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1.
Changes in assemblages of plants, macroinvertebrates and fishes on three eastern Tasmanian reefs were monitored over 12 months in replicated control blocks and adjacent 10×12-m blocks cleared of fucoid, laminarian and dictyotalean algae. Removal of canopy-forming plants produced less change to biotic assemblages than reported in studies elsewhere, with the magnitude of change for fish and invertebrate taxa lower than variation between sites and comparable to variation between months.The introduced annual kelp Undaria pinnatifida exhibited the only pronounced response to canopy removal amongst algal taxa, with a fivefold increase in cleared blocks compared to control blocks. Marine reserves are suggested to assist reef communities resist invasion by U. pinnatifida, through an indirect mechanism involving increased predation pressure on sea urchins and reduced formation of urchin barrens that are amenable to U. pinnatifida propagation.Large invertebrates were more associated with turfing algae or the reef substratum than the macroalgal canopy. The herbivorous sea urchin Heliocidaris erythrogramma and abalone Haliotis ruber showed the strongest response to clearing amongst common macroinvertebrate species, with a halving of population numbers. Observed densities of the common monacanthid fish Acanthaluteres vittiger also declined by about 50%. The relatively high level of resistance shown by eastern Tasmanian reef biota to patch disturbance was attributed largely to high diversity and biomass of turfing macroalgae damping effects of canopy clearance.  相似文献   

2.
Herbivory can be an important factor structuring coastal algal communities. Herbivores may preferentially graze particular algal species or tissue types. Mesograzers, despite their small size, can critically weaken kelp thalli and impact entire kelp beds. We propose that when kelp beds are composed of several kelp cohorts, mesograzers will selectively choose to inhabit younger plants and grazing activities will have a greater impact on younger plants. This study investigated the effects of grazing by the littorinid gastropod, Lacuna vincta, on different age classes of the bull kelp, Nereocystis luetkeana by (1) testing food preference of L. vincta on juvenile, first-year adult, and second-year adult Nereocystis blades in the laboratory, (2) determining substrate (blades of different ages) preference of L. vincta in the laboratory, and by (3) estimating in-situ herbivore abundances and densities on juvenile and adult Nereocystis. Results demonstrated that grazing by L. vincta produced greater damage on juvenile than older Nereocystis tissues. Although L. vincta did not select juvenile versus older kelps as substrate in the laboratory, in situ surveys showed that differences existed between age classes with higher L. vincta densities on juvenile than adult kelp. We conclude that at a local scale, L. vincta can be an important structuring factor in Nereocystis populations due to its high density and grazing ability. Handling editor: K. Martens  相似文献   

3.
Despite recent rapid increases in the occurrence of nonindigenous marine organisms in the marine environment, few studies have critically examined the invasion process for a marine species. Here we use manipulative experiments to examine processes of invasion for the Asian kelp Undaria pinnatifida (Harvey) Suringar at two sites on the east coast of Tasmania. Disturbance to reduce cover of the native algal canopy was found to be critical in the establishment of U. pinnatifida, while the presence of a stable native algal canopy inhibited invasion. In the first sporophyte growth season following disturbance of the canopy, U. pinnatifida recruited in high densities (up to 19 plants m−2) while remaining rare or absent in un-manipulated plots. The timing of disturbance was also important. U. pinnatifida recruited in higher densities in plots where the native canopy was removed immediately prior to the sporophyte growth season (winter 2000), compared with plots where the canopy was removed 6 months earlier during the period of spore release (spring 1999). Removal of the native canopy also resulted in a significant increase in cover of sediment on the substratum. In the second year following canopy removal, U. pinnatifida abundance declined significantly, associated with a substantial recovery of native canopy-forming species. A feature of the recovery of the native algal canopy was a significant shift in species composition. Species dominant prior to canopy removal showed little if any signs of recovery. The recovery was instead dominated by canopy-forming species that were either rare or absent in the study areas prior to manipulation of the canopy.  相似文献   

4.

Aim

In marine ecosystems, habitat‐forming species (HFS) such as reef‐building corals and canopy‐forming macroalgae alter local environmental conditions and can promote biodiversity by providing biogenic living space for a vast array of associated organisms. We examined community‐level impacts of observed climate‐driven shifts in the relative abundances of two superficially similar HFS, the warm‐water kelp Laminaria ochroleuca and the cool‐water kelp Laminaria hyperborea.

Location

Western English Channel, north‐east Atlantic

Methods

We compared algal and invertebrate assemblages associated with kelp stipes and holdfasts, across multiple sites and sampling events. Significant differences were recorded in the structure of assemblages between the host kelp species at each site and event.

Results

Assemblages associated with stipes of the cool‐water HFS were, on average, >12 times more diverse and supported >3600 times more biomass compared with the warm‐water HFS. Holdfast assemblages also differed significantly between species, although to a lesser extent than those associated with stipes. Overall, assemblages associated with the warm‐water HFS were markedly impoverished and comprised far fewer rare or unique taxa.

Main conclusions

While previous research has shown how climate‐driven loss of HFS can cause biodiversity loss, our study demonstrates that climate‐driven substitutions of HFS can also lead to impoverished assemblages. The indirect effects of climate change remain poorly resolved, but shifts in the distributions and abundances of HFS may invoke widespread ecological change, especially in marine ecosystems where facilitative interactions are particularly strong.  相似文献   

5.
Trophic support functions for fishes are a key goal of salt marsh restoration. Food availability in restored sites may be enhanced by creation of shallow pools, which are important sources of prey items in tidal wetlands. Young restored salt marshes are typically sparsely vegetated and are subject to rapidly changing geomorphology. Scouring and sedimentation create and fill shallow depressions, producing a shifting mosaic of tidal pools. In a large (8-ha) southern California experimental restoration site, we created shallow pools and assessed their development of foods for fishes. Created pools quickly developed abundant invertebrate prey, with densities exceeding those found in older, naturally formed pools (P < 0.0001). Opportunistic mobile and disturbance-associated taxa (calanoid copepods, nematodes, Polydora complex, and Trichocorixa reticulata) accounted for higher invertebrate densities in created pools. We repeated experiments in spring, summer, and fall and found seasonal variability in trophic development. We also applied bottom-up (nitrogen addition) and top-down (fish exclusion) treatments to pools. Some measures of algal biomass were increased by nitrogen fertilization (P = 0.001–0.06), but there were no upward-cascading effects on invertebrate composition or abundance. Fish abundance in the site varied seasonally, but there were no compelling effects of fish exclusion treatments on algal or invertebrate abundance. Incorporating shallow depressions into salt marsh restoration projects is a potential tool to jumpstart fish-support functions.  相似文献   

6.
7.
Understanding the impact of multiple stressors on ecosystems is of pronounced importance, particularly when one or more of those stressors is anthropogenic. Here we investigated the role of physical disturbance and increased nutrients on reefs dominated by the canopy-forming kelp Ecklonia radiata. We combined experimental kelp canopy removals and additional nutrient at three different locations in a large embayment in temperate southeastern Australia. Over the following winter recruitment season, Ecklonia recruitment was unaffected by increased nutrients alone, but tripled at all sites where the canopy had been removed. At one site, the combination of disturbance and increased nutrients resulted in more than four times the recruitment of the introduced kelp Undaria pinnatifida. Six months after disturbance, the proliferation of the Undaria canopy in the canopy-removal and nutrient-addition treatment negatively influenced the recovery of the native kelp Ecklonia. Given the otherwise competitive dominance of adult Ecklonia, this provides a mechanism whereby Undaria could maintain open space for the following recruitment season. This interplay between disturbance, nutrients and the response of native and invasive species makes a compelling case for how a combination of factors can influence species dynamics.  相似文献   

8.
The association of the geoduck Panopea abbreviata and the green alga Coccomyxa parasitica is described. The identity of the green alga was confirmed by molecular studies; the alga was found within the hemocytes that infiltrate the connective tissue of the geoduck siphons. Cytological characteristics of hemocytes were not altered by algal infection; very often the algae were seen enveloped by a digestive vacuole within the hemocyte cytoplasm, evidencing diverse degrees of resorption. Connective cells of siphons were rarely infected by C. parasitica. The mean prevalence of C. parasitica was higher (82%) in San Matías Gulf (42°00′S, 65°05′W) than in San José Gulf (45%) (40°32′S, 64°02′W); except for spring, when the two locations showed no differences in prevalences (80%). Independently of location, season and host size, infected geoducks showed lower condition index values than uninfected ones. Regarding other bivalve species, only one specimen of the razor clam Ensis macha was found infected, and none of the oysters Ostrea puelchana and Pododesmus rudis and scallop Aequipecten tehuelchus was parasitized by the green alga.  相似文献   

9.
1. We characterised aquatic and terrestrial invertebrate drift in six south‐western North Carolina streams and their implications for trout production. Streams of this region typically have low standing stock and production of trout because of low benthic productivity. However, little is known about the contribution of terrestrial invertebrates entering drift, the factors that affect these inputs (including season, diel period and riparian cover type), or the energetic contribution of drift to trout. 2. Eight sites were sampled in streams with four riparian cover types. Drift samples were collected at sunrise, midday and sunset; and in spring, early summer, late summer and autumn. The importance of drift for trout production was assessed using literature estimates of annual benthic production in the southern Appalachians, ecotrophic coefficients and food conversion efficiencies. 3. Abundance and biomass of terrestrial invertebrate inputs and drifting aquatic larvae were typically highest in spring and early summer. Aquatic larval abundances were greater than terrestrial invertebrates during these seasons and terrestrial invertebrate biomass was greater than aquatic larval biomass in the autumn. Drift rates of aquatic larval abundance and biomass were greatest at sunset. Inputs of terrestrial invertebrate biomass were greater than aquatic larvae at midday. Terrestrial invertebrate abundances were highest in streams with open canopies and streams adjacent to pasture with limited forest canopy. 4. We estimate the combination of benthic invertebrate production and terrestrial invertebrate inputs can support 3.3–18.2 g (wet weight) m−2 year−1 of trout, which is generally lower than values considered productive [10.0–30.0 g (wet weight) m−2 year−1]. 5. Our results indicate terrestrial invertebrates can be an important energy source for trout in these streams, but trout production is still low. Any management activities that attempt to increase trout production should assess trout food resources and ensure their availability.  相似文献   

10.
Live Norway lobsters (Nephrops norvegicus L.) were trawled at depths of 30 to 55 m off the coast of Jutland (Denmark) in late winter (March) and in summer (August) in 2006. Water temperatures at the bottom and surface of the sea were 7 °C and 2 °C during the winter, and 12 °C and 21 °C in the summer, respectively. The recovery of specific physiological and metabolic variables from the intense stresses associated with capture (trawling and air-exposure during sorting) was followed in seawater at 5 °C in winter or 18 °C in summer. Recovery was compared in lobsters held individually in two different live-storage positions, either resting vertically on the tail or sitting horizontally. In winter, many animals were alive when brought on board and approximately 86% were still alive at the end of experimentation (96 h). In summer very few animals were alive when brought on board and, of these, approximately 95% were dead at 24 h. When compared with values measured in laboratory controls, the stresses of capture elicited very high haemolymph lactate contents in both seasons, although levels recovered within 24 h. Trawling also caused very high haemolymph glucose concentrations, which differed with season. In winter, haemolymph glucose was elevated for 24 h to levels significantly higher than in summer. In summer, glucose had returned to control levels by 4 h. At 4 h after trawling, haemolymph O2 status was not markedly influenced in either season, but there were significant disturbances of acid-base status. In winter, a potential metabolic lactic acidosis was compensated by a marked respiratory alkalosis, with significantly increased haemolymph pH and decreased CO2 total content and partial pressure. These effects disappeared gradually over 96 h. Summer lobsters showed combined metabolic and respiratory acidosis at 4 h, although this had recovered to control values in the small number of survivors sampled at 24 h. The capture stresses elicited very high haemolymph crustacean hyperglycaemic hormone (CHH) titres, significantly higher in summer than in winter. In winter, CHH titre had declined significantly at 24 h, whereas it exhibited a further significant increase at 24 h in summer. Live-storage position had no significant effect on survival or recovery from capture stresses in either season. The results demonstrate that Nephrops were much more stressed by trawling at high summer temperatures and had difficulty recovering from this, with pronounced negative effects on their survival, irrespective of their live-storage position.  相似文献   

11.
The understorey beneath a canopy of the kelp Ecklonia radiata often contains juvenile sporophytes of the same species. When canopy disturbance occurs, these juvenile sporophytes are exposed to new environmental conditions. If these juvenile sporophytes survive these new conditions, then they become a ready source of kelps to rapidly form a new canopy. This study investigated the potential of pre-existing juvenile sporophytes of E. radiata to survive post-disturbance conditions and contribute to the rapid formation of a new canopy. The potential of canopy recovery by recruitment of kelp from zoospores was also investigated. These processes were studied at different times during the summer and on reefs ranging in topographic complexity from simple, flat reefs to highly complex, rugose reefs. By tagging juvenile sporophytes after the adult kelp canopy was removed and monitoring them through time, it was demonstrated that most juveniles (> 50%) survived the change in conditions after canopy loss, with some juveniles going on to become members of a new canopy. Approximately 6-47% of tagged sporophytes died within 3-4 days after canopy removal possibly due to excessive photoinhibition and photostress as demonstrated by changes in photosynthetic performance (decreased alpha values) of juveniles. The potential contribution of juvenile sporophytes to the rapid formation of a new canopy appears to be dependent on the timing of canopy removal with late summer-autumn canopy loss favouring faster recovery. Topographically complex reefs had less short-term (7 days) survival of juvenile sporophytes than topographically simple reefs; however this difference was not carried through to the long-term (6 months) abundance of adult kelp in experimental clearings, which was greater on topographically complex reefs. Clearly, juvenile sporophytes in arrested development under existing canopies of the small kelp E. radiata are important for the rapid recovery of the kelp canopy once adults are lost through physical disturbance.  相似文献   

12.
Molecular analysis of green-tide-forming macroalgae in the Yellow Sea   总被引:2,自引:0,他引:2  
In the summer of 2008, free-floating green algae bloomed in the Yellow Sea. Samples were collected in a wide area (119°32′-122°00′E, 32°25′-36°49′N). We calculated the sequence divergences of nuclear ITS, chloroplast rbcL, and psbA data of free-floating samples collected from the Yellow Sea and Ulvaceae from Europe and Japan. In the ITS sequence, 19 out of the 21 Yellow Sea samples of 2008 were identical to those of a sample taken at Qingdao in 2007. A low divergence (0.2%) was found in remaining two samples. Similar evidence was shown by pairwise distances of rbcL and psbA gene sequence data, implying the uniformity of the Yellow Sea blooms in 2007 and 2008. The ITS sequence of the Yellow Sea samples differed 8.1-10.8% from free-floating Enteromorpha or Ulva reported worldwide. ITS-based molecular phylogenetic results and rbcL sequence data grouped the free-floating alga in the Yellow Sea into one clade with Enteromorpha procera, Enteromorpha linza and Enteromorpha prolifera. Furthermore, both morphological characteristics and ribotype network of the ITS sequences imply that the blooming algae in 2007 and 2008 were E. prolifera. The haplotypes of the Yellow Sea free-floating E. prolifera are closely related to those from the Japanese coast but less to European and American algae.  相似文献   

13.
The fish fauna associated with a Macrocystis pyrifera forest and the effects of the canopy removal on this fauna were studied in the Beagle Channel, Tierra del Fuego. Seasonal changes and differences in the fish communities that inhabit the canopy and the holdfast were also studied. Two patches of kelp forest separated by 200 m were selected. In one, the canopy was removed periodically while the other was used as a control. Samples were collected seasonally, from autumn 1999 to 2001. Fish fauna in the water column was sampled using trammel nets and holdfast fish fauna was sampled by removing the complete holdfast. Different assemblages of fish species were captured in the water column (surface and bottom) and in the holdfast. In the former there were mainly pelagic and benthopelagic species and the latter predominantly demersal species. The principal effect of the canopy removal was a drop in the abundance (and total weight) of Paranotothenia magellanica (doradito), the species with the strongest relationship with the canopy, principally at the surface of the treated patch probably due to a reduction in the availability of refuges. A high degree of seasonality was observed for fish species’ total weight, abundance and diversity, with higher values in summer and autumn. Taking into account our data and available data on kelp growth in the same locality, we suggest of cutting the kelp forest (preferably those farther away from the coast) once a year in winter or early spring, in order to minimize impact on the fish community.  相似文献   

14.
Kelp forests are highly productive and species‐rich benthic ecosystems in temperate regions that provide biogenic habitat for numerous associated species. Diverse epifaunal communities inhabit kelp sporophytes and are subject to variations in the physical environment and to changes experienced by the kelp habitat itself. We assessed seasonal variations in epifaunal invertebrate communities inhabiting giant kelps, Macrocystis pyrifera, and their effects on this seaweed. Six seasonal samplings were conducted over a year at an upwelling‐dominated site in northern‐central Chile where physical conditions are known to fluctuate temporally. More than 30 taxa were identified, among which peracarid crustaceans stood out in both diversity and abundance. Species richness and abundance differed among sporophyte sections (holdfast and fronds) and throughout the year. The frond community was dominated by two grazers (the amphipod Peramphithoe femorata and the isopod Amphoroidea typa), while suspension feeders, grazers, and omnivores (the amphipod Aora typica, the isopod Limnoria quadripunctata, and polychaetes) dominated the holdfasts. Abundances of the dominant species fluctuated throughout the year but patterns of variation differed among species. The most abundant grazer (P. femorata) had highest densities in summer, while the less abundant grazer (A. typa) reached its peak densities in winter. Interestingly, the area of kelp damaged by grazers was highest in autumn and early winter, suggesting that grazing impacts accumulate during periods of low kelp growth, which can thus be considered as ‘vestiges of herbivory past.’ Among the factors determining the observed seasonal patterns, strong variability of environmental conditions, reproductive cycles of associated fauna, and predation by fishes vary in importance. Our results suggest that during spring and early summer, bottom‐up processes shape the community structure of organisms inhabiting large perennial seaweeds, whereas during late summer and autumn, top‐down processes are more important.  相似文献   

15.
Kelp forests are patchy fish-associated habitats, which can vary greatly in their size, foundation species, and several physical habitat attributes. The structure of fish assemblages can vary with these characteristics and with the location of the assemblage within the forest, i.e. edges versus interiors. This study quantified the biological and physical heterogeneity within different sized kelp forests and identified which factors are important in structuring the associated fish assemblages. Fish and habitat surveys were conducted at the edge and interiors of ten kelp forests of varying sizes in Kachemak Bay, south-central Alaska. Fish assemblage structure was not correlated with the species composition of surface canopy forming foundation kelps (Nereocystis leutkeana and Eualaria fistulosa) or with kelp forest size. Instead, it correlated with the abundances of two understory foundation kelps (Agarum clathratum and Saccharina latissima), substratum rugosity, and water depth. Together these benthic attributes explained 53.6 % of the fish assemblage variability. Additionally, significantly different fish assemblages were found at edge compared to interior locations with the relative abundance of seven species (Artedius fenestralis, Ammodytes hexapterus, Blepsias cirrhosis, Gadus macrocephalus, Hexagrammos stelleri, Pholis laeta, and Sebastes melanops) explaining 91.4 % of the variability. This study highlights the importance of habitat characteristics such as understory foundation species, substratum rugosity, water depth and location within a patch on the variability of fish assemblages in high latitude kelp forests.  相似文献   

16.
Persistence of annual plant populations requires sufficient seeds and suitable habitat for development and growth each year. Competition with perennials may prevent within site persistence and result in “fugitive” annual populations. Comparisons have been made between the population biology of annual macroalgae and terrestrial plants, but demographic information necessary to make strong comparisons is lacking for most of these algae, and life history differences may make such comparisons questionable. We studied population dynamics of the kelp Alaria marginata to determine if it was an annual and, if so, how populations persisted. This kelp is the dominant macroalga on exposed mid to low rocky intertidal shores along the Big Sur coast of California. Experimental clearings at two sites were used to assess recruitment timing and survivorship. Sporophytes were collected monthly to determine growth and fecundity. Recruitment occurred in late winter to early spring, primarily on geniculate corallines and residual A. marginata holdfasts. Thinning was inversely related to density, and occurred during the February through July growing season as larger thalli rapidly increased in length (up to 1.4 m month 1) and formed a thick canopy. Sorus development was positively related to size, began as early as March, peaked in late August-October, and decreased as adults were removed by winter surf. Spore release was generally highest (108-109 spores individual 11 h 1) between October and January and associated with high water motion. Survivorship of sporophytes beyond one year was < 1%, showing the populations were annual.Field observations and experiments on effects of canopy clearing, season of clearing, and influence of substrate type on recruitment were done to assess how these annual populations persist. Massive spore production at the onset of fall storms, survival of microscopic stages for 3-4 months facilitated by microhabitat refuges, rapid growth, large size and rapid maturation of sporophytes contributed to persistence. Furthermore, the dense stands with thick canopies may suppress potential competitors via shading and abrasion. Rather than being a fugitive, this combination of growth and life history features enables A. marginata and perhaps other large, annual kelps to maintain perennial populations.  相似文献   

17.
1. Levels of ash-free dry matter (AFDM) and chlorophyll a in epilithon, benthic participate organic matter (BPOM), invertebrate assemblage composition, and biomass of functional feeding groups were compared in winter and summer at forest and pasture sites in three Victorian streams. 2. Chlorophyll a concentrations of epilithon were significantly higher at pasture than forest sites in winter but not in summer while BPOM was not significantly greater at forest sites in either season. Epilithic biomass as AFDM did not show consistent differences between land uses or seasons. 3. Total biomass of invertebrates did not differ between forest and pasture sites but the biomass of shredders was significantly higher, and that of grazers significantly lower, at forest than pasture sites. A site shaded with an artificial canopy behaved as a forest site for grazers but as a pasture site for shredders. 4. Cluster analyses of invertebrate assemblages grouped pasture sites with forest sites on the same stream at the same season, indicating that assemblage composition was less influenced by land-use differences than by between-stream and seasonal differences. 5. Biomass of functional feeding groups appeared to be a more sensitive indicator of invertebrate assemblage response to land-use alteration than either species diversity/ richness measures, or multivariate assemblage composition measures.  相似文献   

18.
The establishment and growth of Nereocystis luetkeana communities are limited by many environmental and biotic factors. Temperature, light quantity and quality, turbidity, salinity, wave motion, and competition for space and light all determine where populations of this annual kelp will establish and persist from season to season. Like much of the nearshore marine vegetation in Puget Sound and the San Juan Archipelago, Nereocystis is negatively impacted by anthropogenic disturbances, such as shoreline development and sediment loading from terrestrial erosion. These events can alter or eliminate optimal habitat and conditions that support adult populations, as well as interfere with the development of the microscopic life stages. Various techniques to establish a Nereocystis bed in the northern waters of Washington, USA are examined and will serve as a model for future restoration efforts. Techniques being investigated are as follows: 1) out‐planting Nereocystis during different seasons; 2) out‐planting at different stages of maturity, i.e., zoospore and microscopic sporophyte (0.5–1.0 mm blade length) from cultures; 3) out‐planting directly on natural substratum versus at elevated positions; 4) transplanting juveniles (10 cm stipe length); and, 5) seeding natural substratum with out‐planted sori (sporophylls), bypassing the lab culturing phase. Results of this study will help define the optimal restoration procedure, including environmental parameters, for Nereocystis in order to aid coastal managers in the implementation of nearshore restoration projects. Preliminary results will be presented.  相似文献   

19.
20.
Hemisphere scale events such as El Niño-Southern Oscillation (ENSO) can alter rainfall regimes worldwide, with important effects on species abundance and distribution. The evidence of ENSO effects on terrestrial communities is, however, restricted to a few ecosystem types. We explored the effects of ENSO episodes on plant/terrestrial-herbivore interactions through changes in the rainfall regime in a southwestern Atlantic salt marsh (Mar Chiquita coastal lagoon, Argentina. 37° 40′S, 57° 23′W). Surveys showed a positive relationship between winter rainfall and the abundance of the wild guinea pig Cavia aperea. The highest salt marsh abundances of C. aperea were associated with rainy periods during El Niño episodes, and the lowest ones were associated with the driest La Niña episodes. Rainfall was negatively associated with marsh sediment salinity, and experiments revealed that increased salinity reduces growth and increases mortality of cordgrass (Spartina densiflora). Salt increase also causes the highest percentage of dry area in S. densiflora leaves and reduced carbon content, and more salt content and secretion in S. densiflora stems. A factorial experiment in which we manipulated C. aperea presence and salinity along the edges of S. densiflora patches showed that plants can asexually invade unvegetated areas when salinity is reduced and C. aperea is excluded. Conversely, S. densiflora edges retracted when salinity was increased or there was C. aperea herbivory. Changes in nutritional quality of S. densiflora could explain the low herbivory of (and lack of impacts from) C. aperea in plots with high salinity. Thus, plant distribution responds directly to climate oscillations through changes in salt stress, and indirectly, through changes in plant-herbivore interactions. Herbivores respond indirectly to climate oscillations through changes in plant food quality, which suggests that top-down effects increase when bottom-up stressors are relaxed. ENSO events have direct and indirect effects on marsh communities that modulate the relative importance of top-down and bottom-up effects and have a considerable effect on the primary productivity of S. densiflora marshes.  相似文献   

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