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1.
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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.  相似文献   

3.
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.  相似文献   

4.
A generalist strategy, as an adaptation to environmental heterogeneity, is common in Arctic freshwater systems, often accompanied, however, by intraspecific divergence that promotes specialization in niche use. To better understand how resources may be partitioned in a northern system that supports intraspecific diversity of Lake Trout, trophic niches were compared among four shallow‐water morphotypes in Great Bear Lake (N65° 56′ 39″, W120° 50′ 59″). Bayesian mixing model analyses of stable isotopes of carbon and nitrogen were conducted on adult Lake Trout. Major niche overlap in resource use among four Lake Trout morphotypes was found within littoral and pelagic zones, which raises the question of how such polymorphism can be sustained among opportunistic generalist morphotypes. Covariances of our morphological datasets were tested against δ13C and δ15N values. Patterns among morphotypes were mainly observed for δ15N. This link between ecological and morphological differentiation suggested that selection pressure(s) operate at the trophic level (δ15N), independent of habitat, rather than along the habitat‐foraging opportunity axis (δ13C). The spatial and temporal variability of resources in Arctic lakes, such as Great Bear Lake, may have favored the presence of multiple generalists showing different degrees of omnivory along a weak benthic–pelagic gradient. Morphs 1–3 had more generalist feeding habits using both benthic and pelagic habitats than Morph 4, which was a top‐predator specialist in the pelagic habitat. Evidence for frequent cannibalism in Great Bear Lake was found across all four morphotypes and may also contribute to polymorphism. We suggest that the multiple generalist morphs described here from Great Bear Lake are a unique expression of diversity due to the presumed constraints on the evolution of generalists and contrast with the development of multiple specialists, the standard response to intraspecific divergence.  相似文献   

5.
Komiya T  Fujita S  Watanabe K 《PloS one》2011,6(2):e17430
Divergent natural selection rooted in differential resource use can generate and maintain intraspecific eco-morphological divergence (i.e., resource polymorphism), ultimately leading to population splitting and speciation. Differing bottom environments create lake habitats with different benthos communities, which may cause selection in benthivorous fishes. Here, we document the nature of eco-morphological and genetic divergence among local populations of the Japanese gudgeon Sarcocheilichthys (Cyprinidae), which inhabits contrasting habitats in the littoral zones (rocky vs. pebbly habitats) in Lake Biwa, a representative ancient lake in East Asia. Eco-morphological analyses revealed that Sarcocheilichthys variegatus microoculus from rocky and pebbly zones differed in morphology and diet, and that populations from rocky environments had longer heads and deeper bodies, which are expected to be advantageous for capturing cryptic and/or attached prey in structurally complex, rocky habitats. Sarcocheilichthys biwaensis, a rock-dwelling specialist, exhibited similar morphologies to the sympatric congener, S. v. microoculus, except for body/fin coloration. Genetic analyses based on mitochondrial and nuclear microsatellite DNA data revealed no clear genetic differentiation among local populations within/between the gudgeon species. Although the morphogenetic factors that contribute to morphological divergence remain unclear, our results suggest that the gudgeon populations in Lake Biwa show a state of resource polymorphism associated with differences in the bottom environment. This is a novel example of resource polymorphism in fish within an Asian ancient lake, emphasizing the importance and generality of feeding adaptation as an evolutionary mechanism that generates morphological diversification.  相似文献   

6.
Organisms with complex life cycles are characterized by a metamorphosis that allows for a major habitat shift and the exploitation of alternative resources. However, metamorphosis can be bypassed in some species through a process called paedomorphosis, resulting in the retention of larval traits at the adult stage and is considered important at both micro‐ and macroevolutionary scales. In facultatively paedomorphic populations of newts, some individuals retain gills and a fully aquatic life at the adult stage (paedomorphs), while others undergo complete metamorphosis (metamorphs), allowing for a terrestrial life‐stage. Because facultative paedomorphosis affects trophic structures and feeding mechanism of newts, one hypothesis is that it may be maintained as a trophic polymorphism, with the advantage to lessen intraspecific competition during the shared aquatic life‐stage. Here, we tested this hypothesis combining stomach content data with stable isotope techniques, using carbon and nitrogen stable isotopes, in facultatively paedomorphic alpine newts Ichthyosaura alpestris. Both stomach content and stable isotope analyses showed that paedomorphs had smaller trophic niches and were more reliant on pelagic resources, while metamorphs relied more on littoral resources, corresponding to a polyphenism along the littoral–pelagic axis and the extension of the population's trophic niche to otherwise ‘underused’ pelagic resources by paedomorphs. Interestingly, stable isotopes revealed that the trophic polyphenism was less marked in males than in females and potentially linked to sexual activity. Although paedomorphosis and metamorphosis are primarily seen as results of tradeoffs between the advantages of using aquatic versus terrestrial habitats, this study provides evidence that additional forces, such as intraspecific trophic niche differences between morphs and trophic niche expansion, may play an important role in the persistence of this dimorphism in heterogeneous environments. Moreover, the different patterns found in males and females show the importance of considering sex to understand the evolutionary ecology of trophic polymorphisms.  相似文献   

7.
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   

8.
Understanding how a monophyletic lineage of a species diverges into several adaptive forms has received increased attention in recent years, but the underlying mechanisms in this process are still under debate. Postglacial fishes are excellent model organisms for exploring this process, especially the initial stages of ecological speciation, as postglacial lakes represent replicated discrete environments with variation in available niches. Here, we combine data of niche utilization, trophic morphology, and 17 microsatellite loci to investigate the diversification process of three sympatric European whitefish morphs from three northern Fennoscandian lakes. The morphological divergence in the gill raker number among the whitefish morphs was related to the utilization of different trophic niches and was associated with reproductive isolation within and across lakes. The intralacustrine comparison of whitefish morphs showed that these systems represent two levels of adaptive divergence: (1) a consistent littoral–pelagic resource axis; and (2) a more variable littoral–profundal resource axis. The results also indicate that the profundal whitefish morph has diverged repeatedly from the ancestral littoral whitefish morph in sympatry in two different watercourses. In contrast, all the analyses performed revealed clustering of the pelagic whitefish morphs across lakes suggesting parallel postglacial immigration with the littoral whitefish morph into each lake. Finally, the analyses strongly suggested that the trophic adaptive trait, number of gill rakers, was under diversifying selection in the different whitefish morphs. Together, the results support a complex evolutionary scenario where ecological speciation acts, but where both allopatric (colonization history) and sympatric (within watercourse divergence) processes are involved.  相似文献   

9.
Summary Adaptive variation can exist at a variety of scales in biological systems, including among species, among local populations of a single species and among individuals within a single population. Trophic or resource polymorphisms in fishes are a good example of the lowest level of this hierarchy. In lakes without bluegill sunfish (Lepomis macrochirus), pumpkinseed sunfish (Lepomis gibbosus) can be trophically polymorphic, including a planktivorous limnetic form found in the pelagic habitat, in addition to the usual benthic form found in the littoral zone. In this paper we examine the degree to which morphological differences between the two forms are caused by genetic differences versus phenotypic plasticity. Adults from pelagic and littoral sites in Paradox Lake, NY, were bred separately and their progeny were raised in cages both in the open water and shallow water habitats of an artificial pond. The experimental design permitted two tests of genetic differences between the breeding stocks (in open and shallow water cages, respectively) and two tests of phenotypic plasticity (in the limnetic and benthic offspring, respectively). Limnetic progeny were more fusiform than benthic progeny raised in the same habitat. In addition, progeny of both stocks displayed limnetic-type characteristics when raised in the open water and benthic-type characteristics in the shallow water. Thus, genetic differences and phenotypic plasticity both contributed to the trophic polymorphism. Phenotypic plasticity and genetic differentiation accounted for 53 and 14%, respectively, of the variation in morphology. This study addresses the nature of subtle phenotypic differences among individuals from a single population that is embedded within a complex community, a condition that is likely to be the norm for most natural populations, as opposed to very large differences that have evolved in relatively few populations that reside in species-poor environments.  相似文献   

10.
Phenotypic divergence in response to divergent natural selection between environments is a common phenomenon in species of freshwater fishes. Intraspecific differentiation is often pronounced between individuals inhabiting lakes versus stream habitats. The different hydrodynamic regimes in the contrasting habitats may promote a variation of body shape, but this could be intertwined with morphological adaptations to a specific foraging mode. Herein, I studied the divergence pattern of the European minnow (Phoxinus phoxinus), a common freshwater fish that has received little attention despite its large distribution. In many Scandinavian mountain lakes, European minnows are considered as being invasive and were found to pose threats to the native fish populations due to resource competition. Minnows were recently found to show phenotypic adaptations in lake versus stream habitats, but the question remained if this divergence pattern is related to differences in resource use. I therefore studied the patterns of minnow divergence in morphology (i.e., using geometric morphometrics) and trophic niches (i.e., using stomach content analyses) in the lake Ånnsjön and its tributaries to link the changes in body morphology to the feeding on specific resources. Lake minnows showed a strong reliance on benthic Cladocera and a more streamlined body shape with a more upward facing snout, whereas stream minnows fed on macroinvertebrates (larvae and adults) to a higher degree and had a deeper body with a snout that was pointed down. Correlations showed a significant relationship of the proportion of macroinvertebrates in the gut and morphological features present in the stream minnows. The results of this study highlight the habitat‐specific divergence pattern in morphology and resource use in this ubiquitous freshwater fish. Consequently, interspecific interactions of invasive minnows and the native fish population could differ in the respective food webs and resource competition could target different native fish species in the contrasting habitats.  相似文献   

11.
1. Fatty acids (FAs) have been widely applied as trophic biomarkers in aquatic food web studies. However, current knowledge of inter‐ and intraspecific variation in consumer FA compositions across spatial and temporal scales is constrained to a few pelagic taxa. 2. We analysed the FAs of 22 taxa of benthic macroinvertebrates, zooplankton and fish collected from the littoral, pelagic and profundal habitats of nine boreal oligotrophic lakes over spring, summer and autumn. We quantified and compared the FA variance partitions contributed by species identity (i.e. an integrative effect of phylogenetic origin, life history and functional feeding guild of individual taxa), site and season using partial redundancy analysis both on all consumers and on benthic arthropods alone. 3. Species identity alone contributed 84.4 and 72.8% of explained FA variation of all consumers and benthic arthropods, respectively. Influences of site, season and all joint effects accounted for 0–11.3% only. Fatty acid profiles of primary consumers differentiated below class level, but those of predators were distinguishable only when they became more taxonomically distinct (i.e. among classes or higher). 4. Pelagic and profundal consumers showed stronger reliance on autochthonous resources than did their littoral counterparts as reflected by their higher ω3 to ω6 FA ratios. Polyunsaturated FAs (PUFAs) were increasingly retained with trophic levels, and saturated FAs (e.g. FA 16 : 0) gradually reduced. Ecologically, this trade‐off enhances the trophic transfer efficiency and confirms the importance of PUFA‐rich autotrophs in aquatic food webs. 5. Our findings indicate strong interspecific differences in FA requirements and assimilation among aquatic consumers from a wide range of taxonomic levels, habitats and lakes. Consumers were able to maintain homoeostasis in FA compositions across spatial and temporal changes in resource FAs, but consumer homoeostasis did not limit the effectiveness of FAs as trophic biomarkers.  相似文献   

12.
Summer distributions of the invasive signal crayfish (Pacifastacus leniusculus) were investigated in relation to physicochemistry in a Kusiro Moor marsh and its inflows and outflows in northern Japan. Maximum crayfish abundance and biomass were 1.04 individuals/m2 and 3.56 g dry mass (DM)/m2 in littoral marsh habitats, and 5.84 individuals/m2 and 13.48 g DM/m2 in stream habitats. Classification tree analysis was used to predict crayfish occurrence at 102 sites from all habitats (i.e. littoral marsh, pelagic marsh and stream) while regression tree analyses were used to predict crayfish abundance at littoral marsh and stream sites separately. The classification tree showed that crayfish occurrence was primarily determined by undercut bank volume regardless of habitat identity. When undercut bank volume was <0.0054 m3, crayfish were predicted to be absent at marsh sites, but expected to occur at stream sites where pH and water temperature exceeded 6.5 and 14.3°C, respectively. The regression tree using only littoral marsh sites showed that undercut bank volume, followed by dissolved oxygen level, determined the splits of the tree. Crayfish abundance was highest when undercut bank volume was >0.61 m3, and moderately high when dissolved oxygen was >9.09 mg/l and undercut bank volume was <0.61 m3. On the other hand, the regression tree using only stream sites showed that water temperature was the major predictor that determined the splits. We discuss the roles of physicochemical factors as limiting factors of the distribution pattern of the invasive crayfish.  相似文献   

13.
  1. Intraspecific trophic variability has important ecological and evolutionary implications, and is driven by multiple interacting factors. Functional traits and environmental conditions are important in mediating the trophic niche of individuals because they determine their ability to consume certain prey, their energetic requirements, and resource availability. In this study, we aimed at investigating the interacting effects of functional traits and environmental conditions on several attributes of trophic niche in natural populations.
  2. Here, we quantified intraspecific variability in the trophic niche of 12 riverine populations of European minnow (Phoxinus phoxinus) using stable isotope analyses. Functional traits (i.e. morpho-anatomical traits) and environmental conditions (i.e. upstream–downstream gradient, forest cover) were quantified to identify the determinants of (1) trophic position and resource origin, (2) trophic niche size, and (3) trophic differentiation (β-diversity) among populations.
  3. We demonstrated that trophic position and resource origin covaried with functional traits related to body size and locomotion performance, and that the strength and shape of these relationships varied according to local environmental conditions. The trophic niche size also differed among populations, although no determinant was identified. Finally, trophic β-diversity was correlated to environmental differentiation among sites.
  4. Overall, the determinants of intraspecific variability in trophic niche appeared highly context-dependent, and related to the interactions between functional traits and environmental conditions. Because populations are currently facing important environmental changes, understanding this context-dependency is important for predicting food web structure and ecosystem dynamics in a changing world.
  相似文献   

14.
15.
Trade-offs in foraging efficiency leading to divergent natural selection between and within populations exploiting different resources are thought to be a primary cause of trophic polymorphism. In this study we focused on the trade-offs in foraging efficiency and growth in a polymorphic perch population. Specifically, we related habitat-specific growth and diet of perch to perch morphology. In a subsequent laboratory study we experimentally tested the trade-off by testing the efficiency of perch with different morphology feeding on pelagic ( Daphnia sp., Chaoborus sp.) and littoral (mayfly larvae) food resources. The feeding performance was tested in different physical environments to see if we could predict growth patterns in the field based on foraging rate and behavior of perch.
In the field study, we found that the perch from the littoral and the pelagic zones differed in both morphology and diet. Within the littoral zone the deeper-bodied individuals grew faster compared to the more streamlined individuals, whereas the opposite pattern was found in the pelagic zone. In the aquarium experiments, perch from the littoral zone had higher capture rates on the pelagic prey types in vegetation trials and on mayfly larvae in both open water and vegetation trials. The pelagic perch had higher capture rates on the pelagic prey types in open water trials. The littoral perch had lower search velocity than the pelagic perch in open water trials whereas the opposite pattern was found in vegetation trials. The attack velocity of the pelagic perch was also higher than that of the littoral perch independent of vegetation structure. Our results suggest that there is a functional trade-off between performance in alternate habitats and general body form in perch. Such trade-offs may promote divergent natural selection and could be the mechanism that give rise to and upholds the pattern in the field.  相似文献   

16.
Two reproductive isolated morphs of Arctic charr (Salvelinus alpinus), termed profundal and littoral charr according to their different spawning habitats, co-occur in the postglacial lake Fjellfr?svatn in North Norway. All profundal charr live in deep water their entire life and have a maximum size of 14cm, while the littoral charr grow to 40cm. Some small and young littoral charr move to the profundal zone in an ontogenetic habitat shift in the ice-free season and the rest of the population remains in epilimnic waters. The two morphs had different diet niches in the profundal zone: the profundal charr ate typical soft-bottom prey (chironomid larvae, pea mussels and benthic copepods), while the young littoral charr mainly consumed crustacean zooplankton. In four other lakes without a profundal morph (i.e. monomorphic populations), young charr also performed ontogenetic habitat shifts to the profundal zone and fed on zooplankton. The profundal morph of Fjellfr?svatn therefore utilize a food resource niche that neither the littoral morph nor comparable monomorphic populations exploit. This suggests that intraspecific resource competition has driven incipient ecological speciation of the profundal charr of Fjellfr?svatn. The exploitation of the soft-bottom resources by the profundal charr supports earlier experimental findings that the profundal morph is genetically different in trophic behaviour and morphology. The sympatric ecological divergence within the profundal habitat is possible because unexploited food resources (soft-bottom profundal prey) are available. Apparently, this represents a case of incipient segregation by expansion to new resource types (niche invasion), and not by subdivision of one broad ancestral niche.  相似文献   

17.
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   

18.
Trophically dimorphic pumpkinseed populations were investigated in five lakes in Ontario, Canada to determine: (1) whether the morphological traits that distinguish littoral from pelagic forms are consistent among populations; (2) whether the pharyngeal jaw apparatus is diminished in pelagic pumpkinseeds because of a lower proportion of hard-bodied prey in their diets; (3) whether there is life-history differentiation between littoral and pelagic forms. Pumpkinseeds captured from the littoral and pelagic zones differed significantly in morphology in four of the five lakes, but the number of external measures that differed significantly within the differentiated populations ranged from zero to six. Littoral pumpkinseeds generally had longer heads, more rearward placement of dorsal and pectoral fins, longer pectoral fins and deeper bodies than pelagic pumpkinseeds. Littoral and pelagic pumpkinseeds were more readily differentiated by internal morphometric measures, with littoral individuals having larger molars and wider spacing between gill rakers than pelagic individuals. Littoral and pelagic differences in age at maturity, size at maturity and gonado-somatic index were present only in one of three populations assessed for these traits, suggesting that morphological divergence is not necessarily accompanied by life-history differentiation.  相似文献   

19.
20.
Predators should stabilize food webs because they can move between spatially separate habitats. However, predators adapted to forage on local resources may have a reduced ability to couple habitats. Here, we show clear asymmetry in the ability to couple habitats by Eurasian perch—a common polymorphic predator in European lakes. We sampled perch from two spatially separate habitats—pelagic and littoral zones—in Lake Erken, Sweden. Littoral perch showed stronger individual specialization, but they also used resources from the pelagic zone, indicating their ability to couple habitats. In contrast, pelagic perch showed weaker individual specialization but near complete reliance on pelagic resources, indicating their preference to one habitat. This asymmetry in the habitat coupling ability of perch challenges the expectation that, in general, predators should stabilize spatially separated food webs. Our results suggest that habitat coupling might be constrained by morphological adaptations, which in this case were not related to genetic differentiation but were more likely related to differences in individual specialization.  相似文献   

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