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1.
Summary We tested Buchsbaum's hypothesis that food palatability in geese is determined by a hierarchy of feeding cues among which deterrent secondary metabolites (mostly phenols) have a primary role (Buchsbaum et al. 1984). In preference tests, greater snow goose feeding was slightly depressed when grass was sprayed with ferulic acid but not when grass was sprayed with p-coumaric and tannic acids. Extracts of Timothy grass, red clover or alfalfa sprayed on grass also failed to depress goose feeding. In a multifactor experiment, phenol and protein content and height of grass were manipulated simultaneously. When ferulic acid was sprayed, protein and phenol content interacted in determining goose feeding preferences; protein content had no effect in the absence of phenol but did have an effect when phenol was added. When tannic acid was used in a similar experiment, results were inconclusive because of a significant and complex interaction between protein content and height of grass. Our results generally failed to support Buchsbaum's hypothesis that phenol content of plants has a primary role in determining food preference in geese. Protein content of plants seemed to be a more important factor.  相似文献   

2.
Little is known about the stoichiometry of nutrient cycling by detritivores. Therefore, we explored stoichiometric relationships in an omnivorous/detritivorous fish (gizzard shad, Dorosoma cepedianum) in three lakes that differed in productivity. Gizzard shad can feed on plankton and sediment detritus, but in all three lakes adult gizzard shad derived >98% of carbon (C) and phosphorus (P), and >90% of nitrogen (N) from sediment detritus, and the remainder from zooplankton.
Gizzard shad selectively consumed detritus with higher C, N and P concentrations than ambient lake sediments. Selective detritivory (i.e. the nutrient content of consumed detritus divided by the nutrient content of ambient detritus) was most pronounced in the lake with the lowest detrital nutrient concentrations. N and P cycling rates per fish were also consistently higher in this lake, in agreement with the prediction of stoichiometry theory that excretion rates should increase with food nutrient content. Among-lake differences in nutrient cycling rates were unrelated to inter-lake variation in fish body nutrient contents, which was minimal. The N:P ratio excreted was near Redfield (∼14:1) in all three lakes.
Stoichiometric analyses showed that the C:N and C:P ratios of sediment detritus were much higher (∼2.8×) than ratios of gizzard shad bodies, revealing substantial N and P imbalances between consumers and their food source. Gizzard shad alleviate N imbalance by selectively feeding on high N detritus (low C:N, high N:P), and apparently alleviate P imbalance by excreting nutrients at a higher N:P than that of their food or their bodies. Thus, this detritivore apparently regulates nutrient acquisition and allocation via both pre-absorption processes (selective feeding) and post-absorptive processes (differential N and P excretion).  相似文献   

3.
Synopsis Detritus is an abundant but poor quality food source for consumers in salt marsh ecosystems. Here we present results of feeding experiments to determine the ability of Fundulus heteroclitus, Cyprinodon variegatus, and Mugil cephalus, three major detritivores in Great Sippewissett Marsh, Massachussetts, to assimilate detritus and use it for growth. C. variegatus, the sheepshead minnow, gained weight on a detrital diet, but F. heteroclitus, the marsh killifish, and juvenile M. cephalus, the striped mullet, lost weight and suffered high mortality on detrital diets. C. variegatus is a herbivore with morphological adaptations for ingesting plant material. F. heteroclitus is a carnivore poorly suited to effectively assimilate detritus from the diet. Although adult M. cephalus are adapted for ingesting and assimilating detrital material, the young may lack these adaptations and thus do not assimilate detritus. C. variegatus excretes ammonium at a lower rate than F. heteroclitus when fed animal food, or when starved for short periods of time. This protein sparing effect could be crucial for survival when detritus is the only food available at certain times of the year. While both C. variegatus and M. cephalus are visitors in the marsh and may leave to exploit food sources elsewhere, F. heteroclitus is a year-round resident and is the most abundant species of fish in the marsh. Yet when high quality food sources become scarce in late summer and fall, detritus, although plentiful, is apparently not a suitable alternative.To whom all correspondences should be addressed.  相似文献   

4.
Stream fungi have the capacity to degrade leaf litter and, through their activities, to transform it into a more palatable food source for invertebrate detritivores. The objectives of the present study were to characterize various aspects of fungal modification of the leaf substrate and to examine the effects these changes have on leaf palatability to detritivores. Fungal species were grown on aspen leaves for two incubation times. Leaves were analyzed to determine the weight loss, the degree of softening of the leaf matrix, and the concentrations of ATP and nitrogen associated with leaves. The activities of a protease and 10 polysaccharide-degrading enzymes produced by each fungus were also determined. Most fungi caused similar changes in physicochemical characteristics of the leaves. All fungi exhibited the capability to depolymerize pectin, xylan, and cellulose. Differences among fungi were found in their capabilities to produce protease and certain glycosidases. Leaf palatability was assessed by offering leaves of all treatments to larvae of two caddisfly shredders (Trichoptera). Feeding preferences exhibited by the shredders were similar and indicated that they perceived distinct differences among fungi. Two fungal species were highly consumed, some moderately and others only slightly. No relationships were found between any of the fungal characteristics measured and detritivore feeding preferences. Apparently, interspecific differences among fungi other than parameters associated with biomass or degradation of structural polysaccharides influence fungal palatability to caddisfly detritivores.  相似文献   

5.
Tomiko Ito 《Limnology》2005,6(2):73-78
Lepidostomatid caddisfly larvae are typical detritivores, but they occasionally eat small dead animals. A laboratory feeding experiment was conducted with Lepidostoma complicatum (Kobayashi) larvae using two different feeding treatments: leaves or leaves and dead chironomids. L. complicatum larvae showed significantly higher growth rates and adult emergence weight and a significantly earlier emergence for the leaves and dead chironomids treatment than for the leaves alone treatment. However, the adult emergence rate was not different between the two feeding treatments. Thus, it is apparent that ingestion of dead chironomids by detritivorous L. complicatum larvae positively influences larval growth rates, adult emergence weight, and larval development time.  相似文献   

6.
Detritivory is the dominant trophic paradigm in most terrestrial, aquatic, and marine ecosystems, yet accurate measurement of consumer trophic position within detrital (=“brown”) food webs has remained unresolved. Measurement of detritivore trophic position is complicated by the fact that detritus is suffused with microbes, creating a detrital complex of living and nonliving biomass. Given that microbes and metazoans are trophic analogues of each other, animals feeding on detrital complexes are ingesting other detritivores (microbes), which should elevate metazoan trophic position and should be rampant within brown food webs. We tested these hypotheses using isotopic (15N) analyses of amino acids extracted from wild and laboratory‐cultured consumers. Vertebrate (fish) and invertebrate detritivores (beetles and moths) were reared on detritus, with and without microbial colonization. In the field, detritivorous animal specimens were collected and analyzed to compare trophic identities among laboratory‐reared and free‐roaming detritivores. When colonized by bacteria or fungi, the trophic positions of detrital complexes increased significantly over time. The magnitude of trophic inflation was mediated by the extent of microbial consumption of detrital substrates. When detrital complexes were fed to vertebrate and invertebrate animals, the consumers registered similar degrees of trophic inflation, albeit one trophic level higher than their diets. The wild‐collected detritivore fauna in our study exhibited significantly elevated trophic positions. Our findings suggest that the trophic positions of detrital complexes rise predictably as microbes convert nonliving organic matter into living microbial biomass. Animals consuming such detrital complexes exhibit similar trophic inflation, directly attributable to the assimilation of microbe‐derived amino acids. Our data demonstrate that detritivorous microbes elevate metazoan trophic position, suggesting that detritivory among animals is, functionally, omnivory. By quantifying the impacts of microbivory on the trophic positions of detritivorous animals and then tracking how these effects propagate “up” food chains, we reveal the degree to which microbes influence consumer groups within trophic hierarchies. The trophic inflation observed among our field‐collected fauna further suggests that microbial proteins represent an immense contribution to metazoan biomass. Collectively, these findings provide an empirical basis to interpret detritivore trophic identity, and further illuminate the magnitude of microbial contributions to food webs.  相似文献   

7.
Several studies have investigated the effect of nematodes on microbial degradation of macrophyte detritus, but little is known about the potential effect of bacterivorous nematodes on productivity of macrofaunal detritivores. We investigated the influence of the bacterivorous nematode Rhabditis (Pellioditis) mediterranea on the incorporation rate of isotopically-labelled macroalgal detritus by the amphipod Paramoera chevreuxi in a laboratory microcosm. The fatty acid composition of amphipods, nematodes, and macroalgal detritus was characterized to help determine the pathway of detritus incorporation by amphipods. The potential importance of R. mediterranea as a source of highly unsaturated fatty acids (HUFAs) to higher trophic levels was also investigated.We found no clear evidence for an effect of nematodes on the incorporation rate of fresh macroalgal detritus by amphipods, although there was some indication that the type of detritus (i.e. the green Chaetomorpha sp. vs the red Polysiphonia sp.) is important in determining the nature and extent of the relationship between nematodes and macrofaunal detritivores. Fatty acid data indicated that nematodes did not contribute significantly to the diet of amphipods when detritus was present, and there was no evidence that nematodes affected the pathway of detritus incorporation by amphipods. Amphipods incorporated Chaetomorpha sp. detritus about 10 times faster than Polysiphonia sp. detritus despite the higher C/N ratio and lower HUFA content of the former. This suggests that other factors, such as the presence of grazer deterrent compounds, are important in determining the palatability of macroalgal detritus. Amphipods fed exclusively on nematodes retained high HUFA levels but suffered high mortality. The burrowing behaviour of nematodes is suggested as the most likely factor limiting their availability to predators.Results suggest a limited interaction between amphipods and bacterivorous nematodes in detrital food webs. Further experiments are needed to test the wider applicability of these findings to different nematode and macrofaunal taxa, and for different types of detritus.  相似文献   

8.
9.
Yang LH 《Oecologia》2006,147(3):522-532
Detritivore communities influence the decomposition of detrital resources in virtually all natural systems. Conversely, detrital resources can also have considerable bottom-up effects on detritivore communities. While many investigations have examined detritivory and decomposition processes, few have considered interactions between detritivores and detritus as concurrent processes in the same system, or in the context of natural detrital pulses. In many systems, resource pulses contribute substantial detrital inputs to belowground systems. These detrital pulses may influence interactions between the detritivore community and detrital decomposition. I conducted field experiments to investigate interactions between detrital resource pulses of periodical cicada (Magicicada spp.) carcasses and scavenging detritivorous macroarthropods. Cicada litterfall pulses influenced several broad groups in the macroarthropod community, including relatively specialized necrophilous taxa and relatively generalized detritivores, omnivores and predators. Conversely, detritivore activity increased the rate of cicada carcass decomposition by 4,082% compared to caged control carcasses. These results suggest that interactions between pulses of cicada detritus and the detritivore community influence both the persistence of ephemeral detrital resources, and the distribution, abundance and behavior of detritivore populations.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.  相似文献   

10.
In different ecosystems herbivores highly prefer particular plant species. This is often explained in a stoichiometric framework of nutrient‐based plant adaptations to herbivory. We hypothesize that such super‐palatability can also arise as an evolutionary by‐product of osmoregulatory adaptations of plants to stressful environmental conditions, as salinity, drought and cold. Here, we investigate in a coastal salt marsh why some plant species are highly preferred by migratory brent geese Branta bernicla bernicla in spring while others are avoided. This salt marsh is an important spring staging site for the geese. Sufficient energy storage in a short period is critical to enable their northward migration to Siberia and subsequent reproduction. We test if geese prefer plants that balance their internal osmotic potential with the saline environment through energy‐rich soluble sugars over plant species that use (compartmentalized) salts for this. We find that plant nitrogen and acid detergent fiber content, classic predictors of herbivore preferences, poorly explain which plants the geese prefer. Instead, plant species that are highly preferred by the geese adapt to salinity by high soluble sugar concentrations while avoided species do this by high plant salt concentrations. Thus, the type of osmoregulatory adaptation to stress displayed by different plant species is a good predictor for the food preference of geese on this salt marsh. We suggest that variation in other types of osmoregulation‐based stress adaptations, as plant cold adaptations in tundras and plant drought adaptations in savannas, have similar important consequences for trophic interactions.  相似文献   

11.
《Global Change Biology》2018,24(7):3170-3186
Warming is among the major drivers of changes in biotic interactions and, in turn, ecosystem functioning. The decomposition process occurs in a chain of facilitative interactions between detritivores and microorganisms. It remains unclear, however, what effect warming may have on the interrelations between detritivores and microorganisms, and the consequences for the functioning of natural freshwater ecosystems. To address these gaps, we performed a field experiment using tank bromeliads and their associated aquatic fauna. We manipulated the presence of bacteria and detritivorous macroinvertebrates (control, “bacteria,” and “bacteria + macroinvertebrates”) under ambient and warming scenarios, and analyzed the effects on the microorganisms and ecosystem functioning (detritus mass loss, colored dissolved organic matter, and nitrogen flux). We applied antibiotic solution to eliminate or reduce bacteria from control bromeliads. After 60 days incubation, bacterial density was higher in the presence than in the absence of macroinvertebrates. In the absence of macroinvertebrates, temperature did not influence bacterial density. However, in the presence of macroinvertebrates, bacterial density decreased by 54% with warming. The magnitude of the effects of organisms on ecosystem functioning was higher in the combined presence of bacteria and macroinvertebrates. However, warming reduced the overall positive effects of detritivores on bacterial density, which in turn, cascaded down to ecosystem functioning by decreasing decomposition and nitrogen flux. These results show the existence of facilitative mechanisms between bacteria and detritivores in the decomposition process, which might collapse due to warming. Detritivores seem to contribute to nutrient cycling as they facilitate bacterial populations, probably by increasing nutrient input (feces) in the ecosystem. However, increased temperature mitigated these beneficial effects. Our results add to a growing research body that shows that warming can affect the structure of aquatic communities, and highlight the importance of considering the interactive effects between facilitation and climatic drivers on the functioning of freshwater ecosystems.  相似文献   

12.
Summary The possible interaction of high soil salinity and low soil nitrogen content in affecting the growth of Spartina alterniflora Loisel in the high and low marshes of the Eastern U.S. was explored. Throughout the whole growing season, the short plants growing in the high marsh, where there was a higher soil salinity and lower available soil nitrogen, contained more proline and glycinebetaine and showed a lower leaf water potential than the tall plants growing in the low marsh. In both short and tall plants, the growing season, with the highest content occurring in spring and fall. In contrast, the glycinebetaine content in both short and tall plants remained fairly constant throughout the growing season, and was consistently at least 10 fold higher than the proline content. It is estimated that 19–30% of the total leaf nitrogen was in the form of proline and glycinebetaine in the short plants, and 14–27% in the tall plants. Ammonium nitrate fertilization in the field resulted in increased growth, higher proline and glycinebetaine contents, and lower water potentials in the short plants, but had little effect on these parameters in the tall plants. We suggest that in the low marsh, the plants can obtain sufficient nitrogen for osmoregulation and other metabolism. In the high marsh with higher soil salinity and lower nitrogen content, the plants have to allocate a even greater proportion of the already limited nitrogen supply for osmoregulation. Thus, nitrogen available for osmoregulation and other nitrogen-requiring metabolism is insufficient, resulting in reduced growth.  相似文献   

13.
1. By selective grazing, invasive grazers can alter macrophyte‐herbivore relationships in shallow freshwater bodies. Evaluating the palatability of macrophytes and understanding the determinants of plant palatability can help predict grazing impact. In no‐choice feeding assays, we tested the palatability of 21 species of freshwater macrophytes to the invasive freshwater apple snail Pomacea canaliculata. 2. Daily feeding rate varied greatly with plant species, ranging from 1.1 to 22% of snail body mass. We assessed six plant properties and examined their correlation with feeding rate. Total nitrogen content was positively related, and C:N ratio and dry matter content (DMC) negatively related, to snail feeding rate. There was no significant correlation between snail feeding rate and plant phenolic content, but the feeding rate on Myriophyllum aquaticum (the plant with the highest phenolic content) was very low. 3. We repeated the feeding assays for 15 species that were not palatable as fresh leaves with reconstituted plant tissues formed by mixing ground up dried leaves with agar. The feeding rate still differed greatly among macrophyte species. Phragmites australis and Vallisneria natans (two species with the highest DMC) were eaten much more as reconstituted plant than as fresh leaves, indicating that structure (i.e. DMC) may be important in their defence against snail herbivory. For two plants (M. aquaticum and Alternanthera philoxeroides) that had moderate amounts of nitrogen/phosphorus but were consumed very little as fresh and reconstituted tissues, we incorporated their extracts into a palatable agar‐based food. The extracts from both species greatly reduced snail feeding rate, indicating the presence of chemical defences in these two species. 4. These results indicated that feeding was affected by several plant traits. The snail favoured plants with a high nitrogen content and avoided plants with a high DMC. Only a few plants possessed chemical feeding deterrents that were effective against this snail. Given the invasive spread of P. canaliculata in Asia, ecologists and managers should consider plant palatability when selecting plants for use in wetland restoration and when predicting the impact of further invasion by this species.  相似文献   

14.
Food assimilation by the detritivorous fish Prochilodus lineatus, throughout a life cycle, was studied by means of fatty acid analysis. Fish length ranged from 2.7 to 55 cm and weight from 2 to 2670 g. Samples for fatty acid determination were taken from the mesenteric deposit in adults and from the whole fish in juveniles. Phyto and zooplankton, and superficial mud were also studied. Samples were taken from the principal channel of the Paraná River and lentic environments in its alluvial plain (60 ° 29W and 31 ° 40S). Fatty acids in the smaller fish (2.8–3.8 cm) indicated the assimilation of zoo and phytoplankton. Longer P. lineatus have a fat-deficient diet, particularly in polyunsaturated acids. At different stages of life, P. lineatus is adapted to different sources of food. When young, it eats zoo- and phytoplankton, becomes gradually detritivorous and has a series of adaptations of the digestive tract (morphological and histological) to enable it to assimilate detritus.  相似文献   

15.
We examined geographic variation in the structure and function of salt marsh communities along the Atlantic and Gulf coasts of the United States. Focusing on the arthropod community in the dominant salt marsh plant Spartina alterniflora, we tested two hypotheses: first, that marsh community structure varies geographically, and second, that two aspects of marsh function (response to eutrophication and addition of dead plant material) also vary geographically. We worked at eleven sites on the Gulf Coast and eleven sites on the Atlantic Coast, dividing each coast up into two geographic areas. Abiotic conditions (tidal range, soil organic content, and water content, but not soil salinity), plant variables (Spartina nitrogen content, height, cover of dead plant material, but not live Spartina percent cover or light interception), and arthropod variables (proportional abundances of predators, sucking herbivores, stem-boring herbivores, parasitoids, and detritivores, but not total arthropod numbers) varied among the four geographic regions. Latitude and mean tidal range explained much of this geographic variation. Nutrient enrichment increased all arthropod functional groups in the community, consistent with previous experimental results, and had similar effects in all geographic regions, contrary to our hypothesis, suggesting widespread consistency in this aspect of ecosystem function. The addition of dead plant material had surprisingly little effect on the arthropod community. Our results caution against the uncritical extrapolation of work done in one geographic region to another, but indicate that some aspects of marsh function may operate in similar ways in different geographic regions, despite spatial variation in community structure.  相似文献   

16.
Combtooth blennies (Blenniidae) have a wide array of diets, including coral polyps, fish scales/mucus/rays, and detritus. However, there is limited knowledge about the diet of some species. We described the diet of 30 blenny species from subtidal, intertidal, or supralittoral habitats in Kochi (warm temperate) and Okinawa (tropical), Japan. Cluster analysis of dietary overlap grouped blennies into seven feeding groups: omnivores, herbivores, detritivores, molluscivores, corallivores, worm-like invertebrate feeders, and fish mucus/scale/ray feeders. The largest cluster, detritivores, contained 17 blenny species from all habitat zones and both climate zones. Our findings suggest that closely related species fill similar feeding niches and detritus is an important component of many blenny diets.  相似文献   

17.
Goranson CE  Ho CK  Pennings SC 《Oecologia》2004,140(4):591-600
Current theories of plant-herbivore interactions suggest that plants may differ in palatability to herbivores as a function of abiotic stress; however, studies of these theories have produced mixed results. We compared the palatability of eight common salt marsh plants that occur across elevational and salinity stress gradients to six common leaf-chewing herbivores to determine patterns of plant palatability. The palatability of every plant species varied across gradients of abiotic stress in at least one comparison, and over half of the comparisons indicated significant differences in palatability. The direction of the preferences, however, was dependent on the plant and herbivore species studied, suggesting that different types of stress affect plants in different ways, that different plant species respond differently to stress, and that different herbivore species measure plant quality in different ways. Overall, 51% of the variation in the strength of the feeding preferences could be explained by a knowledge of the strength of the stress gradient and the type of gradient, plant and herbivore studied. This suggests that the prospects are good for a more complex, conditional theory of plant stress and herbivore feeding preferences that is based on a mechanistic understanding of plant physiology and the factors underlying herbivore feeding preferences.  相似文献   

18.
In Tissawewa, a shallow, eutrophic reservoir in southeastern Sri Lanka, the effect of a major drought on the ecosystem was studied by monitoring the size-structured fish community and its resource base. Primary production was determined as well as the production and diets of ten taxa belonging to four trophic guilds (i.e. herbivorous/detritivorous, benthivorous, zooplanktivorous/insectivorous, piscivorous) that made up more than 98% of the total fish biomass. Two extreme states of the ecosystem were distinguished. Before the drought most primary production was generated byphytoplankton, suspended fine particulate detritus was an important food source and total fish density was high. After the drought the ecosystem was characterised by high macrophyte density, low concentration of suspended detritus and low total fish density. The availability and origin of detritus appeared to be the major factor influencing fish production in Tissawewa. The small pelagic herbivore/detritivore A. melettinus contributed the most biomass and production to the fish community before the drought. After the drought, however, biomass and production dropped considerably. In contrast, the production of the most important species in terms of fisheries yield, the exotic herbivorous/detritivorous tilapias, was hardly affected. Although the composition of their food, benthic detritus, had markedly changed. In Sri Lankan reservoirs a subsidiary fishery for pelagic minor cyprinids was suggested to increase the current yield which is based almost entirely on the exotic tilapia species. The perturbation observed in this study, however, showed that the production of pelagic species was affected particularly by the environmental changes. Exploitation of these species can, therefore, only be considered in combination with hydrological and other management measures that control the environmental conditions of the reservoirs. This revised version was published online in July 2006 with corrections to the Cover Date.  相似文献   

19.
Restoration of salt marsh ecosystems is an important concern in the eastern United States to mitigate damage caused by industrial development. Little attention has been directed to the mycorrhizal influence on plantings of salt marsh species to stabilize estuarine sediments and establish cover. In our study, seedlings of two salt marsh grasses, Spartina alterniflora and Spartina cynosuroides, were grown in soil with a commercial, mixed species inoculum of arbuscular mycorrhizal fungi. Plants were grown in experimental “ebb and flow” boxes, simulating three levels of tidal inundation, to which two levels of applied phosphorus (P) and two levels of salinity were imposed. After 2.5 months, S. alterniflora was poorly colonized by arbuscular mycorrhizae, developing only fungal hyphae and no arbuscules, but S. cynosuroides became moderately colonized. Mycorrhizal inoculation marginally improved growth and P and nitrogen (N) content of both plant species at low levels of P supply but significantly increased tillering in both plant species. This factor could be beneficial in enhancing ground cover during restoration procedures. Greater P availability increased the mycorrhizal status of S. cynosuroides and improved P nutrition of both plant species, despite a reduction in the root‐to‐shoot ratio. Increasing salinity reduced mycorrhizal colonization of S. alterniflora but not of S. cynosuroides. Growth and nutrient content of S. alterniflora was improved at higher levels of salinity, but only increased nutrient content in S. cynosuroides. Increased duration of tidal inundation decreased plant growth in both species, but tissue P and N concentrations were highest with the longest time of inundation in both species.  相似文献   

20.
Biomass, primary production and nutrient budgets associated to Sarcocornia perennis subspecies (ssp.) alpini were studied in the Palmones River estuary salt marsh (Southern Spain) to evaluate the nutrient sequestration capacity of the low marsh. Above- and belowground living and dead biomass, as well as carbon, nitrogen and phosphorus content were monitored during 1 year. Additionally, the fate of aboveground detritus was evaluated in an experiment on litter decomposition. The detritus production of S. perennis ssp. alpini was almost equivalent to its annual primary production indicating a rapid turnover of biomass. We calculated that only 12% of the aboveground detritus was exported out of the low marsh while the rest was decomposed in the sediment with a rate of 0.8 year−1. Changes in concentrations of total carbon, nitrogen and phosphorus in the sediment showed patterns related to S. perennis ssp. alpini belowground biomass. Our results suggested that the sediment functions as a net sink for nutrients accumulating 550 g C m−2 year−1, 55 g N m−2 year−1, and 13 g P m−2 year−1.  相似文献   

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