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1.
In diving seabirds, sexual dimorphism in size often results in sex-related differences of foraging patterns. Previous research on Magellanic penguins, conducted during the breeding season, failed to reveal consistent differences between the sexes on foraging behavior, despite sexual dimorphism. In this paper, we tested the hypothesis that male and female Magellanic penguins differ in diet and foraging patterns during the non-breeding period when the constraints imposed by chick rearing activities vanish. We used stable isotope ratios of carbon and nitrogen in feather and bone to characterize the diet and foraging patterns of male and female penguins in the South Atlantic at the beginning of the 2009–2010 and 2010–2011 post-breeding seasons (feathers) and over several consecutive breeding and migratory seasons (bone). The mean δ13C and δ15N values of feathers showed no differences between the sexes in any of the three regions considered or in the diet composition between the sexes from identical breeding regions; however, Bayesian ellipses showed a higher isotopic niche width in males at the beginning of the post-breeding season. Stable isotope ratios in bone revealed the enrichment of males with δ13C compared with females across the three regions considered. Furthermore, the Bayesian ellipses were larger for males and encompassed those of females in two of the three regions analyzed. These results suggest a differential use of winter resources between the sexes, with males typically showing a larger diversity of foraging/migratory strategies. The results also show that dietary differences between male and female Magellanic penguins may occur once the constraints imposed by chick rearing activities cease at the beginning of the post-breeding season.  相似文献   

2.
Pelagic seabird populations can use several discrete wintering areas, but it is unknown if individuals use the same wintering area year after year. This would have consequences for their population genetic structure and conservation. We here study the faithfulness of individuals to a moulting area within and among years in a small pelagic seabird, the Thin-billed prion, which moult their primary feathers during the early part of the non-breeding period. According to stable carbon isotope ratios (δ13C) of these feathers, 90% of Thin-billed prions moult in Antarctic and 10% in South American waters. Repeated samples from individuals in 2 or 3 years indicated that several birds changed between Antarctic and South American moulting areas or vice versa. However, individuals moulting in an area in one year were more likely to do so again. Four out of five adults maintained highly conserved δ13C over the extended moulting period. One bird, however, had systematic changes in δ13C indicating latitudinal movements between the two areas during moult. Thus, the present data show that this seabird species has a highly flexible migratory strategy, not only at the population level, but also at the individual level, enabling these seabirds to exploit a highly unpredictable environment.  相似文献   

3.
We examined the isotopic signatures (δ13C, δ15N) of adult body feathers from southern giant petrels Macronectes giganteus collected at two breeding colonies in Antarctica (Potter Peninsula and Cape Geddes) and one in southern Patagonia (Observatorio Island), as well as in whole blood collected from adults of both sexes at each Antarctic colonies and from chicks at Potter Peninsula. As body feather moult is a continuous process in giant petrels, feathers provide an integrated annual signal of an adult’s diets and foraging habitats. In contrast, the stable isotope values of adult and chick blood are reflective of their diets during the breeding season. We found that sex-specific dietary segregation in adults breeding in Antarctica was notable during the breeding season (blood samples) but absent when examined across the entire year (feather samples). In addition, blood stable isotope values differed between chicks and adults, indicating that adults provision their offspring with a relatively higher amount of penguin and seal prey that what they consume themselves. This finding confirms previous work that suggests that chicks are preferentially fed with prey of presumably higher nutritional value such as carrion. Finally, based on isotopic differences between major oceanographic zones in the Southern Ocean, our data indicate population-specific differences in foraging distribution, with Antarctic populations move seasonally between Antarctic and subantarctic zones, while Patagonian populations likely forage in subtropical waters and in continental shelf habitats year-round.  相似文献   

4.
Carbon and nitrogen stable isotope ratios of Japanese anchovy (Engraulis japonicus) and their stomach contents were examined and compared among various regions around Japan. Geographical variations in the isotope ratios were found between inshore and Pacific offshore regions. While most of the anchovy samples had isotope ratios around −17.6‰ for δ13C and 10.0‰ for δ15N as median values, higher (more enriched) isotope values were found in the anchovy sampled from inshore regions. On the contrary, lower (more depleted) values were found mostly in the anchovy from the Pacific offshore region including the Kuroshio Extension and Kuroshio-Oyashio transition zones. Higher carbon isotope ratios in the inshore regions may reflect a carbon source from benthic primary producers in addition to phytoplankton possibly through the consumption of the larvae of benthic organisms such as bivalves or decapods, which were found in the stomach contents of the inshore anchovy. Variations in the nitrogen isotope ratio may reflect not only differences in the trophic level of prey species, but also variations in the baseline level of food webs. Stable isotope ratios are potentially a useful tool for understanding the stock/population structure and migration of anchovy. The present findings indicate the potential importance of the “inshore–offshore” variations in the biology of Japanese anchovy populations in the northwestern Pacific waters.  相似文献   

5.
Longline fisheries, oil spills, and offshore wind farms are some of the major threats increasing seabird mortality at sea, but the impact of these threats on specific populations has been difficult to determine so far. We tested the use of molecular markers, morphometric measures, and stable isotope (delta15N and delta13C) and trace element concentrations in the first primary feather (grown at the end of the breeding period) to assign the geographic origin of Calonectris shearwaters. Overall, we sampled birds from three taxa: 13 Mediterranean Cory's Shearwater (Calonectris diomedea diomedea) breeding sites, 10 Atlantic Cory's Shearwater (Calonectris diomedea borealis) breeding sites, and one Cape Verde Shearwater (C. edwardsii) breeding site. Assignment rates were investigated at three spatial scales: breeding colony, breeding archipelago, and taxa levels. Genetic analyses based on the mitochondrial control region (198 birds from 21 breeding colonies) correctly assigned 100% of birds to the three main taxa but failed in detecting geographic structuring at lower scales. Discriminant analyses based on trace elements composition achieved the best rate of correct assignment to colony (77.5%). Body measurements or stable isotopes mainly succeeded in assigning individuals among taxa (87.9% and 89.9%, respectively) but failed at the colony level (27.1% and 38.0%, respectively). Combining all three approaches (morphometrics, isotopes, and trace elements on 186 birds from 15 breeding colonies) substantially improved correct classifications (86.0%, 90.7%, and 100% among colonies, archipelagos, and taxa, respectively). Validations using two independent data sets and jackknife cross-validation confirmed the robustness of the combined approach in the colony assignment (62.5%, 58.8%, and 69.8% for each validation test, respectively). A preliminary application of the discriminant model based on stable isotope delta15N and delta13C values and trace elements (219 birds from 17 breeding sites) showed that 41 Cory's Shearwaters caught by western Mediterranean long-liners came mainly from breeding colonies in Menorca (48.8%), Ibiza (14.6%), and Crete (31.7%). Our findings show that combining analyses of trace elements and stable isotopes on feathers can achieve high rates of correct geographic assignment of birds in the marine environment, opening new prospects for the study of seabird mortality at sea.  相似文献   

6.
The use of stable-hydrogen isotopes (deltaD) has become a common tool for estimating geographic patterns of movement in migratory animals. This method relies on broad and relatively predictable geographic patterning in deltaD values of precipitation, but these patterns are not estimated without error. In addition, deltaD measurements are relatively imprecise, particularly for organic tissue. Most models for estimating geographic locations have ignored these sources of error. Common modeling approaches include regression, range-matching, and likelihood-based assignment tests (including discriminant analysis). Here, we show the benefits of a simple stochastic extension to likelihood-based assignment tests that incorporates two estimable sources of error and describe the resulting influence on the certainty of assigning breeding origins for wintering American Redstarts (Setophaga ruticilla), a small Nearctic-Neotropical migratory bird. Through simulation, we incorporated both spatial interpolation error associated with models of deltaD in precipitation and analytical error associated with the measurement of deltaD in tissue samples. In general, assignments that did not include these sources of error fell within the ranges of the stochastic results, but the difference in proportion of birds assigned to any one breeding region varied by as much as 54%. To explore how the distribution of assignments generated from error models influenced the application of these results, we developed a simple model of winter habitat loss. We removed the proportion of Redstarts wintering at a particular site from the global population and then used the isotope-based assignments to predict the resulting population declines for each breeding region. This gave distributions of change in population sizes, some of which included no change or even a population increase. The sources of error we modeled may challenge the degree of certainty in the use of stable-isotope-based data on connectivity to predict population dynamics of migratory animals. We suggest that stronger inference will result from incorporating these sources of error into future studies that use deltaD or other stable isotopes to infer the geographic origin of individuals.  相似文献   

7.
Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ13C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ15N values. Feather δ13C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.  相似文献   

8.
Jennings  S.  Warr  K. J. 《Marine Biology》2003,142(6):1131-1140
Marine Biology - Nitrogen stable isotopes can be used to estimate the trophic position of consumers in food webs. However, the nitrogen stable isotope ratios (δ15N) of primary producers at the...  相似文献   

9.
To aid in the management and conservation of Southwestern Willow Flycatcher (Empidonax traillii extimus, hereafter “Flycatcher”), we developed numerous models of flycatcher breeding habitat at Roosevelt Lake, AZ. For model development and testing, we compiled 10 years of flycatcher territory data that were obtained from intensive fieldwork between 1996 and 2005. We identified riparian vegetation annually in the project area from Landsat Thematic Mapper images, and extracted floodplain features from a digital elevation model. We created a novel class of temporal (i.e., multiyear) variables by characterizing the stability and variability in breeding habitat over a 6-year time interval. We used logistic regression to determine associations between environmental variables and flycatcher territory occurrence, and to test specific hypotheses. We mapped the probability of territory occurrence with a GIS and determined model accuracies with a classification table and a 10-year population database. Environmental features that were associated with breeding flycatchers included floodplain size, proximity to water, and the density, heterogeneity, age and stability of riparian vegetation. Our best model explained 79% of the variability in the flycatcher breeding population at Roosevelt Lake. The majority of predicted flycatcher habitat formed between 1996 and 2004 on an exposed lakebed ~3 years after water levels receded during a prolonged drought. A high correlation between annual reservoir levels and predicted breeding habitat (r = ?0.82) indicates that we can create and manage habitat for conservation purposes. Our predictive models quantify and assess the relative quality of flycatcher breeding habitat remotely, and can be used to evaluate the effectiveness of habitat restoration activities. Numerous techniques we developed can be used to characterize riparian vegetation and patch dynamics directly off of satellite imagery, thereby increasing its utility for conservation purposes.  相似文献   

10.
Climate in low-latitude wintering areas may influence temperate and high-latitude breeding populations of birds, but demonstrations of such relationships have been rare because of difficulties in linking wintering with breeding populations. We used long-term aerial surveys in Mexican wintering areas and breeding areas in Alaska, USA, to assess numbers of Black Brant (Branta bernicla nigricans; hereafter brant) on their principal wintering and breeding area in El Ni?o and non-El Ni?o years. We used Pollock's robust design to directly estimate probability of breeding and apparent annual survival of individually marked brant at the Tutakoke River (TR) colony, Alaska, in each year between 1988 and 2001. Fewer brant wintered in Mexico during every El Ni?o event since 1965. Fewer brant were observed on the principal breeding area following each El Ni?o since surveys began in 1985. Probability of breeding was negatively related to January sea surface temperature along the subtropical coast of North America during the preceding winter. Between 23% (five-year-olds or older) and 30% (three-year-olds) fewer brant nested in 1998 following the strong El Ni?o event in the winter of 1997-1998 than in non-El Ni?o years. This finding is consistent with life history theory, which predicts that longer-lived species preserve adult survival at the expense of reproduction. Oceanographic conditions off Baja California, apparently by their effect on Zostera marina (eelgrass), strongly influence winter distribution of brant geese and their reproduction (but not survival), which in turn affects ecosystem dynamics in Alaska.  相似文献   

11.
Stable-isotope profiles of feathers can reveal the location or habitat used by individual birds during the molting period. Heterogeneity in isotope profiles will reflect heterogeneity in molt locations, but also heterogeneity in breeding locations, because spatial heterogeneity in molt locations will be congruent with spatial heterogeneity in breeding locations in species with high connectivity between breeding and molting sites. We used information on the congruence of spatial heterogeneity in molt and breeding location to study population processes in Barn Swallows (Hirundo rustica) from a region. near Chernobyl, Ukraine, that has been radioactively contaminated since 1986; from an uncontaminated control region near Kanev, Ukraine; and from a sample of pre-1986 museum specimens used to investigate patterns prior to the nuclear disaster at Chernobyl, from both regions. Previous studies have revealed severe reductions in Barn Swallow reproductive performance and adult survival in the Chernobyl region, implying that the population is a sink and unable to sustain itself. Female Barn Swallows are known to disperse farther from their natal site than males, implying that female stable-isotope profiles should tend to be more variable than profiles of males. However, if the Barn Swallows breeding at Chernobyl are not self-sustaining, we would expect males there also to originate from a larger area than males from the control region. We found evidence that the sample of adult Barn Swallows from the Chernobyl region was more isotopically heterogeneous than the control sample, as evidenced from a significant correlation between feather sigma13C and sigma15N values in the control region, but not in the Chernobyl region. Furthermore, we found a significant difference in feather sigma15N values between regions and periods (before and after 1986). When we compared the variances in sigma13C values of feathers, we found that variances in both sexes from post-1986 samples from Chernobyl were significantly larger than variances for feather samples from the control region, and than variances for historical samples from both regions. These findings suggest that stable-isotope measurements can provide information about population processes following environmental perturbations.  相似文献   

12.
Stable isotope ratios (typically of carbon and nitrogen) provide one representation of an organism's trophic niche and are widely used to examine aspects of food web structure. Yet stable isotopes have not been applied to quantitatively characterize community-wide aspects of trophic structure (i.e., at the level of an entire food web). We propose quantitative metrics that can be used to this end, drawing on similar approaches from ecomorphology research. For example, the convex hull area occupied by species in delta13C-delta15N niche space is a representation of the total extent of trophic diversity within a food web, whereas mean nearest neighbor distance among all species pairs is a measure of species packing within trophic niche space. To facilitate discussion of opportunities and limitations of the metrics, we provide empirical and conceptual examples drawn from Bahamian tidal creek food webs. These examples illustrate how this methodology can be used to quantify trophic diversity and trophic redundancy in food webs, as well as to link individual species to characteristics of the food web in which they are embedded. Building from extensive applications of stable isotope ratios by ecologists, the community-wide metrics may provide a new perspective on food web structure, function, and dynamics.  相似文献   

13.
We investigated migration and wintering of adult northern gannets (Morus bassanus) breeding in east Scotland, North Sea, by deploying geolocation loggers over three winters. The wintering ranges of these birds varied from the North Sea to the Atlantic off West Africa. Flight time was taken as a proxy for migration and foraging effort. Gannets wintering off Africa had higher total flight times during migration than birds wintering further north. Total flight times in different wintering regions were generally low. Birds off West Africa consistently spent < 25 % of daylight hours in flight, but birds further north showed more variable values that may reflect more variable weather or food availability. Winter sea surface temperatures ranged from 9 °C (North Sea) to 16 °C (West Africa). Thermostatic costs in winter as estimated by measuring thermal conductance in carcasses in still air and water were 28 % higher in North Sea than off West Africa. This effect is aggravated by higher thermostatic costs caused by stronger wind chills in the North Sea compared to the conditions off West Africa. Birds wintering close to the UK arrived at the colony on average 12 days earlier than birds wintering off West Africa. We conclude that the net cost-benefit analysis may be similar for all wintering areas investigated.  相似文献   

14.
Stable isotope analysis of carbon and nitrogen is frequently used to study the diets and foraging ecology of marine predators. However, isotopic values may also be affected by an individual’s nutritional status and associated physiological processes. Here, we use C and N stable isotopes in blood and feathers of blue-footed booby chicks at the Galápagos Islands to examine how isotopic values are related to body condition and growth rate, and to assess the consistency in the isotope ratios of individuals during growth. Size dimorphism in blue-footed boobies provided an additional opportunity to examine how isotope ratios differ between sexes in relation to body size and growth rate. There was no significant difference between sexes but both C and N stable isotopes were significantly negatively related to the body condition of chicks. These data were consistent with individual variation in physiological processes affecting fractionation, although we cannot rule out the possibility that they were also influenced to some extent by population-level variation in the stable isotope ratios of prey fed to chicks, for instance related to prey size, depth or lipid content. Our results highlight the need for methods that take proper account of confounding physiological factors in isotopic studies of foraging ecology and diet.  相似文献   

15.
Stable isotopes of growing feathers and blood both represent assimilated diet, and both tissues are used to study the diet and foraging distribution of marine and terrestrial birds. Although most studies have assumed that both tissues represent a difference of one trophic level to diet, the enrichment factors of blood and feathers may differ, especially where endogenous reserves are used as precursors during feather synthesis. In this study, we compare carbon and nitrogen stable isotopes of blood and simultaneously growing feathers of five species of Procellariiformes, representing five genera, different geographical regions and different life stages (chicks and adults). In all species, feathers were enriched in 15N and 13C compared with blood. Isotopic values of carbon and nitrogen were correlated in different tissues growing simultaneously for most species analyzed, suggesting that mathematical corrections could be used to compare different tissues. Our results imply that more care needs to be taken when comparing stable isotope signatures across studies assuming different tissues are equivalent indicators of trophic ecology.  相似文献   

16.
Summary Explanations of delayed plumage maturation (DPM) in passerines have focused on potential breeding season advantages for non-definitive (subadult) plumage. In contrast, the molt constraint hypothesis (Rohwer and Butcher 1988) proposes that subadult plumage is a winter adaptation, increasing winter survivorship by decreasing intraspecific aggression (the winter status signaling hypothesis) or predation (the winter crypsis hypothesis). Under the molt constraint hypothesis, nondefinitive breeding plumage is non-adaptive, resulting from species-specific constraints on the number of feathers replaced during the pre-alternate (spring) molt. In studies conducted on an overwintering population of orchard orioles in Panama, I tested predictions of both the winter status signaling and molt constraint hypotheses. Contrary to predictions of winter status signaling, I found no evidence that subadult plumage reduces adult male aggression toward subadults. Agonistic encounters occurred at random with respect to plumage and the intensity of adult male/subadult male encounters was not lower than the intensity of encounters occurring within age classes. Contrary to the molt constraint hypothesis, I found no evidence that the number of feathers exchanged by subadults in their pre-alternate molt is the sole constraint on the development of subadult breeding plumage. The majority of feathers grown by molting subadult orioles during January and February were of non-definitive coloration. These results, together with results of earlier breeding season experiments which tested summer communication hypotheses for DPM in this species, suggest that subadult plumage in the orchard oriole may be non-adaptive and the result of constraints on plumage development. They also indicate, however, that the extent of the pre-alternate molt is not the sole source of that constraint.  相似文献   

17.
Effects of deployment of miniaturised transmitters and loggers have been studied mainly in diving seabirds such as penguins, and less so in flying seabirds. However, some studies of albatrosses and petrels recorded extended trip durations and elevated rates of nest desertion following device attachment, especially if transmitter loads exceeded 3 % of adult mass. Studies have usually compared performance parameters such as trip duration, meal mass, breeding success or rate of return in the next season between birds with devices and controls. We here examined the effects of geolocator loggers (Global Location Sensing, (GLS)) on thin-billed prions Pachyptila belcheri (130 g), by comparing performance parameters and additionally eco-physiological parameters. GLS weighed ca. 1 % of the body mass, and were fixed on leg rings, which may influence the flight efficiency by creating an asymmetric load. We found no differences in the performance parameters, either in the season of attachment or the season following recovery. Similar stable isotope ratios in adult blood and feather samples further indicated that the foraging ecology was not influenced. However, after 1 year of logger deployment, adults differed in their hormonal response to stress: while baseline corticosterone levels were not influenced, corticosterone levels in response to handling were elevated. Moreover, increased heterophil/lymphocyte ratios and a decreased tail growth in winter suggest that carrying the GLS was energetically costly, and adults adapted physiologically to the higher work load, while keeping up a normal breeding performance.  相似文献   

18.
Many species of marine organisms go through ontogenetic shifts that occur in unknown or inaccessible locations. Finding these areas is crucial to understand connectivity and resilience of populations, both of which have conservation implications. When extrinsic markers are not suitable to track organisms, intrinsic markers can be useful to infer the location of inaccessible areas where these cryptic stages occur. Our study focuses on the location of oceanic foraging areas of the cryptic early juvenile stage of green turtles, Chelonia mydas, in the Atlantic. Due to the small size of hatchlings, the use of telemetry is limited to short periods of time and small spatial ranges, which do not allow determining the location of oceanic foraging areas. We used lead (Pb) stable isotopes to determine the possible location of oceanic foraging areas of small green turtles in the Atlantic Ocean. Pb stable isotope ratios in the scute tissue deposited when turtles were in the oceanic habitat were compared to ratios of major sources of lead in the Atlantic and oceanic areas in the Atlantic to determine the location of oceanic foraging grounds. The Pb isotope ratios in the scute of oceanic-stage green turtles indicated that turtles use different regions in the Atlantic and that they are capable of transatlantic migrations. We compare the oceanic locations identified by this study with those suggested by two previous studies.  相似文献   

19.
Doucett RR  Marks JC  Blinn DW  Caron M  Hungate BA 《Ecology》2007,88(6):1587-1592
Understanding river food webs requires distinguishing energy derived from primary production in the river itself (autochthonous) from that produced externally (allochthonous), yet there are no universally applicable and reliable techniques for doing so. We compared the natural abundance stable isotope ratios of hydrogen (deltaD) of allochthonous and autochthonous energy sources in four different aquatic ecosystems. We found that autochthonous organic matter is uniformly far more depleted in deuterium (lower deltaD values) than allochthonous: an average difference of approximately 100% per hundred. We also found that organisms at higher trophic levels, including both aquatic invertebrates and fish, have deltaD values intermediate between aquatic algae and terrestrial plants. The consistent differences between leaves and algae in deltaD among these four watersheds, along with the intermediate values in higher trophic levels, indicate that natural abundance hydrogen isotope signatures are a powerful tool for partitioning energy flow in aquatic ecosystems.  相似文献   

20.
We utilised stable isotope ratios to assess differences in diet among three indigenous and syntopic rocky shore suspension-feeders (mussels Perna perna, barnacles Tetraclita serrata, and polychaetes Gunnarea gaimardi). We also determined the spatial and temporal variability in the suspension-feeder diets by collecting specimens on two occasions from two regions adjacent to hydrologically distinct river mouths (i.e. one with larger annual freshwater throughput than the other). The results showed that the isotopic niches (used as proxies for trophic niches) of the three species did not overlap and that the barnacles occupied a trophic position (3.4) well above those of the mussels (2) and polychaetes (2.6). We ascribed the interspecific differences primarily to the disparate feeding mechanisms used by the species. Large-scale regional (~50 km) differences in suspension-feeder diets were apparent, but not small-scale (up to a few km north and south of each estuary mouth). The regional differences in diet resulted from the increased availability of estuarine-origin suspended particulate matter (SPM) and zooplankton in the region adjacent to the river with relatively larger freshwater output, although overall incorporation of zooplankton versus mixed SPM into consumer diets was relatively consistent between regions and through time. Temporal shifts in suspension-feeder diets were apparent from stable carbon isotope ratios in the consumers. Our results showcase the measurable effects of regional-scale processes that can alter the food sources for dominant primary consumers in the rocky intertidal, thus potentially affecting entire food webs through bottom-up processes. The clear evidence for trophic niche partitioning offers valuable insights into how potentially strong competitors can coexist.  相似文献   

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