高营养级捕食者在浅水湖泊沿岸带与敞水区能量耦合的维持作用

研究利用保安湖沿岸带与敞水区两种生境中高营养级捕食者(达氏鲌与红鳍原鲌)的碳(δ13C)、氮(δ15N)稳定同位素数据,通过稳定同位素质量平衡混合模型、非度量多维尺度分析(NMDS)等方法,分析了两种生境中鲌类食物来源的差异.结果表明,两种生境中鲌类食物来源基本一致,且食物来源较为广泛,包括沿岸带饵料鱼类、敞水区饵料鱼...     

Genetic and morphological variation between two forms of lacustrine brook charr

Genetic data suggest that the littoral and pelagic forms of brook charr Salvelinus fontinalis in Lake Bondi are two populations with partial reproductive isolation and non-random mating. Genetic differentiation between the two groups was supported by differences in allele frequencies and by deviation from Hardy-Weinberg equilibrium when the two groups were pooled; no such deviation was observed when fish were divided into littoral and pelagic groups. In contrast to Lake Bondi, no clear evidence of genetic differentiation was observed in Lake Ledoux. Discriminant function analyses of morphological characters support the existence of littoral and pelagic groups in Bondi and Ledoux Lakes. In Lake Bondi, the two groups differed significantly in two shape variables (pelagic fish had shorter dorsal fins, and longer body length posterior to the dorsal fin than littoral ones) whereas in Lake Ledoux, the groups differed in four shape variables (pelagic fish had shorter pectoral fins, shorter dorsal fins, and a shorter and higher caudal peduncle than littoral ones). Discriminant analyses of these characters were effective in reclassifying fish into their appropriate groups in both populations, with an efficiency of 78% for juveniles in Lake Bondi and 69% for adults in Lake Ledoux. Differences in morphology between the two forms are consistent with adaptations required to forage in each zone, i.e. benthic form in the littoral zone and planktivorous form in the pelagic zone.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Trophic polymorphism in introduced pumpkinseed (<Emphasis Type="Italic">Lepomis gibbosus</Emphasis>) inhabiting Iberian reservoirs

Introduced pumpkinseed in Iberian reservoirs display marked external morphological differentiation along two simultaneous dimensions of flow and trophic structure. We assessed the degree of internal morphological differentiation using gill rakers, pharyngeal jaws and the levator posterior muscle among pumpkinseeds occupying four different habitats and determined whether prey consumption accounted for any discernible differences in feeding structures among ecomorphs. Results showed significant differentiation by habitat based on pharyngeal muscle and jaw dimensions in all study reservoirs, with pelagic pumpkinseeds having smaller jaws than littoral pumpkinseeds in four of the five reservoirs. Gill rakers, however, differentiated morphs in only three of the five reservoirs, corresponding to differences in zooplankton consumption among pelagic and littoral individuals in those reservoirs. Based on all internal morphological traits, greater divergence was seen along the littoral-pelagic trophic axis in the lacustrine zones of reservoirs compared to the fluvial zone. Overall differences noted in internal morphology are likely the result of phenotypic plasticity; the ability of this species to readily adapt to changing physical environments may explain the success of the pumpkinseed in its introduced range.     

Deterioration of cichlid habitat by increased sedimentation in the rocky littoral zone of Lake Malawi

Lake Malawi has been affected by soil erosion across the catchment area. We experimentally investigated habitat preferences of cichlids in the rocky littoral zone to examine the influence of increased sedimentation on bottom-dwelling cichlids. Numbers of individuals and species, as well as the feeding rate of some of these cichlids, increased following the clearing of sediment. The results showed that these cichlids preferred sediment-free bottom and that the sedimentation causes deterioration of their habitats. Studies on the consequences of the habitat deterioration caused by increased sedimentation are needed in the management of these most diverse freshwater fish communities.     

Diet overlap between the newly introduced <Emphasis Type="Italic">Lamprichthys tanganicanus</Emphasis> and the Tanganyika sardine in Lake Kivu,Eastern Africa

This study evaluates the possible competition for food between Lamprichthys tanganicanus, recently introduced in Lake Kivu, and Limnothrissa miodon, which has been the basis of the pelagic fishery in this lake for several decades. Since 2006, L. tanganicanus has expanded in the lake and its numbers have increased in the captures, raising concern for the sardine fishery. We carried out a 2-year monthly survey, based on experimental captures in littoral and pelagic stations, which demonstrated the invasive dispersal of L. tanganicanus in littoral and pelagic waters. The diet of both species was determined on the basis of gut content analyses, taking into account the influence of site and season, and a diet overlap index was calculated. In the pelagic zone, where almost all size classes of both species were present and essentially fed upon mesozooplankton, the diet overlap was high. This situation stems from the fact that L. tanganicanus has colonized the pelagic zone in Lake Kivu, likely in search for more abundant mesozooplankton. Inshore, the diet overlap between the two species was lower, as L. tanganicanus consumed a broad range of food, whereas L. miodon strongly selected insects and, chiefly for the largest specimens, fishes. These results suggest a likelihood of interspecific competition, particularly offshore, where mesozooplankton is the main available food type, and call for further monitoring of the sardine fishery, to assess a possible impact of the invader.     

Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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Trophic ecology of the Lake Superior wave zone: a stable isotope approach

Stable carbon and nitrogen isotope ratio analyses were used to characterize the primary energy sources and trophic positions of 16 common Lake Superior wave zone invertebrate species. Isotope data from six tributary species that were taxonomically and ecologically matched with common wave zone species revealed broad energetic separation between these similarly structured benthic food webs. Previously published stable isotope data for Lake Superior wetland and pelagic food webs were used to assess the relative importance of inter-habitat energy flow within the Lake Superior ecosystem. The results of these comparisons indicate that the Lake Superior wave zone is energetically distinct from its tributaries, wetlands, and to a lesser extent from its vast pelagic realm. This information and approach should prove useful in future studies on the bioenergetics of inter-zonal migrants and other species that forage in multiple habitats within the lake and also in revealing energetic connections among terrestrial, riverine, littoral, and pelagic food webs in the coastal ecosystems of Lake Superior.     

Habitats and Littoral Zone Fish Community Structure of Two Natural Lakes in Southeast Brazil

The fish community and habitat structure in the littoral zone of lakes Pedra and Hortência, middle Rio Doce Valley, Brazil, were investigated in three sampling periods from August 1992 to May 1993. A total of 9106 fishes were collected, including 11 species in Lake Hortência and 12 species in Lake Pedra. Diversity was higher in Lake Hortência, although total fish abundance was the same for both lakes, with no significant temporal differences. The length distribution of the majority of fish species was similar between lakes. Geophagus brasiliensis was the dominant species (number/biomass) in all habitats of the two lakes in all three sampling periods. A striking difference in total fish abundance in relation to habitat types, which we attribute to physical variables, especially the abundance of macrophytes was observed. Negative impacts attributable to the local Eucalyptus sp. plantations were not detected in the fish community structure of Lake Hortência, which is probably due to the maintenance of the riparian vegetation contributing to similar physical environmental conditions in the littoral zone and high similarity in fish community structure between the two lakes. However, further studies on fish community structure and potential Eucalyptus impacts are suggested as necessary to improve conservation measures of the lakes and their fish faunas.     

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

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Effects of habitat and food resources on morphology and ontogenetic growth trajectories in perch

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.     

Morphological differentiation in introduced pumpkinseed Lepomis gibbosus (L.) occupying different habitat zones in Portuguese reservoirs

Introduced pumpkinseed Lepomis gibbosus sampled from four habitat zones (fluvial pelagic, fluvial littoral, lacustrine pelagic and lacustrine littoral) in three Portuguese reservoirs were used to test the hypotheses that habitats with the least similar characteristics will show the most differentiation, and that morphological differences will relate to functional adaptations to flow and trophic habitats. Results from discriminant function analysis and ANCOVA showed that there were significant differences in external morphology in pumpkinseed captured from the four habitat zones in all three reservoirs. Littoral and pelagic differentiation was stronger than fluvial and lacustrine differentiation in all of the reservoirs, and the most significant variable that differentiated pumpkinseed from the littoral and pelagic habitats was body depth. The illustration of external morphological differentiation in pumpkinseed along both habitat dimensions demonstrates the high degree of morphological plasticity of this introduced species.     

Chaos and stability of age-0 fish assemblages in a temperate deep reservoir: unpredictable success and stable habitat use

Large year-to-year variability in different fish species recruitment has been confirmed by previous studies while diurnal patterns of occupation in two basic reservoir habitats (pelagic and littoral) by different age-0 fish species in late summer are still unclear. Data collected over an 11-year period regarding late-summer age-0 fish assemblages in pelagic and littoral habitats of a reservoir were used to test the recruitment instability and to investigate diurnal habitat use. Trawling was conducted in the pelagic habitat at night while beach seining was conducted in the littoral habitat during day and night. Fluctuations in age-0 fish abundance and species composition were observed with both sampling methods; however, the following spatio-temporal patterns were relatively stable in most investigated years: (1) pelagic species (pikeperch; Sander lucioperca, small perch; Perca fluviatilis, bream; Abramis brama at night), (2) littoral species (large perch, asp; Leuciscus aspius, dace; Leuciscus leuciscus), (3) migratory species likely performing diel horizontal migrations (bleak; Alburnus alburnus), (4) species abundant in the littoral habitat both during day and night and also in pelagic habitat at night (roach; Rutilus rutilus) and (5) species detected in both habitats exclusively at night (ruffe; Gymnocephalus cernuus).     

Stable resource polymorphism along the benthic littoral–pelagic axis in an invasive crayfish

Although intraspecific variability is now widely recognized as affecting evolutionary and ecological processes, our knowledge on the importance of intraspecific variability within invasive species is still limited. This is despite the fact that understanding the linkage between within‐population morphological divergences and the use of different trophic or spatial resources (i.e., resource polymorphism) can help to better predict their ecological impacts on recipient ecosystems. Here, we quantified the extent of resource polymorphism within populations of a worldwide invasive crayfish species, Procambarus clarkii, in 16 lake populations by comparing their trophic (estimated using stable isotope analyses) and morphological characteristics between individuals from the littoral and pelagic habitats. Our results first demonstrated that crayfish occured in both littoral and pelagic habitats of seven lakes and that the use of pelagic habitat was associated with increased abundance of littoral crayfish. We then found morphological (i.e., body and chelae shapes) and trophic divergence (i.e., reliance on littoral carbon) among individuals from littoral and pelagic habitats, highlighting the existence of resource polymorphism in invasive populations. There was no genetic differentiation between individuals from the two habitats, implying that this resource polymorphism was stable (i.e., high gene flow between individuals). Finally, we demonstrated that a divergent adaptive process was responsible for the morphological divergence in body and chela shapes between habitats while difference in littoral reliance neutrally evolved under genetic drift. These findings demonstrated that invasive P. clarkii can display strong within‐population phenotypic variability in recent populations, and this could lead to contrasting ecological impacts between littoral and pelagic individuals.     

Trophic polymorphism in brook charr revealed by diet, parasites and morphometrics

Stomach contents, parasite assemblages and morphometrics were compared in brook charr Salvelinus fontinalis from the littoral and pelagic zone of two adjacent lakes on the Canadian Shield. In lac Baie des Onze Îles, fish from the littoral zone had greater abundance of benthic prey in their stomach and were more heavily infected by parasites that use intermediate hosts associated with the littoral zone than fish captured in the pelagic zone. Littoral and pelagic brook charr from this lake also differed in regard to body shape and fin length, with each group being anatomically adapted to exploit their respective habitats. The highly significant correlation between morphometric and parasite canonical scores supports the hypothesis of functional diversification of individuals within lac Baie des Onze Îles. While fish from littoral and pelagic zones of lac Caribou did not differ in terms of diet, parasite assemblages or morphometrics, they were different to fish from lac Baie des Onze Îles in that they were less frequently infected with parasites that use gastropods as intermediate hosts, and had shorter pectoral fins. The inter-lake comparisons suggested that parasite assemblages and morphometrics of brook charr reflected the dominance of the limnetic and littoral habitats in lacs Caribou and lac Baie des Onze Îles, respectively.     

Intraspecific competition drives multiple species resource polymorphism in fish communities

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.     

Determinants of the distribution of juvenile fish in the littoral area of a shallow lake

1. Research has often focused on the pelagic areas of lakes; the littoral zone has received less attention. The few studies concerning fish distribution in littoral habitats have concentrated on stands of submersed macrophytes, whereas other littoral habitat types have seldom been investigated. 2. This study aimed to predict the occurrence of juvenile fish in several littoral habitats of a shallow lake as a function of food availability, complexity of habitat structure, water depth and substrate. Habitats comprising reed, woody structures, and two open water areas differing in depth were sampled for fish and invertebrate biomasses on two shores, over 6 months and during both daylight and at night. 3. The juvenile fish community consisted almost exclusively of 0+ and 1+ roach and perch. There was a strong diel component in habitat use, with a predominant occurrence of fish in complex habitats (mainly woody structures) during the day, and a partial migration towards the open habitats at night, more strongly expressed in roach than in perch. 4. The diet of all fish groups was relatively constant over the seasonal cycle, and was independent of habitat. There was a higher degree of planktivory in roach than in perch, but both species fed on benthic macroinvertebrates to a substantial extent. 5. According to a logistic regression model, the biomass of potential food organisms in the different habitats had little predictive effect on the spatial distribution of the fish, whereas the structural complexity of the habitats combined with the diel cycle explained about 28% of the occurrence patterns in 0+ and 1+ perch and 1+ roach.     

Discrete foraging niches promote ecological, phenotypic, and genetic divergence in sympatric whitefish (Coregonus lavaretus)

Natural populations often vary in their degree of ecological, morphological and genetic divergence. This variation can be arranged along an ecological speciation continuum of increasingly discrete variation, with high inter-individual variation at one end and well defined species in the other. In postglacial fishes, evolutionary divergence has commonly resulted in the co-occurrence of a pelagic and a benthic specialist. We studied three replicate lakes supporting sympatric pelagic and benthic European whitefish (Coregonus lavaretus (L.)) morphs in search for early signs of possible further divergence into more specialized niches. Using stomach content data (recent diet) and stable isotope analyses (time-integrated measure of trophic niche use), we observed a split in the trophic niche within the benthic whitefish morph, with individuals specializing on either littoral or profundal resources. This divergence in resource use was accompanied by small but significant differences in an adaptive morphological trait (gill raker number) and significant genetic differences between fish exploiting littoral and profundal habitats and foraging resources. The same pattern of parallel divergence was found in all three lakes, suggesting similar natural selection pressures driving and/or maintaining the divergence. The two levels of divergence (a clear and robust benthic – pelagic and a more subtle littoral – profundal divergence) observed in this study apparently represent different stages in the process of ecological speciation.     

Some like it deep: Intraspecific niche segregation in ruffe (Gymnocephalus cernua)

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