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1.
Sea water temperature affects all biological and ecological processes that ultimately impact ecosystem functioning. In this study, we examine the influence of temperature on global biomass transfers from marine secondary production to fish stocks. By combining fisheries catches in all coastal ocean areas and life‐history traits of exploited marine species, we provide global estimates of two trophic transfer parameters which determine biomass flows in coastal marine food web: the trophic transfer efficiency (TTE) and the biomass residence time (BRT) in the food web. We find that biomass transfers in tropical ecosystems are less efficient and faster than in areas with cooler waters. In contrast, biomass transfers through the food web became faster and more efficient between 1950 and 2010. Using simulated changes in sea water temperature from three Earth system models, we project that the mean TTE in coastal waters would decrease from 7.7% to 7.2% between 2010 and 2100 under the ‘no effective mitigation’ representative concentration pathway (RCP8.5), while BRT between trophic levels 2 and 4 is projected to decrease from 2.7 to 2.3 years on average. Beyond the global trends, we show that the TTEs and BRTs may vary substantially among ecosystem types and that the polar ecosystems may be the most impacted ecosystems. The detected and projected changes in mean TTE and BRT will undermine food web functioning. Our study provides quantitative understanding of temperature effects on trophodynamic of marine ecosystems under climate change.  相似文献   

2.
Loss of biodiversity and nutrient enrichment are two of the main human impacts on ecosystems globally, yet we understand very little about the interactive effects of multiple stressors on natural communities and how this relates to biodiversity and ecosystem functioning. Advancing our understanding requires the following: (1) incorporation of processes occurring within and among trophic levels in natural ecosystems and (2) tests of context‐dependency of species loss effects. We examined the effects of loss of a key predator and two groups of its prey on algal assemblages at both ambient and enriched nutrient conditions in a marine benthic system and tested for interactions between the loss of functional diversity and nutrient enrichment on ecosystem functioning. We found that enrichment interacted with food web structure to alter the effects of species loss in natural communities. At ambient conditions, the loss of primary consumers led to an increase in biomass of algae, whereas predator loss caused a reduction in algal biomass (i.e. a trophic cascade). However, contrary to expectations, we found that nutrient enrichment negated the cascading effect of predators on algae. Moreover, algal assemblage structure varied in distinct ways in response to mussel loss, grazer loss, predator loss and with nutrient enrichment, with compensatory shifts in algal abundance driven by variation in responses of different algal species to different environmental conditions and the presence of different consumers. We identified and characterized several context‐dependent mechanisms driving direct and indirect effects of consumers. Our findings highlight the need to consider environmental context when examining potential species redundancies in particular with regard to changing environmental conditions. Furthermore, non‐trophic interactions based on empirical evidence must be incorporated into food web‐based ecological models to improve understanding of community responses to global change.  相似文献   

3.
Anthropogenic activities such as land‐use change, pollution and fishing impact the trophic structure of coral reef fishes, which can influence ecosystem health and function. Although these impacts may be ubiquitous, they are not consistent across the tropical Pacific Ocean. Using an extensive database of fish biomass sampled using underwater visual transects on coral reefs, we modelled the impact of human activities on food webs at Pacific‐wide and regional (1,000s–10,000s km) scales. We found significantly lower biomass of sharks and carnivores, where there were higher densities of human populations (hereafter referred to as human activity); however, these patterns were not spatially consistent as there were significant differences in the trophic structures of fishes among biogeographic regions. Additionally, we found significant changes in the benthic structure of reef environments, notably a decline in coral cover where there was more human activity. Direct human impacts were the strongest in the upper part of the food web, where we found that in a majority of the Pacific, the biomass of reef sharks and carnivores were significantly and negatively associated with human activity. Finally, although human‐induced stressors varied in strength and significance throughout the coral reef food web across the Pacific, socioeconomic variables explained more variation in reef fish trophic structure than habitat variables in a majority of the biogeographic regions. Notably, economic development (measured as GDP per capita) did not guarantee healthy reef ecosystems (high coral cover and greater fish biomass). Our results indicate that human activities are significantly shaping patterns of trophic structure of reef fishes in a spatially nonuniform manner across the Pacific Ocean, by altering processes that organize communities in both “top‐down” (fishing of predators) and “bottom‐up” (degradation of benthic communities) contexts.  相似文献   

4.
Inverse trophic cascades are a well explored and common consequence of the local depletion or extinction of top predators in natural ecosystems. Despite a large body of research, the cascading effects of predator removal on ecosystem functions are not as well understood. Developing microcosm experiments, we explored food web changes in trophic structure and ecosystem functioning following biomass removal of top predators in representative temperate and tropical rock pool communities that contained similar assemblages of zooplankton and benthic invertebrates. We observed changes in species abundances following predator removal in both temperate and tropical communities, in line with expected inverse effects of a trophic cascade, where predation release benefits the predator’s preys and competitors and impacts the preys of the latter. We also observed several changes at the community and ecosystem levels including a decrease in total abundance and mean trophic level of the community, and changes in chlorophyll-a and total dissolved particles. Our results also showed an increase in variability of both community and ecosystem processes following the removal of predators. These results illustrate how predator removal can lead to inverse trophic cascades both in structural and functioning properties, and can increase variability of ecosystem processes. Although observed patterns were consistent between tropical and temperate communities following an inverse cascade pattern, changes were more pronounced in the temperate community. Therefore, aquatic food webs may have inherent traits that condition ecosystem responses to changes in top-down trophic control and render some aquatic ecosystems especially sensitive to the removals of top predators.  相似文献   

5.
Reynolds PL  Bruno JF 《PloS one》2012,7(5):e36196
Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.  相似文献   

6.
Species diversity affects the functioning of ecosystems, including the efficiency by which communities capture limited resources, produce biomass, recycle and retain biologically essential nutrients. These ecological functions ultimately support the ecosystem services upon which humanity depends. Despite hundreds of experimental tests of the effect of biodiversity on ecosystem function (BEF), it remains unclear whether diversity effects are sufficiently general that we can use a single relationship to quantitatively predict how changes in species richness alter an ecosystem function across trophic levels, ecosystems and ecological conditions. Our objective here is to determine whether a general relationship exists between biodiversity and standing biomass. We used hierarchical mixed effects models, based on a power function between species richness and biomass production (Y = a × Sb), and a database of 374 published experiments to estimate the BEF relationship (the change in biomass with the addition of species), and its associated uncertainty, in the context of environmental factors. We found that the mean relationship (b = 0.26, 95% CI: 0.16, 0.37) characterized the vast majority of observations, was robust to differences in experimental design, and was independent of the range of species richness levels considered. However, the richness–biomass relationship varied by trophic level and among ecosystems; in aquatic systems b was nearly twice as large for consumers (herbivores and detritivores) compared to primary producers; in terrestrial ecosystems, b for detritivores was negative but depended on few studies. We estimated changes in biomass expected for a range of changes in species richness, highlighting that species loss has greater implications than species gains, skewing a distribution of biomass change relative to observed species richness change. When biomass provides a good proxy for processes that underpin ecosystem services, this relationship could be used as a step in modeling the production of ecosystem services and their dependence on biodiversity.  相似文献   

7.
Competition between marine mammals and fisheries for marine resources-whether real or perceived-has become a major issue for several countries and in international fora. We examined trophic interactions between marine mammals and fisheries based on a resource overlap index, using seven Ecopath models including marine mammal groups. On a global scale, most food consumed by marine mammals consisted of prey types that were not the main target of fisheries. For each ecosystem, the primary production required (PPR) to sustain marine mammals was less than half the PPR to sustain fisheries catches. We also developed an index representing the mean trophic level of marine mammal's consumption (TL(Q)) and compared it with the mean trophic level of fisheries' catches (TL(C)). Our results showed that overall TL(Q) was lower than TL(C) (2.88 versus 3.42). As fisheries increasingly exploit lower-trophic level species, the competition with marine mammals may become more important. We used mixed trophic impact analysis to evaluate indirect trophic effects of marine mammals, and in some cases found beneficial effects on some prey. Finally, we assessed the change in the trophic structure of an ecosystem after a simulated extirpation of marine mammal populations. We found that this lead to alterations in the structure of the ecosystems, and that there was no clear and direct relationship between marine mammals' predation and the potential catch by fisheries. Indeed, total biomass, with no marine mammals in the ecosystem, generally remained surprisingly similar, or even decreased for some species.  相似文献   

8.
Humans have drastically altered the abundance of animals in marine ecosystems via exploitation. Reduced abundance can destabilize food webs, leading to cascading indirect effects that dramatically reorganize community structure and shift ecosystem function. However, the additional implications of these top‐down changes for biogeochemical cycles via consumer‐mediated nutrient dynamics (CND) are often overlooked in marine systems, particularly in coastal areas. Here, we review research that underscores the importance of this bottom‐up control at local, regional, and global scales in coastal marine ecosystems, and the potential implications of anthropogenic change to fundamentally alter these processes. We focus attention on the two primary ways consumers affect nutrient dynamics, with emphasis on implications for the nutrient capacity of ecosystems: (1) the storage and retention of nutrients in biomass, and (2) the supply of nutrients via excretion and egestion. Nutrient storage in consumer biomass may be especially important in many marine ecosystems because consumers, as opposed to producers, often dominate organismal biomass. As for nutrient supply, we emphasize how consumers enhance primary production through both press and pulse dynamics. Looking forward, we explore the importance of CDN for improving theory (e.g., ecological stoichiometry, metabolic theory, and biodiversity–ecosystem function relationships), all in the context of global environmental change. Increasing research focus on CND will likely transform our perspectives on how consumers affect the functioning of marine ecosystems.  相似文献   

9.
Climate change is altering the rate and distribution of primary production in the world's oceans. Primary production is critical to maintaining biodiversity and supporting fishery catches, but predicting the response of populations to primary production change is complicated by predation and competition interactions. We simulated the effects of change in primary production on diverse marine ecosystems across a wide latitudinal range in Australia using the marine food web model Ecosim. We link models of primary production of lower trophic levels (phytoplankton and benthic producers) under climate change with Ecosim to predict changes in fishery catch, fishery value, biomass of animals of conservation interest, and indicators of community composition. Under a plausible climate change scenario, primary production will increase around Australia and generally this benefits fisheries catch and value and leads to increased biomass of threatened marine animals such as turtles and sharks. However, community composition is not strongly affected. Sensitivity analyses indicate overall positive linear responses of functional groups to primary production change. Responses are robust to the ecosystem type and the complexity of the model used. However, model formulations with more complex predation and competition interactions can reverse the expected responses for some species, resulting in catch declines for some fished species and localized declines of turtle and marine mammal populations under primary productivity increases. We conclude that climate‐driven primary production change needs to be considered by marine ecosystem managers and more specifically, that production increases can simultaneously benefit fisheries and conservation. Greater focus on incorporating predation and competition interactions into models will significantly improve the ability to identify species and industries most at risk from climate change.  相似文献   

10.
Experiments with realistic scenarios of species loss from multitrophic ecosystems may improve insight into how biodiversity affects ecosystem functioning. Using 1000 L mesocoms, we examined effects of nonrandom species loss on community structure and ecosystem functioning of experimental food webs based on multitrophic tropical floodplain lagoon ecosystems. Realistic biodiversity scenarios were developed based on long-term field surveys, and experimental assemblages replicated sequential loss of rare species which occurred across all trophic levels of these complex food webs. Response variables represented multiple components of ecosystem functioning, including nutrient cycling, primary and secondary production, organic matter accumulation and whole ecosystem metabolism. Species richness significantly affected ecosystem function, even after statistically controlling for potentially confounding factors such as total biomass and direct trophic interactions. Overall, loss of rare species was generally associated with lower nutrient concentrations, phytoplankton and zooplankton densities, and whole ecosystem metabolism when compared with more diverse assemblages. This pattern was also observed for overall ecosystem multifunctionality, a combined metric representing the ability of an ecosystem to simultaneously maintain multiple functions. One key exception was attributed to time-dependent effects of intraguild predation, which initially increased values for most ecosystem response variables, but resulted in decreases over time likely due to reduced nutrient remineralization by surviving predators. At the same time, loss of species did not result in strong trophic cascades, possibly a result of compensation and complexity of these multitrophic ecosystems along with a dominance of bottom-up effects. Our results indicate that although rare species may comprise minor components of communities, their loss can have profound ecosystem consequences across multiple trophic levels due to a combination of direct and indirect effects in diverse multitrophic ecosystems.  相似文献   

11.
In light of current global changes to ecosystem function (e.g. climate change, trophic downgrading, and invasive species), there has been a recent surge of interest in exploring differences in nutrient cycling among ecosystem types. In particular, a growing awareness has emerged concerning the importance of scavenging in food web dynamics, although no studies have focused specifically on exploring differences in carrion consumption between aquatic and terrestrial ecosystems. In this forum we synthesize the scavenging literature to elucidate differences in scavenging dynamics between terrestrial and marine ecosystems, and identify areas where future research is needed to more clearly understand the role of carrion consumption in maintaining ecosystem function within each of these environments. Although scavenging plays a similar functional role in terrestrial and aquatic food webs, here we suggest that several fundamental differences exist in scavenging dynamics among these ecosystem types due to the unique selection pressures imposed by the physical properties of water and air. In particular, the movement of carcasses in marine ecosystems (e.g. wave action, upwelling, and sinking) diffuses biological activity associated with scavenging and decomposition across large, three‐dimensional spatial scales, creating a unique spatial disconnect between the processes of production, scavenging, and decomposition, which in contrast are tightly linked in terrestrial ecosystems. Moreover, the limited role of bacteria and temporal stability of environmental conditions on the sea floor appears to have facilitated the evolution of a much more diverse community of macrofauna that relies on carrion for a higher portion of its nutrient consumption than is present in terrestrial ecosystems. Our observations are further discussed as they pertain to the potential impacts of climate change and trophic downgrading (i.e. removal of apex consumers from ecosystems) on scavenging dynamics within marine and terrestrial ecosystems.  相似文献   

12.
One challenge in merging community and ecosystem ecology is to integrate the complexity of natural multitrophic communities into concepts of ecosystem functioning. Here, we combine food‐web and allometry theories to demonstrate that primary production, as measured by the total nutrient uptake of the multitrophic community, is determined by vertical diversity (i.e. food web's maximum trophic level) and structure (i.e. distributions of species and their abundances and metabolic rates across trophic levels). In natural ecosystems, the community size distribution determines all these vertical patterns and thus the total nutrient uptake. Our model suggests a vertical diversity hypothesis (VDH) for ecosystem functioning in complex food webs. It predicts that, under a given nutrient supply, the total nutrient uptake increases exponentially with the maximum trophic level in the food web and it increases with its maximum body size according to a power law. The VDH highlights the effect of top–down regulation on plant nutrient uptake, which complements traditional paradigms that emphasised the bottom–up effect of nutrient supply on vertical diversity. We conclude that the VDH contributes to a synthetic framework for understanding the relationship between vertical diversity and ecosystem functioning in food webs and predicting the impacts of global changes on multitrophic ecosystems.  相似文献   

13.
The organic content of sediments, often considered as an important trophic source for the benthos, can be used as a measurement of the trophic status of marine ecosystems. Soft-bottom communities were studied between Leghorn and Elba Island, from 5 to 500 m depth. Total biomass and trophic structure (percent dominance of trophic guilds) were related to organic carbon content and C/N ratio of sediments. Sediment organic matter, depending on its quantity and quality, could act as a food source or a stress source for the community.  相似文献   

14.
Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.  相似文献   

15.
The size composition of primary producers is important for how energy is channeled through a food web and on to the higher trophic levels and eventually to fisheries. To evaluate this, we studied the productive patterns for large (micro) versus small (nano) phytoplankton in two south marine Patagonian ecosystems: The Inner Sea of Chiloe—ISCh and, Moraleda Channel—MCh. We built Ecopath models (EwE), and evaluated the hypothesis that the overall primary productivity—rather than the ratio of large to small primary producers—constitutes an adequate proxy for predicting the amount of secondary and tertiary production and biomass (up to the fisheries). The EwE model included four small-scale fisheries and 36 functional groups. The functioning of both ecosystems was similar but the ecosystem parameters (biomass, energy transfer efficiencies from primary producers, secondary, and tertiary production) were twice as much in the basin with more microphytoplankton biomass. Overall, the hypothesis was rejected, albeit it was possible to highlight the importance of the quality and size spectrum of plankton on the structure of marine ecosystem, and to demonstrate the key role of the microbial loop over traditional food web in the functioning of the carbon biological pump in Patagonia ecosystems.  相似文献   

16.
The exceptional diversity of large mammals in African savannas provides an ideal opportunity to explore the relative importance of top‐down and bottom‐up controls of large terrestrial herbivore communities. Recent work has emphasized the role of herbivore and carnivore body size in shaping these trophic relationships. However, the lack of across‐ecosystem comparisons using a common methodology prohibits general conclusions. Here we used published data on primary production, herbivore and carnivore densities and diets to estimate the consumption fluxes between three trophic levels in four African savanna ecosystems. Our food web approach suggests that the body size distribution within and across trophic levels has a strong influence on the strength of top‐down control of herbivores by carnivores and on consumption fluxes within ecosystems, as predicted by theoretical food web models. We generalize findings from the Serengeti ecosystem that suggest herbivore species below 150 kg are more likely to be limited by predation. We also emphasize the key functional role played by the largest species at each trophic level. The abundance of the largest herbivore species largely governs the consumption of primary production in resident communities. Similarly, predator guilds in which the largest carnivore species represent a larger share of carnivore biomass are likely to exert a stronger top‐down impact on herbivores. Our study shows how a food web approach allows integrating current knowledge and offers a powerful framework to better understand the functioning of ecosystems.  相似文献   

17.
Soil fauna can be an important regulator of community parameters and ecosystem processes, but there have been few quantitative syntheses of the role of soil fauna in terrestrial soil communities and ecosystems. Here, we conducted a meta‐analysis to investigate the impacts of invertebrate soil micro‐ and mesofauna (grazers and predators) on plant productivity and microbial biomass. Overall our results indicate that an increase in the biomass of soil fauna increased aboveground plant productivity across ecosystems by 35% and decreased microbial biomass by 8%. In addition, we found no evidence for trophic cascades in terrestrial soil food webs, but the bacterivorous component of soil fauna influenced plant productivity and microbial biomass more than did the fungivorous component. Furthermore, changes in the biomass of soil fauna differentially affected plant productivity among plant functional groups: a higher biomass of soil fauna increased aboveground productivity by 70% in coniferous systems. However, in ecosystems dominated by legumes, a functional group with lower inorganic nitrogen requirements, there was no response of aboveground productivity to increases in the biomass of soil fauna. In sum, the results of this meta‐analysis indicate that soil fauna help to regulate ecosystem production, especially in nutrient‐limited ecosystems.  相似文献   

18.
Global patterns of aquatic food chain length   总被引:5,自引:1,他引:4  
Food chain length is a fundamental ecosystem property, and plays a central role in determining ecosystem functioning. Recent advances in the field of stable isotope ecology allow the estimation of food chain length (FCL) from stable nitrogen isotope (δ15N) data. We conducted a global literature synthesis and estimated FCL for 219 lake, stream, and marine ecosystems. Streams had shorter food chains (∼3.5 trophic levels) than marine and lake ecosystems (∼4.0 trophic levels). In marine systems, inclusion of marine mammals increased FCL by 2/3 of a trophic level. For each ecosystem type, estimates of FCL were normally distributed and spanned two full trophic levels. Comparison with published connectance food webs revealed similar mean FCL values, though stable isotope-derived FCL estimates were less variable. At the global scale, FCL showed weak or no relationships with ecosystem size, mean annual air temperature, or latitude. Our study highlights the utility of stable isotopes for quantifying among-system food web variability, and the application of this approach for assessing global-scale patterns of food chain length.  相似文献   

19.
Viral lysis of microbial hosts releases organic matter that can then be assimilated by nontargeted microorganisms. Quantitative estimates of virus-mediated recycling of carbon in marine waters, first established in the late 1990s, were originally extrapolated from marine host and virus densities, host carbon content and inferred viral lysis rates. Yet, these estimates did not explicitly incorporate the cascade of complex feedbacks associated with virus-mediated lysis. To evaluate the role of viruses in shaping community structure and ecosystem functioning, we extend dynamic multitrophic ecosystem models to include a virus component, specifically parameterized for processes taking place in the ocean euphotic zone. Crucially, we are able to solve this model analytically, facilitating evaluation of model behavior under many alternative parameterizations. Analyses reveal that the addition of a virus component promotes the emergence of complex communities. In addition, biomass partitioning of the emergent multitrophic community is consistent with well-established empirical norms in the surface oceans. At steady state, ecosystem fluxes can be probed to characterize the effects that viruses have when compared with putative marine surface ecosystems without viruses. The model suggests that ecosystems with viruses will have (1) increased organic matter recycling, (2) reduced transfer to higher trophic levels and (3) increased net primary productivity. These model findings support hypotheses that viruses can have significant stimulatory effects across whole-ecosystem scales. We suggest that existing efforts to predict carbon and nutrient cycling without considering virus effects are likely to miss essential features of marine food webs that regulate global biogeochemical cycles.  相似文献   

20.
The Chukchi Sea is a seasonally ice-covered, marginal Arctic-shelf sea that possesses both large petroleum reserves and abundant biological communities, including migratory mammals and seabirds. We developed a mass balance food web model for the eastern Chukchi Sea to evaluate the trophic structure of this ecosystem and to compare food web properties of the Chukchi Sea to those of other high-latitude marine ecosystems. We compiled data on biomass levels, diet composition, demographic rates (production, consumption), and fishery removals, and used these data to construct an Ecopath trophic mass balance model. The majority of biomass was concentrated in benthic invertebrates and most of the mass flow above trophic level 2.0 was through these groups. We found that density estimates of most fish groups derived from trawl survey data using area-swept methods were insufficient to match the consumptive demands of predators, and that densities needed to be several-fold greater to meet modeled demand. We also used a set of system metrics derived from a common modeling framework to highlight differences in ecosystem structure between the eastern Chukchi Sea and other high-latitude systems. The extent of benthic dominance observed in the eastern Chukchi Sea was unique among the systems examined, both in terms of food web structure and associated mass flows between benthic and pelagic components. In relation to total biomass density, the eastern Chukchi Sea had low production when compared with the other systems, and this lower turnover rate suggests that recovery from disturbance might be slow.  相似文献   

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