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1.
Inter-individual diet variation within populations is likely to have important ecological and evolutionary implications. The diet-fitness relationships at the individual level and the emerging population processes are, however, poorly understood for most avian predators inhabiting complex terrestrial ecosystems. In this study, we use an isotopic approach to assess the trophic ecology of nestlings in a long-lived raptor, the Bonelli’s eagle Aquila fasciata, and investigate whether nestling dietary breath and main prey consumption can affect the species’ reproductive performance at two spatial scales: territories within populations and populations over a large geographic area. At the territory level, those breeding pairs whose nestlings consumed similar diets to the overall population (i.e. moderate consumption of preferred prey, but complemented by alternative prey categories) or those disproportionally consuming preferred prey were more likely to fledge two chicks. An increase in the diet diversity, however, related negatively with productivity. The age and replacements of breeding pair members had also an influence on productivity, with more fledglings associated to adult pairs with few replacements, as expected in long-lived species. At the population level, mean productivity was higher in those population-years with lower dietary breadth and higher diet similarity among territories, which was related to an overall higher consumption of preferred prey. Thus, we revealed a correspondence in diet-fitness relationships at two spatial scales: territories and populations. We suggest that stable isotope analyses may be a powerful tool to monitor the diet of terrestrial avian predators on large spatio-temporal scales, which could serve to detect potential changes in the availability of those prey on which predators depend for breeding. We encourage ecologists and evolutionary and conservation biologists concerned with the multi-scale fitness consequences of inter-individual variation in resource use to employ similar stable isotope-based approaches, which can be successfully applied to complex ecosystems such as the Mediterranean.  相似文献   

2.
Understanding the mechanisms that shape animal population dynamics is of fundamental interest in ecology, evolution and conservation biology. Food supply is an important limiting factor in most animal populations and may have demographic consequences. Optimal foraging theory predicts greater consumption of preferred prey and less diet diversity when food is abundant, which may benefit key fitness parameters such as productivity and survival. Nevertheless, the correspondence between individual resource use and demographic processes in populations of avian predators inhabiting large geographic areas remains largely unexplored, particularly in complex ecosystems such as those of the Mediterranean basin. Based on a long‐term monitoring program of the diet and demography of Bonelli's eagle Aquila fasciata in western Europe, here we test the hypothesis that a predator's diet is correlated to its breeding productivity and survival at both the territorial and population levels, and ultimately to its population growth rate. At the territorial level, we found that productivity increased with greater consumption of European rabbits Oryctolagus cuniculus, the Bonelli's eagle's preferred prey, and pigeons, an important alternative prey for this predator. The survival of territorial pairs was negatively affected by higher diet diversity, which probably reflected the inability to find sufficient high quality prey. Diet effects at the population level were similar but more noticeable than at the territorial level, i.e. a greater consumption of rabbits, together with lesser consumption of small‐to‐medium avian species (‘other birds’; non‐preferred prey), increased productivity, while greater diet diversity and lower consumption of rabbits was associated with reduced survival and population growth rate. Overall, our study illustrates how the diet of a predator species can be closely related to key individual vital rates, which, in turn, leave a measurable fingerprint on population dynamics within and among populations across large spatial scales.  相似文献   

3.
Age-dependent breeding performance is arguably one of the best-documented phenomena in ornithology. The existence of age-related trends has major implications for life-history theory, but the proximate reasons for these patterns remain poorly understood. It has been proposed that poor breeding performance of young individuals might reflect lack of foraging skills. We investigated this possibility in a medium-sized, powerful raptor-the northern goshawk Accipiter gentilis. Male goshawks are responsible for providing their females and their offspring with food. We hypothesized that young males may generally show poor breeding performance or even delay breeding, because they lack the experience to hunt efficiently-especially, their principal avian prey, the feral pigeon Columba livia. Our study exploited a rare 'natural experiment', the expansion phase of an urban population, where intraspecific interference was negligible and many young males bred successfully. This enabled us to examine the improvement of foraging skills in a larger sample of young individuals, and in more controlled conditions than usually possible. Using data from individually identified male breeders, we show that, consistent with our hypothesis, the proportion of pigeons in the diet increased significantly with male age, for at least the first three years of life. Other studies have shown a parallel increase in productivity, and a positive effect of a pigeon-rich diet on brood size and nestling condition, stressing the potential fitness relevance of this prey species for goshawks. Our results suggest a causal link between patterns of age-dependence in foraging ecology and reproductive performance. Furthermore, our study is, to our knowledge, the first demonstration that prey choice of breeders, which might reflect individual hunting skills, is age-dependent in a raptor.  相似文献   

4.
We studied a population of 23–25 Eagle Owl Bubo bubo pairs between 1994 and 2000 in a 1330-km2 study plot in the central-eastern Italian Alps. Compared to random sites, territories were located at lower elevation and closer to intensively cultivated-urbanized valley floors. Early laying was associated with low elevation and negatively affected productivity. Diet was dominated by rats, hedgehogs and dormice ( n  = 978 prey items), all of them typical of low-elevation habitats. Higher productivity was associated with a higher proportion of rats in the diet of individual pairs. Low availability of rats resulted in a more diverse diet, in turn associated with low productivity. Territories were occupied every year in a non-random fashion, and those most occupied were characterized by higher productivity and higher occurrence of the favoured prey types in the diet, suggesting they were of superior quality. Eagle Owls also paid a cost associated with nesting near human-altered habitats: the main cause of mortality reported to local authorities was electrocution. This is an increasing cause of death for many European populations and may be a cause for conservation concern. Human persecution is also an important cause of mortality in some parts of the European range. Apart from such costs, the study population appeared to have adapted well to the proximity of humans: estimates of density and productivity were comparable to those recorded elsewhere in Europe. The pattern found in our population also held at higher spatial scales: data from 17 European populations showed density to be highest in low-elevation, human-altered landscapes.  相似文献   

5.
Between 2000 and 2009 we studied the diet and breeding success of Egyptian Vultures Neophron percnopterus in southern Spain. Wild species accounted for 74.9% of prey items (n = 1071) with a predominance of mammals (62.3%), followed by birds (20.8%) and reptiles (13.1%). Spatially, the diet was highly varied and not restricted to carcasses of livestock; wild Rabbits Oryctolagus cuniculus accounted for 54% of the overall remains. The spatial variability may reflect regional and local disparity in the availability of main prey. The temporal relationship between variation in trophic diversity and Vulture nesting productivity (both values showing a long‐term decrease) might suggest a causal link between variation in diet and reproductive output. We hypothesize that high turnover rates could explain productivity variation as a consequence of the recruitment of less experienced individuals to the breeding population. This could in turn generate covariation between diet and reproductive output.  相似文献   

6.
While intra‐population variability in resource use is ubiquitous, little is known of how this measure of niche diversity varies in space and its role in population dynamics. Here we examined how heterogeneous breeding environments can structure intra‐population niche variation in both resource use and reproductive output. We investigated intra‐population niche variation in the Arctic tundra ecosystem, studying peregrine falcon (Falco peregrinus tundrius, White) breeding within a terrestrial‐marine gradient near Rankin Inlet, Nunavut, Canada. Using stable isotope analysis, we found that intra‐population niches varied at the individual level; we examined within‐nest and among‐nest variation, though only the latter varied along the terrestrial‐marine gradient (i.e., increased among‐nest variability among birds nesting within the marine environment, indicating higher degree of specialization). Terrestrial prey species (small herbivores and insectivores) were consumed by virtually all falcons. Falcons nesting within the marine environment made use of marine prey (sea birds), but depended heavily on terrestrial prey (up to 90% of the diet). Using 28‐years of peregrine falcon nesting data, we found a positive relationship between the proportion of terrestrial habitat surrounding nest sites and annual nestling production, but no relationship with the likelihood of successfully rearing at least one nestling reaching 25 days old. Annually, successful inland breeders raised 0.47 more young on average compared to offshore breeders, which yields potential fitness consequences for this long‐living species. The analyses of niche and reproductive success suggest a potential breeding cost for accessing distant terrestrial prey, perhaps due to additional traveling costs, for those individuals with marine nest site locations. Our study indicates how landscape heterogeneity can generate proximate (niche variation) and ultimate (reproduction) consequences on a population of generalist predator. We also show that within‐individual and among‐individual variation are not mutually exclusive, but can simultaneously arise and structure intra‐population niche variation.  相似文献   

7.
The breeding population of the black-browed albatross Thalassarche melanophrys has increased at Heard Island since the first census data were obtained in 1947/1948. Four breeding localities are known, and all populations have increased in the period 1947/1948-2000/2001. The breeding population is estimated to have been approximately 200 pairs in 1947/1948. Based on 2000/2001 census data, the population has increased to a minimum of approximately 600 pairs over the 53 years. Two mechanisms, that of increased prey availability through scavenging discards from trawlers operating within their foraging range, and climatic amelioration, are proposed as hypotheses for this increase.  相似文献   

8.
1. Understanding how density-dependent and independent processes influence demographic parameters, and hence regulate population size, is fundamental within population ecology. We investigated density dependence in growth rate and fecundity in a recovering population of a semicolonial raptor, the osprey Pandion haliaetus [Linnaeus, 1758], using 31 years of count and demographic data in Corsica. 2. The study population increased from three pairs in 1974 to an average of 22 pairs in the late 1990s, with two distinct phases during the recovery (increase followed by stability) and contrasted trends in breeding parameters in each phase. 3. We show density dependence in population growth rate in the second phase, indicating that the stabilized population was regulated. We also show density dependence in productivity (fledging success between years and hatching success within years). 4. Using long-term data on behavioural interactions at nest sites, and on diet and fish provisioning rate, we evaluated two possible mechanisms of density dependence in productivity, food depletion and behavioural interference. 5. As density increased, both provisioning rate and the size of prey increased, contrary to predictions of a food-depletion mechanism. In the time series, a reduction in fledging success coincided with an increase in the number of non-breeders. Hatching success decreased with increasing local density and frequency of interactions with conspecifics, suggesting that behavioural interference was influencing hatching success. 6. Our study shows that, taking into account the role of non-breeders, in particular in species or populations where there are many floaters and where competition for nest sites is intense, can improve our understanding of density-dependent processes and help conservation actions.  相似文献   

9.
Breeding ecology of brown skuas (Catharacta antarctica lonnbergi) was studied at Bird Island, South Georgia in the austral summers of 2000/2001–2003/2004. A complete census recorded 467 breeding pairs in 3.55 km2 of suitable habitat (132 pairs per km2), and an additional 312 nonbreeders at club-sites. Comparison with previous counts indicates two phases of population change: an initial rapid increase (3.6% per annum) from the late 1950s to early 1980s, probably attributable to increased carrion availability from the expanding Antarctic fur seal (Arctocephalus gazella) population, followed by slower growth (0.9% p.a.). Currently, seal carrion dominates the diet of skuas during incubation, with a switch to seabird prey during chick-rearing. Breeding is now later, chick growth poorer, and productivity significantly lower than in the early 1980s. There is also a strong seasonal decline in adult attendance, and chicks that hatch later and are in poorer condition are less likely to fledge. These results suggest a long-term increase in competition for carrion that is particularly apparent once fur seal pupping has ceased.  相似文献   

10.
Amongst raptor species, individuals with specialized diets are commonly observed to have higher reproductive output than those with general diets. A suggested cause is that foraging efficiency benefits accrue to diet specialists. This diet specificity hypothesis thus predicts that diet breadth and reproductive success should be inversely related within species. We highlight, however, that a prey availability hypothesis also makes the same prediction in some circumstances. Hence, when high diet specificity results from high encounter rates with an abundant, preferred prey, then prey availability may affect reproductive success, with diet specialization as an incidental correlate. Using three insular study areas in western Scotland, we examine diet specificity and reproductive success in Golden Eagles Aquila chrysaetos. Diet breadth and breeding productivity were not negatively related in any of our study areas, even though birds with specific diets did tend to have a higher incidence of preferred prey (grouse and lagomorphs) in the diet. Indeed, in two study areas there was evidence that diet generalists had higher breeding productivity. Our results therefore failed to support the diet specificity hypothesis but were consistent with the prey availability hypothesis. We highlight that although many other studies are superficially consistent with the diet specificity hypothesis, our study is not alone in failing to provide support and that the hypothesis does not provide a generic explanation for all relevant results. Diet specificity in predators can be at least partially a response to prey diversity, availability and distribution, and benefits associated with different prey types, so that being a generalist is not necessarily intrinsically disadvantageous. We suggest that the available evidence is more consistent with variation in prey abundance and availability as a more influential factor explaining spatial and temporal variation in breeding productivity of ‘generalist’ species such as the Golden Eagle. Under this argument, prey abundance and availability are the main drivers of variation in reproductive output. Diet specificity is a consequence of variation in prey availability, rather than a substantial cause of variation in reproductive success.  相似文献   

11.
Understanding variation in food requirements of wild animals is of central importance in population ecology and conservation, as it helps to identify where and when food may be limiting. Studies on diet variation or prey provisioning rates may give useful insights when direct information on prey availability is lacking. We assess spatial and temporal variation in the diet of an endangered predator, the Black Harrier Circus maurus. This raptor is endemic to southern Africa and specializes on small mammals but also feeds on birds and reptiles as alternative prey. Using data on 1679 prey identified in 953 pellets collected in inland and coastal regions from 2006 to 2015, we show that diet composition changed little throughout the breeding season in the coastal region, whereas there was a marked seasonal decline in the occurrence of small mammal prey in the inland region, with a concomitant increase in alternative prey. The proportion of small mammals in the diet declined with increasing maximum temperature, the latter being highest at the inland region late in the breeding season. Using camera recordings at nests in 2014, we further analysed daily patterns of prey provisioning to nestlings. A marked reduction in small mammal provisioning rates occurred during the middle of the day in the hotter inland region but not in the cooler coastal region. Reduced availability of the primary prey, small mammals, in hotter conditions, through a reduction in activity or overall abundance, could explain these patterns. Finally, we show a positive relationship between winter rainfall and interannual differences in the proportion of small mammals in the diet of Black Harriers breeding in the coastal region, suggesting relationships between diet and prey abundance that are mediated through rainfall. We discuss the need to consider spatial variation in food availability in conservation strategies.  相似文献   

12.
A better understanding of the mechanisms driving superpredation, the killing of smaller mesopredators by larger apex predators, is important because of the crucial role superpredation can play in structuring communities and because it often involves species of conservation concern. Here we document how the extent of superpredation has changed over time, and assessed the impact of such temporal variation on local mesopredator populations using 40 yr of dietary data collected from a recovering population of northern goshawks Accipiter gentilis, an archetypical avian superpredator. We then assessed which mechanisms were driving variation in superpredation, e.g. was it opportunistic, a response to food becoming limited (due to declines in preferred prey) or to reduce competition. Raptors comprised 8% of goshawk diet on average in years when goshawk abundance was high, which is higher than reported elsewhere. Additionally, there was a per capita increase in superpredation as goshawks recovered, with the proportion of goshawk diet comprising raptors increasing from 2 to 8% as the number of goshawk home‐ranges increased from ≤ 14 to ≥ 25. This increase in superpredation coincided with a population decline in the most commonly killed mesopredator, the Eurasian kestrel Falco tinnunculus, which may represent the reversal of the ‘mesopredator release’ process (i.e. mesopredator suppression) which occurred after goshawks and other large raptors declined or were extirpated. Food limitation was the most likely driver of superpredation in this system given: 1) the substantial decline of two main prey groups in goshawk diet, the increase in diet diversity and decrease in goshawk reproductive success are all consistent with the goshawk population becoming food‐limited; 2) it's unlikely to be purely opportunistic as the increase in superpredation did not reflect changes in the availability of mesopredator species; and 3) the majority of mesopredators killed by goshawks do not compete with goshawks for food or nest sites.  相似文献   

13.
The conservation status of the hen harrier Circus cyaneus in Britain, and its pivotal role in the raptor–grouse moor conflict, has rendered research on the factors influencing its population dynamics of crucial importance. Using over 20 years of data, we examine influences on the reproductive output of the small, but recently expanding, population of Welsh hen harriers. Productivity (fledglings per breeding attempt) has steadily increased with time, being the highest in recent years. Available evidence suggested that food availability could not account for these recent increases. Human interference (apparently, largely or entirely in the form of nest destruction by grouse moor gamekeepers) had a large influence on productivity, despite being restricted to a minority of the breeding population. This negative effect, and a positive effect of May temperature, was most influential on variation in breeding productivity. This study quantifies, for the first time, that cessation of persecution can result in a marked improvement in hen harrier reproductive output. Nest predation by red foxes Vulpes vulpes had no effect on variation in breeding productivity and there was no evidence that control of fox numbers by gamekeepers compensated for their depression of productivity through destroying harrier nests. Our analyses also indicated that gamekeepers probably destroyed an unknown number of nests before they were discovered. We argue that a low proportion of territorial females that apparently breed, in cases when clutch size and fledged brood size are high, may be an indication of persecution. The recent increase in the breeding productivity of Welsh harriers has probably been influential in the recent recovery of the Welsh harrier population and has apparently been due to a combination of cessation of human interference and warmer temperatures.  相似文献   

14.
The recovery of the peregrine falcon (Falco peregrinus anatum) in California has taken place amid strong geographical differences in habitat quality, potentially creating a sink population in the southern coastal habitat and source populations in the northern interior and urban habitats. We analyzed long-term monitoring data to investigate the mechanisms and consequences of spatial structuring for the recovery of this set of nonstable subpopulations. Dispersal rates between habitats were asymmetric, with extremely limited dispersal out of the interior habitat and a strong tendency for birds in the southern coast to disperse to the urban habitats. We used these dispersal estimates and habitat-specific productivity rates to build a set of regional population models that describe population growth within and dispersal between each subpopulation. We tested for the existence of habitat-specific survival and territory acquisition rates by comparing model projections with the number of breeding pairs censused annually in each subpopulation. Our analyses indicate a high rate of survival for interior birds and suggest that both the interior and urban subpopulations were regulated by territory availability over the study period. The inherent spatial structure of this regional peregrine falcon population has had a considerable influence on its recovery and management.  相似文献   

15.
Using records from archives detailing bounties for wolves killed in northern Spain during the eighteenth and nineteenth centuries, we investigated demographic and spatial distribution parameters of the population to determine whether direct persecution or prey availability was responsible for the observed population decline. Captures of adult, subadult, and young individuals, including those of litters, showed a downward trend. Progressive decreases in age ratio and litter size, and the increase in the proportion of males, were compatible with a population under food stress, driven by the extinction of wild ungulates, the sharp reduction in livestock numbers, and the lack of alternative prey. The immigration and dispersal process does not seem to have functioned under such conditions. In the study area, where strychnine was not used until the end of the nineteenth century, the broadly accepted idea of human persecution having an exclusive or primary role in wolf decline does not necessarily apply.  相似文献   

16.
17.
Specialist predators are generally negatively impacted by habitat change. Predators that inhabit transformed areas are usually forced to diversify their diet and this departure away from traditional resources can have negative consequences for fitness and demographic parameters. We consider this relationship as it applies to Verreaux's eagles Aquila verreauxii, which is typically considered to be a highly specialised predator of hyraxes (Procavia and Heterohyrax spp.). We investigate diet in relation to land cover in two adjacent areas of South Africa and explore the links between diet diversity, the percentage of hyrax consumed, and the breeding performance of eagles. We also examine these same patterns using data from other studies. We found that diet diversity was greater in the agriculturally developed Sandveld region compared to the natural Cederberg region. Proportions of the three main prey types were correlated with the proportion of agriculturally developed land around the nest site. Breeding performance was correlated with the diet, but not in the manner expected, with breeding productivity being greater in regions with large diet diversity and a small proportion of hyrax in the diet. We found similar patterns when placing our results into a broader geographical context using other dietary studies of Verreaux's eagles, suggesting our results were not unique to our study system. Thus, our results suggest that diet diversification does not necessarily impinge on breeding performance in the presence of adequate alternative prey resources. This research adds to the growing number of studies suggesting that some predators may be adaptable up to a threshold level of habitat transformation. These results have implications for predicting changes on such species by anthropogenic habitat transformation and highlight the potential for agriculturally developed areas to maintain a conservation value when habitat heterogeneity is maintained.  相似文献   

18.
The demographic consequences of within-population variability in predator foraging are not well understood. We assessed the relationship between the degree of diet specialization and two demographic parameters, population density and reproductive output, within a single population of Imperial Eagles Aquila heliaca at a nature reserve in north-central Kazakhstan. Nearest-neighbour distances between eagle nests throughout the reserve, and thus population density, were correlated with the degree to which diets were specialized. Diet diversity showed an extensive regional variability that was linked to prey distributions, but within-year analyses of reproductive output did not show similar linkages. However, multi-year analyses of breeding performance showed inter-regional differences in nesting success that were paralleled, and probably driven by, similar trends in diet diversity. In contrast, brood size at fledging was not linked to diet diversity and was more probably driven by reserve-wide influences such as weather. Finally, the decision to initiate breeding was associated neither with diet diversity nor with environmental variability. Our results indicate that the degree of dietary specialization is linked to the demographics of Imperial Eagle populations. For these and other raptor populations, it is possible that management could be used separately to increase or decrease nesting success, brood size at fledging, and the likelihood that a pair will initiate breeding.  相似文献   

19.
The Mauritius Kestrel Falco punctatus, once the rarest kestrel worldwide, became an icon of bird conservation after it recovered from four to six individuals in 1974 to some 800 by 2005 following intense conservation management. Its population however then halved within about a decade prompting a re-evaluation of the IUCN status and up listing of the species in 2014 and an increased conservation attention. Drivers of this new decline are unclear and the influence of habitat structure and diet on breeding success may be important contributors but have received relatively little attention, particularly in the way they may interact to influence production of new fledglings. We address this knowledge gap by studying whether breeding success is influenced by habitat structure (in terms of cover of the invasive Ravenala in native habitats, an alien plant causing strong structural shift in the forests that it invades, and extent of cleared area), diet composition and food pass frequency (as a proxy for food intake) and food quality at 28 nests of a re-introduced kestrel population in south east Mauritius during the 2015–2016 breeding season. The kestrel’s diet comprised native and alien birds, reptiles, insects, and small alien mammals, with a disproportionately high proportion of Phelsuma gecko. A higher frequency of food provisioning and percentage cover of Ravenala both contributed to higher breeding success. Ravenala may increase gecko density or increase gecko detectability and predation by the kestrel, or both, while changed land use (pasture and sugar cane fields) may increase prey diversity in the form of non-forest prey known to be eaten by Kestrels (e.g. alien agamids, small mammals and birds). These prey related influences on breeding suggest that the Bambou mountain range provides a human-generated novel ecosystem altering food availability and increasing the kestrel’s breeding success. However, Ravenala is an invasive alien species harmful to the wider forest biodiversity. Progressive weeding of Ravenala and concurrent re-introduction and augmentation of native palms and Pandanus species which geckos can use at comparable densities to Ravenala, is recommended. This would likely improve the kestrel’s hunting habitat quality and maintain high gecko density or detectability and the vegetation structure required for hunting manoeuvrability and prey availability without the negative consequences of invasive Ravenala.  相似文献   

20.
Shifting prey availability can lead to altered species interactions, indicated by variation in the dietary niche breadth and position of species within an assemblage. On the Newfoundland coast, annual inshore spawning migration of the dominant forage fish, Capelin Mallotus villosus, provides an excellent opportunity to investigate the influence of varying prey availability on dietary niche breadth and position among species. During June–August 2017, we investigated species‐ and assemblage‐level dietary responses to shifting Capelin availability of three Capelin‐eating, sympatrically breeding auk species, the Atlantic Puffin Fratercula arctica, Razorbill Alca torda and Common Murre Uria aalge. The diet of Leach's Storm Petrels Oceanodroma leucorhoa, which breed alongside the three auk species but are not known to rely on Capelin, was also examined to determine dietary shifts throughout breeding that were unrelated to Capelin availability. We quantified stable isotope ratios (δ15N, δ13C) in seabird blood components (plasma, cellular component) collected both before and after spawning Capelin arrived in the study area and compared isotopic niche breadth within a Bayesian framework. At the species level, auk trophic position increased and isotopic niche breadth narrowed after Capelin arrived, suggesting a more Capelin‐based diet. Simultaneously, trophic diversity of the auk assemblage, reflecting the extent of spacing among niches of species, decreased after spawning Capelin arrived inshore. Contrastingly, increased trophic position but broader isotopic niche breadth during higher relative to lower Capelin availability for Leach's Storm Petrel confirm that this species is probably not affected by the inshore arrival of Capelin, but instead that isotopic changes may be more related to a shift in breeding stage to chick‐rearing. Overall, our findings reiterate the importance of Capelin as a prey resource for breeding auks in coastal Newfoundland, but that the degree of reliance on Capelin varies among species, possibly allowing coexistence of these ecologically similar species. The findings highlight potential changing species interactions, such as increased competition, under declines in Capelin biomass.  相似文献   

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