresholds to resource pulses. The review concludes with an outlook on global change: does the hierarchical view of precipitation effects in ecosystems provide new perspectives on the future of arid/semiarid lands?

Adaptation, Physiological↗

Are boreal ecosystems susceptible to alien plant invasion? Evidence from protected areas.

Although biological invasion by alien species is a major contributor to loss of indigenous biological diversity, few studies have examined the susceptibility of the boreal biome to invasion. Based on studies of other ecosystems, we hypothesized that alien plants will be restricted to disturbed areas near human activity and will not be found in natural areas of boreal ecosystems in Gros Morne National Park (Canada), a protected area experiencing a wide range of disturbance regimes. The distribution of alien plants in the region was evaluated using surveys, and study sites were established in naturally and anthropogenically disturbed habitats that had been invaded. Within study sites, randomization tests evaluated the importance of disturbance to alien plant invasion by examining changes in environmental conditions and species abundance within various disturbance regimes, while the importance of site characteristics limiting the distribution of alien plants were examined using Canonical Correspondence Analysis. Consistent with studies in a variety of biomes, areas of high disturbance and human activity had the greatest abundance of resources and the highest percentage of alien species. However, contrary to our hypothesis, natural areas of boreal ecosystems were found susceptible to alien plant invasion. Vegetation types vulnerable to invasion include forests, riparian areas, fens, and alpine meadows. Natural disturbance occurring in these vegetation types caused increases in bare ground and/or light availability facilitating alien plant invasion. Although high soil pH was associated with alien plants in these areas, disturbance was not found to cause changes in soil pH, suggesting susceptibility to invasion is pre-determined by bedrock geology or other factors influencing soil pH. Moose (Alces alces), a non-native herbivore, acts as the primary conduit for alien plant invasion in GMNP by dispersing propagules and creating or prolonging disturbance by trampling and browsing vegetation. The recurrent nature of disturbance within the boreal biome and its interaction with site conditions and herbivores enables alien plants to persist away from areas of high human activity. Managers of natural lands should monitor such interactions to decrease the invasion potential of alien plants.

Analysis of Variance↗

Ecological effects of low-level phosphorus additions on two plant communities in a neotropical freshwater wetland ecosystem.

We conducted a low-level phosphorus (P) enrichment study in two oligotrophic freshwater wetland communities (wet prairies [WP] and sawgrass marsh [SAW]) of the neotropical Florida Everglades. The experiment included three P addition levels (0, 3.33, and 33.3 mg P m(-2) month(-1)), added over 2 years, and used in situ mesocosms located in northeastern Everglades National Park, Fla., USA. The calcareous periphyton mat in both communities degraded quickly and was replaced by green algae. In the WP community, we observed significant increases in net aboveground primary production (NAPP) and belowground biomass. Aboveground live standing crop (ALSC) did not show a treatment effect, though, because stem turnover rates of Eleocharis spp., the dominant emergent macrophyte in this community, increased significantly. Eleocharis spp. leaf tissue P content decreased with P additions, causing higher C:P and N:P ratios in enriched versus unenriched plots. In the SAW community, NAPP, ALSC, and belowground biomass all increased significantly in response to P additions. Cladium jamaicense leaf turnover rates and tissue nutrient content did not show treatment effects. The two oligotrophic communities responded differentially to P enrichment. Periphyton which was more abundant in the WP community, appeared to act as a P buffer that delayed the response of other ecosystem components until after the periphyton mat had disappeared. Periphyton played a smaller role in controlling ecosystem dynamics and community structure in the SAW community. Our data suggested a reduced reliance on internal stores of P by emergent macrophytes in the WP that were exposed to P enrichment. Eleocharis spp. rapidly recycled P through more rapid aboveground turnover. In contrast, C. jamaicense stored added P by initially investing in belowground biomass, then shifting growth allocation to aboveground tissue without increasing leaf turnover rates. Our results suggest that calcareous wetland systems throughout the Caribbean, and oligotrophic ecosystems in general, respond rapidly to low-level additions of their limiting nutrient.

Analysis of Variance↗

Dimensional approaches to designing better experimental ecosystems: a practitioners guide with examples.

Enclosed, experimental ecosystems ("mesocosms") are now widely used research tools in ecology. However, the small size, short duration and often simplified biological and physical complexity of mesocosm experiments raises questions about extrapolating results from these miniaturized ecosystems to nature. Dimensional analysis, a technique widely used in engineering to create scale models, employs "compensatory distortion" as a means of maintaining functional similarity in properties and relationships of interest. An earlier paper outlined a general approach to applying dimensional analysis to the construction and interpretation of mesocosm experiments (Petersen and Hastings in Am Nat 157:324, 2001). In this paper we use examples, largely drawn from the aquatic literature, to illustrate how dimensional approaches might be used to maintain key ecological properties. Such key properties include effective habitat size, environmental variability, vertical and horizontal gradients, and interactions among habitats. We distinguish both continuous and discrete approaches that can be used to achieve functional similarity through compensatory distortion. In addition to its potential as a tool for improving the realism of experimental ecosystems, the dimensional approach points towards new options for developing, testing and advancing our understanding of scaling relationships in nature.

Body Size↗

Meta-ecosystems and biological energy transport from ocean to coast: the ecological importance of herring migration.

Ecosystems are not closed, but receive resource subsidies from other ecosystems. Energy, material and organisms are moved between systems by physical vectors, but migrating animals also transport resources between systems. We report on large scale energy transport from ocean to coast by a migrating fish population, the Norwegian spring-spawning (NSS) herring Clupea harengus. We observe a rapid body mass increase during parts of the annual, oceanic feeding migration and we use a bioenergetics model to quantify energy consumption. The model predicts strong seasonal variation in food consumption with a marked peak in late May to July. The copepod Calanus finmarchicus is the most important prey and 23 x 10(6) tones (wet weight) of C. finmarchicus is consumed annually. The annual consumption-biomass ratio is 5.2. During the feeding migration 17% of consumed energy is converted to body mass. The biomass transported to the coast and left as reproductive output is estimated from gonad weight and is about 1.3 x 10(6) tones for the current population. This transport is to our knowledge the world's largest flux of energy caused by a single population. We demonstrate marked temporal variation in transport during the last century and discuss the effects of NSS herring in the ocean, as a major consumer, and at the coast, where eggs and larvae are important for coastal predators. In particular, we suggest that the rapid decline of lobster Homarus gammarus landings in Western Norway during the 1960s was related to the collapse of NSS herring. We also discuss spatial variation in energy transport caused by changed migration patterns. Both climate and fisheries probably triggered historical changes in the migration patterns of NSS herring. New migration routes emerge at the level of individuals, which in turn determines where resources are gathered and delivered, and therefore, how meta-ecosystems function.

Animal Migration↗

Habitat structure, trophic structure and ecosystem function: interactive effects in a bromeliad-insect community.

Although previous studies have shown that ecosystem functions are affected by either trophic structure or habitat structure, there has been little consideration of their combined effects. Such interactions may be particularly important in systems where habitat and trophic structure covary. I use the aquatic insects in bromeliads to examine the combined effects of trophic structure and habitat structure on a key ecosystem function: detrital processing. In Costa Rican bromeliads, trophic structure naturally covaries with both habitat complexity and habitat size, precluding any observational analysis of interactions between factors. I therefore designed mesocosms that allowed each factor to be manipulated separately. Increases in mesocosm complexity reduced predator (damselfly larva) efficiency, resulting in high detritivore abundances, indirectly increasing detrital processing rates. However, increased complexity also directly reduced the per capita foraging efficiency of the detritivores. Over short time periods, these trends effectively cancelled each other out in terms of detrital processing. Over longer time periods, more complex patterns emerged. Increases in mesocosm size also reduced both predator efficiency and detritivore efficiency, leading to no net effect on detrital processing. In many systems, ecosystem functions may be impacted by strong interactions between trophic structure and habitat structure, cautioning against examining either effect in isolation.

Animals↗

Threshold levels for effects of insecticides in freshwater ecosystems: a review.

A literature review of freshwater (model) ecosystem studies with neurotoxic insecticides was performed to assess ecological threshold levels, to compare these levels with the first tier approach within European Union (EU) administration procedures, and to evaluate the ecological consequences of exceeding these thresholds. Studies published between 1980 and 2001 were reviewed. Most studies covered organophosphates and synthetic pyrethroids in lentic waters. The most sensitive taxa were representatives of crustaceans, insects and fish. Based on toxic units, threshold values were equivalent for compounds with a similar mode of action. This also accounted for the nature and magnitude of direct effects at higher concentrations. Although laboratory single species toxicity tests may not allow predictions on precise ecological effects, some generalisations on effects and recovery can be made with respect to acute standard laboratory EC50 data. The NOEC(ecosystem) usually is a factor of 10 or more higher than first tier acceptable concentrations, particularly in the case of single applications and acetylcholinesterase inhibitors. Acceptable concentrations, as set by the EU first tier approach, appear to be protective. Recovery of sensitive endpoints usually occurs within 2 months of the (last) application when peak concentrations remain lower than (0.1-1) x EC50 of the most sensitive standard test species. The consistency of response patterns found in model ecosystem studies can be useful when estimating the ecological risks of pesticides. The use of an effect classification system was also helpful in evaluating effects.

Animals↗

Landscape change and ecosystem classification in a municipal district of a small city (Isernia, Central Italy).

Landscape changes taking place from 1954 to 1992 in the muncipal district of Isernia city (Central Italy) were described in relation to a system of ecosystem classification. Isernia municipal district was selected for study because recent historic changes in this area represent a typical example of landscape transformation similar to many small cities of Italy and other Mediterranean countries. To assess overall changes, three land cover maps (scale 1:25,000) were derived from panchromatic aerial photographs and field surveys. These were then digitalised in a Geographic Information System. A Land Facet (LF) map was derived by combining a phytoclimatic, a lithostatigrafic and a topographic map, and then digitalised as data layers in the same GIS. Results demonstrated two main landscape transformation trends: forest and semi-natural areas increased (8%), whereas agricultural areas decreased (12%). The urban area was relatively small during the entire analysed period, growing from 1% in 1954, to just 5% in 1992. Forest coverage was significant on reliefs, on hillside ecosystems such as limestone and on clay and marl hills LF. Arable land was particularly significant in flat ecosystems with deeper soils, such as on recent alluvial plain LF. These temporal changes were interpreted as being related to the replacement of traditional farming methods (grazing pastures) with more intensive methods (crop fields), especially on alluvial plains.

Agriculture↗

Quantifying carbon budgets of conifer Mediterranean forest ecosystems, Turkey.

Aboveground biomass, aboveground litterfall, and leaf litter decomposition of five indigenous tree stands (pure stands of Pinus brutia, Pinus nigra, Cedrus libani, Juniperus excelsa, and a mixed stand of Abies cilicica, P. nigra, and C. libani) were measured in an eastern Mediterranean evergreen needleleaf forest of Turkey. Measurements were converted to regional scale estimates of carbon (C) stocks and fluxes of forest ecosystems, based on general non-site-specific allometric relationships. Mean C stock of the conifer forests was estimated as 97.8 +/- 79 Mg C ha(-1) consisting of 83.0 +/- 67 Mg C ha(-1) in the aboveground and 14.8 +/- 12 Mg C ha(-1) in the belowground biomass. The forest stands had mean soil organic carbon (SOC) and nitrogen (SON) stocks of 172.0 +/- 25.7 Mg C ha(-1) and 9.2 +/- 1.2 Mg N ha(-1), respectively. Mean total monthly litterfall was 376.2 +/- 191.3 kg C ha(-1), ranging from 641 +/- 385 kg C ha(-1) for Pinus brutia to 286 +/- 82 kg C ha(-1) for Cedrus libani. Decomposition rate constants (k) for pine needles were 0.0016 for Cedrus libani, 0.0009 for Pinus nigra, 0.0006 for the mixed stand, and 0.0005 day(-1) for Pinus brutia and Juniperus excelsa. Estimation of components of the C budgets revealed that the forest ecosystems were net C sinks, with a mean sequestration rate of 2.0 +/- 1.1 Mg C ha(-1) yr(-1) ranging from 3.2 +/- 2 Mg C ha(-1) for Pinus brutia to 1.6 +/- 0.6 Mg C ha(-1) for Cedrus libani. Mean net ecosystem productivity (NEP) resulted in sequestration of 98.4 +/- 54.1 Gg CO2 yr(-1) from the atmosphere when extrapolated for the entire study area of 134.2 km2 (Gg = 10(9) g). The quantitative C data from the study revealed the significance of the conifer Mediterranean forests as C sinks.

Biomass↗