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1.
The presence of and mechanisms behind density-dependent growth and resource limitation in larval and juvenile stages of organisms with high mortality such as fish are much debated. We compare observed consumption and growth rates with maximum consumption and growth rates to study the extent of resource limitation in young-of-the-year (YOY) roach (Rutilus rutilus) and perch (Perca fluviatilis). Diet, habitat use, consumption rate and growth rate were measured under varying YOY fish densities over 2 years in four lakes. In the first year, YOY roach and perch were studied under allopatric conditions. Experimental addition of perch roe in the second year also allowed study of YOY of the two species under sympatric conditions in two of the lakes. The diet of YOY roach was dominated by cladoceran zooplankton and YOY roach habitat use was restricted to the shore region in both years. This restricted habitat use did not involve any cost in foraging gain in the first year as consumption and growth rates were very close to maximum rates. During the second year, when the two species coexisted, resources were limited in late season, more so in the littoral than in the pelagic habitat in one lake while the reverse was the case in the other lake. The diet of YOY perch was also dominated by zooplankton, and with increasing perch size the proportion of macroinvertebrate prey in the diet increased. After hatching, YOY perch first utilized the pelagic habitat restricting their habitat use to the shore after 1 to several weeks in the pelagic zone. During the larval period, perch were not resource limited whereas juvenile perch were resource limited in both years. The fact that YOY perch were more resource limited than YOY roach was related to the higher handling capacity and lower attack rate of perch relative to roach, rendering perch more prone to resource limitation. Estimates of resource limitation based on consumption rates and growth rates yielded similar results. This supports the adequacy of our approach to measure resource limitation and suggests that this method is useful for studying resource limitation in organisms with indeterminate growth. Our results support the view that density-dependent growth is rare in larval stages. We suggest that density-dependent growth was absent because larval perch and roach were feeding at maximum levels over a wide range of larvae densities. Received: 14 June 1999 / Accepted: 29 October 1999  相似文献   

2.
Horizontal and vertical heterogeneity as a result of size‐structured processes are important factors influencing indirect effects in food webs. In a whole‐lake experiment covering 5 years, we added the intermediate consumer roach (Rutilus rutilus) to two out of four lakes previously inhabited by the omnivorous top predator perch (Perca fluviatilis). We focused our study on the direct consumption effect of roach presence on zooplankton (and indirectly phytoplankton) versus the indirect effect of roach on zooplankton (and phytoplankton) mediated via effects on perch reproductive performance. The patterns in zooplankton and phytoplankton abundances were examined in relation to population density of roach and perch including young‐of‐the‐year (YOY) perch in the light of non‐equilibrium dynamics. The presence of roach resulted in changed seasonal dynamics of zooplankton with generally lower biomasses in May–June and higher biomasses in July–August in roach lakes compared to control lakes. Roach presence affected perch recruitment negatively and densities of YOY perch were on average higher in control lakes than in treatment lakes. In years when perch recruitment did not differ between lakes as a result of experimental addition of perch eggs, total zooplankton biomass was lower in treatment lakes than in control lakes. Phytoplankton biomass showed a tendency to increase in roach lakes compared to control lakes. Within treatment variation in response variables was related to differences in lake morphometry in treatment lakes. Analyses of the trophic dynamics of each lake separately showed strong cascading effects of both roach and YOY perch abundance on zooplankton and phytoplankton dynamics. Consideration of the long transients in the dynamics of top predators (fish) in aquatic systems that are related to their long life span involving ontogenetic niche shifts is essential for making relevant interpretations of experimental perturbations. This conclusion is further reinforced by the circumstance that the intrinsic dynamics of fish populations may in many cases involve high amplitude dynamics with long time lags.  相似文献   

3.
Studies on resource polymorphism have mainly been considered at the end stage of ontogeny, whereas many species undergo diet changes as they grow. We conducted a field survey to analyze the role of adaptive variation during ontogeny in Eurasian perch (Perca fluviatilis). We caught perch from the littoral and pelagic zones of a lake to investigate whether perch differ in morphology and diet between these habitats. We also investigated whether there were any differences in morphological trajectories during the ontogeny of perch collected from the two habitats. We found that perch caught in the littoral habitat, independently of size, had a deeper body, larger head and mouth and longer fins than perch caught in the pelagic zone. Macroinvertebrates and fish dominated the diet of littoral perch, whereas the diet of the pelagic perch consisted mainly of zooplankton and to some extent fish. Independently of size, the more streamlined individuals had a larger proportion of zooplankton and a smaller proportion of macroinvertebrates in their diet than the deeper-bodied individuals, indicating a relation between diet and morphology. Some morphological characters followed different ontogenetic trajectories in the two habitats; e.g. the changes to a deeper body and a larger head were faster in the littoral than in the pelagic perch. The relationship between the length of perch and the size of the mouth and fins also differed between perch from the two habitats, where the increase in the length of the pelvic fin and the area of the mouth increased faster with size in the littoral perch. Our findings show that variation in morphology between habitats differs during ontogeny in a way that corresponds to functional expectations for fish species that occupy these habitats.  相似文献   

4.
In August, growth rate of young–of–the–year (YOY) Perch In lake enclosures could be explained by both YOY density and mean cladoceran biomass, suggesting that in a lake where YOY perch are dominant, growth may be density dependent in late summer and mediated through top–down control on daphnid biomass. In June, growth rate of YOY perch could not be fully explained by YOY density or by mean cladoceran biomass, suggesting that growth and survival during the first part of the summer is negatively affected by a diet of Bosmina and cyclopoid copepods only. The experiments also suggest why YOY perch have a slow growth and a low abundance in eutrophic lakes where small zooplankton dominate. The June experiment also indicated that growth of late larval or early juvenile perch improved when a larger cladoceran became available and was included in the diet.  相似文献   

5.
Despite the common occurrence of ontogenetic niche shifts, their consequences for morphological adaptations have been little studied. To address this question, we studied morphological adaptations related to ontogenetic niche shifts in Eurasian perch (Perca fluviatilis) in eight lakes that varied in density of benthic resources and planktivorous fish biomass. Perch start to feed on pelagic zooplankton, then shift to benthic resources at intermediate sizes, and finally, when large enough, mainly feed on fish. These three functional niches over ontogeny are expected to set constraints on the morphology and size-specific growth of perch. The growth of perch was negatively related to planktivorous fish biomass in the zooplanktivorous niche, but positively related to planktivorous fish biomass in the piscivorous niche. The number of gill rakers of perch was negatively related to the biomass of planktivorous fish, providing evidence for the occurrence of character displacement as a consequence of competition in the zooplanktivorous niche. Perch in lakes with low densities of predator-sensitive macroinvertebrates had greater body height measurements and a larger mouth early during ontogeny. This pattern is suggested to be a result of a selection for increased efficiency in the benthic niche when the availability of benthic resources is low. Perch in lakes with a high biomass of planktivorous fishes had fusiform body morphology, a thicker tail and a larger mouth then the average piscivorous perch. The different responses of perch morphology to variation in the availability of benthic resources compared to variation in planktivore biomass are suggested to be partly because the availability of the former resource to a larger extent is set by abiotic conditions (humic content). We suggest that the key factors affecting size-specific growth and body morphology of perch in the system studied are the availability of resources in the benthivorous and piscivorous niches. We also provide evidence for morphological trade-offs, especially between the benthivorous and the piscivorous ontogenetic niches. Received: 6 July 1999 / Accepted: 8 September 1999  相似文献   

6.
Selection of habitat to avoid predation may affect the diet of young-of-year (YOY) lake trout (Salvelinus namaycush). YOY lake trout may use inshore habitat to avoid predation; this habitat may be sub-optimal for growth. To test this, YOY lake trout were penned in nearshore and offshore pelagic areas of two arctic lakes. Toolik Lake had a lake trout population, the other lake, S6, did not. YOY lake trout in Toolik Lake lost weight, but those offshore lost less weight. The YOY lake trout in Lake S6 gained weight and those offshore gained more weight. The primary diet item of the YOY lake trout in both lakes during this experiment was the zooplankter Diaptomis probilofensis; it was also one of the most abundant species. However, its density inshore in Lake S6 was similar to inshore and offshore densities in Toolik Lake. The increased availability of alternative zooplankton prey in Lake S6 may account for the growth differential of YOY lake trout in Lake S6 relative to Toolik Lake. Bioenergetic modeling of YOY lake trout suggests that growth similar to that in the offshore of Lake S6 would be necessary for successful recruitment. If the reduced zooplankton availability in Toolik Lake leads to the reduced growth of YOY in the inshore and offshore pelagic areas, then these fish will be more susceptable to winter predation/starvation. For YOY lake trout to survive in Toolik Lake they most likely shift to feeding on benthic prey before the end of their first summer. Dept. of Chemical Engineering  相似文献   

7.
Pär Byström  Jens Andersson 《Oikos》2005,110(3):523-536
Intraspecific competition for resources is strongly influenced by the size of competitors. In this study, we estimated the size‐scaling of the foraging capacities on zooplankton and benthic macroinvertebrates in Arctic char (Salvelinus alpinus) to link size‐dependent performance to effects from competition. The competitive interactions between two size‐classes (YOY and 1‐y) of char were then studied in a large‐scale pond experiment and in two small subarctic lakes. The attack rate function on zooplankton was hump‐shaped whereas the attack rate on benthic chironomids increased monotonically with size. The size‐scaling exponent's for zooplankton and chironomids were 0.65 and 0.30, respectively, leading to that critical resource density (CRD) and maximum growth resource density (GMRD) increases with size, suggesting an exploitative competitive advantage of small individuals over large. Correspondingly, large (1‐y) char growth was negatively affected by cohort competition whereas small (YOY) char growth was not. Diets of both size classes were dominated by macroinvertebrates with large overlap in prey size suggesting only small gape size advantages for large char. Small char fed to a larger extend on cladocerans which, due to the hump‐shaped foraging efficiency function on zooplankton, were a relatively more profitable resource for small than large char. Estimates of CRD and GmRD were in correspondence with observed growth responses and resource estimates for zooplankton, whereas for macroinvertebrates only qualitatively correspondence with foraging estimates and char performance was found. Although we were able to explain our results with exploitative competition only, we suggest a general need for size‐dependent foraging estimates on prey in more complex habitats in order to quantitatively link performance and resource abundances. Interference and size‐dependent resource use as mechanisms for observed stable population dynamics in char was not supported by this study and instead a low per capita fecundity and early cannibalism on recruits are suggested to be potential mechanisms that may stabilize char dynamics.  相似文献   

8.
Bream (Abramis brama) undergo ontogenetic diet shift from zooplankton to benthic macroinvertebrates, but the switching size may be highly variable. To unravel under what conditions bream are pelagic versus benthic foragers, we experimentally determined size‐dependent foraging capacities on three prey types from the planktivory and benthivory niche; zooplankton, visible and buried macroinvertebrates. From these data we derived predictions of size‐dependent diet preferences from estimates of prey value and competitive ability, and tested these predictions on diet data from the field. Planktivorous foraging capacity described a hump‐shaped relationship with bream length that peaked for small bream of 67 mm total length. Benthivory capacity increased with increasing bream size, irrespective if benthic prey were visible on the sediment surface or buried in the sediment. From the experimental data and relationships of metabolic demand we calculated minimum resource requirement for maintenance (MRR) for each of the prey categories used in experiments. MRR increased with bream size for both zooplankton and visible chironomids, but decreased with bream size for buried chironomids, suggesting that intermediate sized bream (120–300 mm) may be competitively sandwiched between small and large bream that are more competitive planktivores and benthivores, respectively. Prey value estimates and competitive abilities qualitatively predicted diet shift in a bream population being released from competition. Competitive release did not change the diet of the largest size‐class feeding on an optimal diet of benthic invertebrates both before and after competitive release. However, profound diet shifts towards benthic macroinvertebrates were recorded for intermediate size‐classes that fed on a suboptimal diet prior to competitive release. Thus, laboratory estimates of size‐dependent foraging capacity of bream in planktivorous and benthivorous feeding niches provided useful information on size‐specific competitive ability, and successfully predicted diet preference in the field.  相似文献   

9.
  1. Browning of waters, coupled to climate change and land use changes, can strongly affect aquatic ecosystems. Browning-induced light limitation may have negative effects on aquatic consumers via shifts in resource composition and availability and by negatively affecting foraging of consumers relying on vision. However, the extent to which light limitation caused by browning affects fish via either of these two pathways is largely unknown.
  2. Here we specifically test if fish growth responses to browning in a pelagic food web are best explained by changes in resource availability and composition due to light limitation, or by reduced foraging rates due to decreased visual conditions.
  3. To address this question, we set up a mesocosm experiment to study growth responses of two different fish species to browning and conducted an aquaria experiment to study species-specific fish foraging responses to browning. Furthermore, we used a space-for-time approach to analyse fish body length-at-age across >40 lakes with a large gradient in lake water colour to validate experimental findings on species-specific fish growth responses.
  4. With browning, we found an increase in chlorophyll a concentrations, shifts in zooplankton community composition, and a decrease in perch (Perca fluviatilis) but not roach (Rutilus rutilus) body growth. We conclude that fish growth responses are most likely to be linked to the observed shift in prey (zooplankton) composition. In contrast, we found limited evidence for reduced perch, but not roach, foraging rates in response to browning. This suggests that light limitation led to lower body growth of perch in brown waters mainly through shifts in resource composition and availability, perhaps in combination with decreased visibility. Finally, with the lake study we confirmed that perch but not roach body growth and length-at-age are negatively affected by brown waters in the wild.
  5. In conclusion, using a combination of experimental and observational data, we show that browning of lakes is likely to (continue to) result in reductions in fish body growth of perch, but not roach, as a consequence of shifts in prey availability and composition, and perhaps reduced foraging.
  相似文献   

10.
It has been hypothesized that inter-specific competition will reduce species niche utilization and drive morphological evolution in character displacement. In the absence of a competitor, intra-specific competition may favor an expansion of the species niche and drive morphological evolution in character release. Despite of this theoretical framework, we sometimes find potential competitor species using the same niche range without any partitioning in niche. We used a database on test fishing in Sweden to evaluate the factors (inter- and intraspecific competition, predation, and abiotic factors) that could influence habitat choice of two competitor species. The pattern from the database shows that the occurrence of perch and roach occupying both littoral and pelagic habitats of lakes in Sweden is a general phenomenon. Furthermore, the results from the database suggest that this pattern is due to intra-specific competition rather than inter-specific competition or predation. In a field study, we estimated the morphological variation in perch and roach and found that, individuals of both species caught in the littoral zone were more deeper bodied compared to individuals caught in the pelagic zone. Pelagic perch fed more on zooplankton compared to littoral perch, independent of size, whereas the littoral perch had more macroinvertebrates and fish in their diet. Pelagic roach fed more on zooplankton compared to littoral roach, whereas littoral individuals fed more on plant material. Furthermore, we sampled littoral and pelagic fish from another lake to evaluate the generality of our first results and found the same habitat associated morphology in both perch and roach. The results show a consistent multi-species morphological separation in the littoral and pelagic habitats. This study suggests that intra-specific competition is possibly more important than inter-specific competition for the morphological pattern in the perch-roach system.  相似文献   

11.
1. Yellow perch (Perca flavescens) are often the only surviving fish species in acidified lakes. We studied four lakes along a gradient of recovery from acidification and that had different food web complexities. All had abundant yellow perch, two had low piscivore abundance, one had a well‐established piscivore population and one was manipulated by introducing piscivorous smallmouth bass (Micropterus dolomieu). We hypothesised that there would be strong effects on perch abundance, behaviour and diet induced by the presence of piscivores. 2. In the manipulated lake, the bass reduced yellow perch abundance by 75% over a 2‐year period. Concomitantly, perch use of the pelagic habitat fell from 48 to 40%. 3. In contrast to findings from less disturbed systems, yellow perch in the littoral zone of the manipulated lake did not strongly shift from zooplankton to benthic food sources after the arrival of piscivores. Diet analysis using stable carbon isotopes revealed a strong continued reliance on zooplankton in all lakes, independent of the degree of piscivory. The failure to switch to benthos in the refuge area of the littoral zone is most likely related to the depauperate benthos communities in these formerly acidified lakes. 4. Yellow perch in lakes recovering from acidification face a considerable ecological challenge as the necessary switch to benthic diet is hindered by a low abundance of benthos. The arrival of piscivores in these recovering lakes imposes further restrictions on perch access to food items. We infer that future recovery of perch populations (and higher trophic levels) will have to be preceded by the re‐establishment of diverse benthic macroinvertebrate communities in these lakes.  相似文献   

12.
Spatial distribution of young-of-the-year (YOY) and older roach, rudd, perch and ruffe was compared in two artificial lakes with macrophytes present and absent, and a valley reservoir, using gillnets. Almost all species of interest and both age categories preferred benthic habitats. The depth distribution in benthic habitats was relatively consistent across water bodies with the highest fish densities found in the shallowest depths. In the macrophyte-rich lake, YOY roach and perch utilize the 3–6 m benthic layer the most, whereas the fish preferred the 0–3 m benthic layer in the macrophyte-poor lake and reservoir. No differences were found in the depth distribution in pelagic habitats sampled by pelagic gillnets for YOY fish between the water bodies. Older fish usually utilized the surface water layer. Macrophytes influenced the depth distribution of YOY fish in benthic habitats, where their density maximum shifted deeper in the macrophyte-rich lake when fewer macrophytes were present in the shallowest benthic depth. In lakes, YOY fish utilized a wider depth spectrum due to the deeper thermocline when compared to the reservoir. Oxygen and temperature stratification are the main factors influencing fish distribution, whereas macrophyte presence particularly influences the depth distribution of YOY fish in benthic habitats.  相似文献   

13.
Variation in growth and habitat use is closely connected to individual responses to habitat specific resource levels and predation risk. In three mountain lakes which differed in the density of young-of-the-year (YOY) arctic char ( Salvelinus alpinus ), we studied the growth, diets and habitat use of YOY char in relation to resource levels. With two model approaches, we further examined the extent to which YOY and 1-yr old char were resource limited and, if so, whether resource limitation was associated with habitat use and small char densities. Benthic prey dominated the diet of YOY char and YOY char habitat use was restricted to the near-shore habitat in all lakes. In all lakes were chironomid densities higher in the near-shore habitat than in the offshore benthic habitat whereas zooplankton densities were higher in the pelagic than in the near-shore habitat. Growth of YOY char in all lakes was close to predicted maximum growth despite large variation in YOY densities between lakes. Model results suggested that density dependent resource limitation in YOY char is unlikely to occur despite restricted near-shore habitat use. In contrast, strong density dependent resource limitation was predicted in 1-yr old char with a restricted habitat use to the near-shore habitat. Correspondingly, field data suggested that the habitat use of 1-yr old char was density dependent as the use of the offshore habitat increased earlier in time and to a larger extent at high densities. As small individuals are vulnerable to predation but constrained by their low food processing capacity relative to their encounter capacity, we suggest that resource limitation in small individuals should be less pronounced and habitat use should mainly depend on predation risk. A trade-off in habitat use between foraging gain and predation risk is therefore expected to be more likely for individuals, large enough to be resource limited but still small enough to be vulnerable to predation.  相似文献   

14.
While phenotypic responses to direct species interactions are well studied, we know little about the consequences of indirect interactions for phenotypic divergence. In this study we used lakes with and without the zebra mussel to investigate effects of indirect trophic interactions on phenotypic divergence between littoral and pelagic perch. We found a greater phenotypic divergence between littoral and pelagic individuals in lakes with zebra mussels and propose a mussel-mediated increase in pelagic and benthic resource availability as a major factor underlying this divergence. Lakes with zebra mussels contained higher densities of large plankton taxa and large invertebrates. We suggest that this augmented resource availability improved perch foraging opportunities in both the littoral and pelagic zones. Perch in both habitats could hence express a more specialized foraging morphology, leading to an increased divergence of perch forms in lakes with zebra mussels. As perch do not prey on mussels directly, we conclude that the increased divergence results from indirect interactions with the mussels. Our results hence suggest that species at lower food web levels can indirectly affect phenotypic divergence in species at the top of the food chain.  相似文献   

15.
Trait combinations that lead to a higher efficiency in resource utilization are important drivers of divergent natural selection and adaptive radiation. However, variation in environmental features might constrain foraging in complex ways and therefore impede the exploitation of critical resources. We tested the effect of water transparency on intra-population divergence in morphology of Eurasian perch (Perca fluviatilis) across seven lakes in central Sweden. Morphological divergence between near-shore littoral and open-water pelagic perch substantially increased with increasing water transparency. Reliance on littoral resources increased strongly with increasing water transparency in littoral populations, whereas littoral reliance was not affected by water transparency in pelagic populations. Despite the similar reliance on pelagic resources in pelagic populations along the water transparency gradient, the utilization of particular pelagic prey items differed with variation in water transparency in pelagic populations. Pelagic perch utilized cladocerans in lakes with high water transparency and copepods in lakes with low water transparency. We suggest that under impaired visual conditions low utilization of littoral resources by littoral perch and utilization of evasive copepods by pelagic perch may lead to changes in morphology. Our findings indicate that visual conditions can affect population divergence in predator populations through their effects on resource utilization.  相似文献   

16.
17.
Body shape of young‐of‐the‐year (YOY) perch Perca fluviatilis , and number and size of potential predators (perch and pike Esox lucius ) were compared across five lakes in northern Sweden. Body depth and dorsal fin ray length of YOY perch differed between lakes, with high relative body depth and long dorsal fin rays found in the lakes where the number of large piscivores was the highest. The most slender fish were found in the lake where the perch population had the smallest individuals and pike occurred in very low numbers. The average body mass in YOY perch from the two lakes with highest and lowest body depth, respectively, were the same, which indicates a difference between lakes in the relation between growth in length and in depth. Both body depth and fin ray length were correlated with predation risk by pike. Fin ray length was also correlated with number of piscivorous perch. Selection for different body shapes can be caused by different biotic and abiotic factors, singly or in combination, and the results from this study indicates that predation risk is one of these factors affecting body depth and fin ray length in perch.  相似文献   

18.
The foraging related capacities, energy requirements and the ability of individuals to withstand starvation are strongly dependent on body size and temperature. In this study, we estimated size-dependent foraging rates and critical resource density (CRD) in small Arctic char ( Salvelinus alpinus ) under winter conditions and compared these with previous observations under summer conditions. We investigated if starvation mortality is size-dependent in the laboratory, and we assessed the potential for winter growth and the occurrence and size dependency of winter mortality both in a large scale pond experiment and in natural lakes. The efficiency of foraging on macroinvertebrates increased with size but was lower at 4 than 12°C, still CRD was lower at 4 than 12°C as metabolic rates decreased faster than foraging efficiency with temperature. When starved, small char died before large and at rates which suggest that YOY char need to feed during winter to avoid starvation. Results from both our pond experiment and field study indicate that survival of YOY char over winter is high, despite severe winter conditions, because YOY char are able to feed and grow during winter. In seasonal environments with declining resources, the size scaling and temperature dependency of foraging and metabolic demands may provide conditions which can favour either small or large individuals. This size advantage dichotomy relates to that larger individual's by having a higher CRD are more likely to start starving, but once resource levels are below CRD for all size classes, small individuals starve to death at a higher rate. Negative size-dependent winter mortality from starvation is suggested to be more pronounced in species that are not adapted to feed at low temperatures and in species feeding on zooplankton, since zooplankton abundance, in contrast to macroinvertebrate abundance, is generally low during winter.  相似文献   

19.
20.
Summary Densities of the cladoceran, Holopedium gibberum, were manipulated in 18 enclosures containing juvenile (age 0+) yellow perch (Perca flavescens) and mean-lake densities of other zooplankton. In enclosures, where nearlake densities of all zooplankton species including Holopedium were maintained, young-of-the-year perch grew significantly heavier and longer than in experimental enclosures where Holopedium was excluded. Holopedium comprised between 15–45% of the diet (wet weight) of perch in the first 2 weeks of July in the control treatment (Holopedium at or near ambient lake densities) and only 3–7% of total biomass ingested in the experimental treatment (Holopedium density selectively reduced). Predation on Holopedium decreased dramatically after the 2nd week of July in the control treatment after which Chaoborus, chironomids, and Sida became dominant prey items (by weight) of juvenile perch. These findings suggest that growth and survivorship of age 0+ perch in Precambrian Shield lakes may be coupled to Holopedium abundance. Thus, utilization of Holopedium by young-of-the-year yellow perch may affect recruitment of this species since overwintering survivorship, range of accessible prey sizes or species, and vulnerability of juvenile perch to predation by larger fish depend on body size, which is reduced when Holopedium is excluded from the diet.  相似文献   

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