Rearing environment affects spatial learning in juvenile Chinook salmon Oncorhynchus tshawytscha

We tested the prediction that a complex physical rearing environment would enhance short-term spatial memory as assessed by learning ability in a spatial navigation task in juvenile Chinook salmon Oncorhynchus tshawytscha. We reared fish in two low-density treatments, where fish were either in bare fiberglass tanks (bare) or in tanks with physical structure (complex). We also tested conventionally reared high-density hatchery fish to compare with these other experimental treatments. Our reason for including this third hatchery treatment is that the two low-density treatments, aside from the manipulation of structure, followed a rearing programme that is designed to produce fish with more wild-like characteristics. We tested individually marked fish for seven consecutive days and recorded movement and time to exit a testing maze. Stimulus conspecific fish outside the exit of the maze provided positive reinforcement for test fish. Fish from the bare treatment were less likely to exit the start box compared with fish in the complex and hatchery treatments. However, fish in the hatchery treatment were significantly more likely to exit the maze on their own compared with both the bare and complex treatments. Hatchery fish effectively learned the task as shown by a decrease in the number of mistakes over time, but the number of mistakes was significantly greater on the first day of trials. Increasing habitat complexity with structure may not necessarily promote spatial learning ability, but differences between hatchery and experimental treatments in rearing density and motivation to be near conspecifics likely led to observed behavioural differences.     

Predation risk and competition affect habitat use of adult perch,Perca fluviatilis

The aim of this study was to examine how the presence of a predator and an interspecific competitor influence the habitat use of adult perch (Perca fluviatilis; size: 15.1 ± 0.5 cm) when given the choice between two adjacent habitats. By conducting aquarium experiments, the habitat occupancy of P. fluviatilis was documented in the presence and absence of a predator (pike Esox lucius; size: 25.4 ± 2.1 cm) and a potential competitor (ruffe Gymnocephalus cernuus; size: 14.1 ± 0.3 cm) fish species. Two P. fluviatilis individuals generally shared the same habitat. In the presence of a conspecific, P. fluviatilis favoured the structurally more-complex, artificial macrophyte habitat over the less-structured rock and sand habitat, which in turn were used equally. In the predator- and competitor treatments, P. fluviatilis seemed to adapt their habitat use to the habitat occupancy of E. lucius and G. cernuus in the Macrophyte vs. Rock and, in the predator treatment, also in the Macrophyte vs. Sand habitat combination, by increasingly occupying a habitat that was used less by the predator or competitor species, respectively. This behaviour suggests that P. fluviatilis tried to avoid the other fish species by choosing a, in some cases less preferred, predator- or competitor-free habitat. This study emphasizes the importance of biological interactions illustrated by the potential of predation risk and competition to structure fish communities by influencing habitat use at small spatial scales.     

Habitat-mediated survival of newly settled red king crab in the presence of a predatory fish: Role of habitat complexity and heterogeneity

Red king crab (RKC) (Paralithodes camtschaticus) are generally associated with structurally complex habitats during the first 2 years of benthic life. In this first experimental laboratory study with a fish predator, survival of newly settled juvenile RKC was tested in eight different habitat treatments with varying amounts and types of physical structure, open sand, gravel bottom, and habitat islands. Video observations provided insights on habitat-mediated interactions between Pacific halibut predators (Hippoglossus stenolepis) and crab prey. Survival of RKC increased with amount of physical structure and was highest in the most heterogeneous habitat and in habitats characterized by high density patches. Predator activity decreased with increasing amount of structure, and attacks on RKC were correlated with predator activity. Low survival in open sand habitat was associated with both high attack rate and high capture success (captures per attack). Lower levels of capture success did not vary among the habitats containing algae and other complex physical structures, but attack rates declined with increasing amount of structure, and encounter rate (i.e., prey detection and attack) was the primary determinant of mortality. RKC were capable of detecting predators and adjusted their behavior to avoid predation by sheltering in dense microhabitat patches. Successful stock enhancement for greatly reduced populations of RKC in the Gulf of Alaska will depend upon placing seed stock in habitats with abundant protective habitat, and high quality microhabitats may serve as well as continuous cover.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

Short‐term Costs and Benefits of Habitat Complexity for a Territorial Fish

An increase in habitat complexity is thought to decrease visibility and the territory size of visually oriented animals. Hence, the addition of physical structure has been viewed as a useful restoration technique to increase the density of territorial species, particularly in stream fishes. However, a decrease in territory size may have a negative effect on the fitness of individual organisms. We attempted to quantify some of the positive and negative effects of increasing habitat structure on the behaviour and growth rate of wild young‐of‐the‐year (YOY) Atlantic salmon. Fish were exposed to one of two habitat treatments in mesh enclosures in Catamaran Brook, New Brunswick: a gravel substrate (low complexity) or a gravel substrate with boulders added (high complexity). Wild‐caught individuals were tagged, weighed and measured before being stocked at densities of 1·m^?2 for 7‐d trials. While fish from high‐complexity treatments benefited from lower rates of aggression, they also had lower foraging rates and smaller territories compared to those in low‐complexity treatments. Specific growth rate, however, did not differ significantly between treatments. While the addition of structure to a habitat may be beneficial at the population level in terms of an increase in population density, our results suggested that individual fish may pay some short‐term costs in these environments. Further research is needed to evaluate the longer term costs and benefits of adding structure to improve the habitat quality for stream salmonids.     

Testes Investment along a Vertical Depth Gradient in an Herbivorous Fish

Males are expected to adjust testes investment according to the varying level of sperm competition that they experience. Spatial and temporal variation in population density likely influences sperm competition. In herbivorous aquatic organisms, densities often decrease along a vertical depth gradient, because their food is photosynthetic and thus becomes less abundant in deeper regions where less light penetrates. This decrease should be dramatic on a steep slope, which allows testing of the association between density and testes investment at the within‐population level. We tested the effect in the socially monogamous herbivorous cichlid fish Variabilichromis moorii living on a steep slope in Lake Tanganyika. We examined competitor density and food abundance as ecological factors, territory defense behaviors and phenotypic traits (testes investment and somatic investment), and compared them between shallow (4–6 m depth) and deep habitats (10–13 m depth) separated by several dozen meters. We found that food availability drastically decreased with increasing depth and that V . moorii was much more abundant in shallower habitats. Males in shallower habitats were in better physical condition (based on fat and liver mass) despite experiencing greater costs in terms of territory defense. Testes investment differed in areas with different competitor density and food abundance along a vertical depth gradient, but competitor density was the most explanatory factor of the difference. This suggests that this herbivorous fish would change testes investment in response to population density.     

Habitat structural complexity mediates food web dynamics in a freshwater macrophyte community

A considerable amount of research has investigated the influence of habitat structure on predator success, yet few studies have explored the implications for community structure and food-web dynamics. The relative importance of macrophyte structure and fish predation on the composition of the macroinvertebrate and periphyton communities in a lowland river was investigated using a multifactorial caging experiment. We hypothesised that: (1) fish predators are less effective in a more structurally complex macrophyte analogue; (2) strong direct and indirect effects of fish predators (e.g. trophic cascades) are less likely to occur in a structurally complex habitat; and (3) the strength of these patterns is influenced by the composition of the prevailing community assemblage. We measured the abundance and composition of the macroinvertebrate and periphyton communities associated with three different-shaped macrophyte analogues, under different fish predator treatments and at different times. Macrophyte analogue architecture had strong, consistent effects on both the macroinvertebrate and periphyton communities; both were most abundant and diverse on the most structurally complex plant analogue. In contrast, the fish predators affected only a subset of the macroinvertebrate community and there was a suggestion of minor indirect effects on periphyton community composition. Contrary to expectations, the fish predators had their strongest effects in the most structurally complex macrophyte analogue. We conclude that in this system, macrophyte shape strongly regulates the associated freshwater assemblage, resulting in a diverse community structure less likely to exhibit strong effects of fish predation.     

Competitive exclusion after invasion?

The ‘Competitive Exclusion Principle’ is a foundation stone in the understanding of interspecific competition and niche relationships between species. In spite of having the status of a biological law, the principle has limited empirical support. In this study, we document strong effects of competition from the invading fish species vendace Coregonus albula over a 14-year period in the sub-arctic Pasvik watercourse. The native d.r. whitefish, that shared food and habitat niche with the invader, was displaced from its original niche and showed a more than 90% decline in population density over the study period. The study thus provides a unique record of how an exotic fish species excludes a native species from its original niche. Our data support the competitive exclusion principle, but also indicate that the vulnerability of the inferior competitor depends on a lack of alternative resources and on indirect ecological interactions.     

Alternative forms of competition and predation dramatically affect habitat selection under foraging--predation-risk trade-offs

Habitat selection under foraging—predation-risk trade-offshas been a frequent topic of interest to theoretical behavioraland evolutionary ecologists. However, most habitat selectionmodels assume that individuals compete exploitatively for resourcesand that predation is either density independent or dilutedcompletely by competitor number, despite empirical evidencethat other forms of competition and predation also occur innature. I developed an individual-based model for studyingthe effects of alternative forms of competition and predationon the process of habitat selection under foraging—predation-risktrade-offs. To make the model more relevant to natural populations,I assumed that individuals vary continuously in traits relatedto competitive ability and vulnerability to predation and allowed resources and predators to be distributed across more than twohabitats. The results of my investigation demonstrate thatthe predicted pattern of habitat selection can be affecteddramatically by the form predation is assumed to take. Whenpredation is density dependent or frequency dependent, individuals will tend to be distributed across habitats according to theirabsolute vulnerability to predation. In contrast, when predationis density dependent or vulnerability dependent, individualswill tend to segregate by competitive ability. Whether oneassumes that individuals compete for resources via exploitationor interference also influences the predicted pattern of habitat selection. In general, interference competition results in amore even distribution of competitors across habitats.     

Environmental drivers of demographics, habitat use, and behavior during a post-Pleistocene radiation of Bahamas mosquitofish (Gambusia hubbsi)

A fundamental goal of evolutionary ecology is to understand the environmental drivers of ecological divergence during the early stages of adaptive diversification. Using the model system of the post-Pleistocene radiation of Bahamas mosquitofish (Gambusia hubbsi) inhabiting blue holes, we used a comparative field study to examine variation in density, age structure, tertiary (adult) sex ratio, habitat use, as well as adult feeding and social behaviors in relation to environmental features including predation risk, interspecific competition, productivity (e.g. chlorophyll a, zooplankton density), and abiotic factors (e.g. salinity, surface diameter). The primary environmental factor associated with ecological differentiation in G. hubbsi was the presence of piscivorous fish. Gambusia hubbsi populations coexisting with predatory fish were less dense, comprised of a smaller proportion of juveniles, and were more concentrated in shallow, near-shore regions of blue holes. In addition to predation risk, the presence of a competitor fish species was associated with G. hubbsi habitat use, and productivity covaried with both age structure and habitat use. Feeding and social behaviors differed considerably between sexes, and both sexes showed behavioral differences between predator regimes by exhibiting more foraging behaviors in the absence of predators and more sexual behaviors in their presence. Males additionally exhibited more aggressive behaviors toward females in the absence of predators, but were more aggressive toward other males in the presence of predators. These results largely matched a priori predictions, and several findings are similar to trends in other related systems. Variation in predation risk appears to represent the primary driver of ecological differentiation in this system, but other previously underappreciated factors (interspecific competition, resource availability) are notable contributors as well. This study highlights the utility of simultaneously evaluating multiple environmental factors and multiple population characteristics within a natural system to pinpoint environmental drivers of ecological differentiation.     

Density‐dependent effects of non‐native brown trout Salmo trutta on the species–area relationship in stream fish assemblages

The spatial scale and density‐dependent effects of non‐native brown trout Salmo trutta on species richness of fish assemblages were examined at 48 study sites in Mamachi Stream, a tributary of Chitose River, Hokkaido, Japan. The density of age ≥1 year S. trutta was high in the upstream side of the main stem of Mamachi Stream. Fish species richness increased with increasing area of study sites (habitat size), but the increasing magnitude of the species richness with area decreased with increasing age of ≥1 year S. trutta density. The relationships between age ≥1 year S. trutta, however, and presence–absence of each species seemed to be different among species. Species richness was also determined by location and physical environmental variables, i.e. it was high on the downstream side and in structurally complex environments.     

Structural complexity in relation to the habitat preferences,territoriality, and hatchery rearing of juvenile China rockfish (<Emphasis Type="Italic">Sebastes nebulosus</Emphasis>)

Conservation efforts require an understanding of the basic behavior and ecology of target species. However, limited information exists for a wide range of taxa, including declining species of rockfish (genus Sebastes). First, we observed captive juvenile China rockfish (S. nebulosus) to determine how they interact with their environment and conspecifics. Juveniles exhibited site fidelity and territoriality. These aggressive interactions occurred within the context of size-biased dominance, centered on competition for structurally complex habitat. Given the apparent importance of structure and the absence of structure in typical hatchery environments, we then asked how the absence of structure affects future behavior. When barren-reared and structure-reared juveniles were combined into a structurally complex aquarium, barren-reared fish displayed less structure use and less site fidelity than structure-reared fish. However, after 1 to 2 weeks, barren-reared fish began to use structure and showed site fidelity that eventually equaled that of structure-reared juveniles, showing that those behavioral effects of the rearing environment were not permanent. Though these short-term effects may still impact survival after hatchery release, we were unable to detect significant effects on vulnerability to a predator (lingcod, Ophiodon elongatus) in laboratory trials.     

The influence of competitor density on space use in juvenile striped plateau lizards (Sceloporus virgatus)

To investigate the role of competitor density in influencing space use patterns of juvenile striped plateau lizards (Sceloporus virgatus), a density manipulation experiment was conducted within large (800 m²) field enclosures treated with low (20) and high (60) densities of hatchlings. Enclosures were monitored for 10 months, after which the experiment was replicated. Home range and core area sizes after release of lizards and initial establishment in the fall were significantly reduced in the high-density treatments; fall home range shape, measured as the perimeter-to-area ratio, was significantly reduced in low-density treatments; no significant differences were detected between treatments in core area shape or overlap. During the spring/early summer activity season after the lizard's first winter, no significant differences between density treatments were detected for any of these variables, as enclosure densities had converged between treatments. Individuals in high-density enclosures had increased their space use as competitor density had declined. These results illustrate that competitor density has significant influence on space use by juvenile lizards.     

Effect of competitor abundance on feeding territoriality in a grazing fish,the ayuPlecoglossus altivelis

The grazing fish, ayu,Plecoglossus altivelis Temminck & Schlegel, establishes feeding territorialiry during the young stage. The population density fluctuates from year to year by more than a hundredfold, but the determinant of territory size is less well known. The feeding territoriality of ayu was examined under simulated habitat conditions where fish density was manipulated and food resources were renewable. Fish competed for algae attached to the substrata and were divided into residents with territories, and floaters without territories. By experimental alteration of fish density the number of residents increased with density and rerritory size decreased with density. Floaters intruded into territories in a school to feed on algae, which induced overt aggression of the resident and reduced the productivity of algae growing there. Both the intruding frequency of floaters over territorial areas and their feeding pressure on algae increased at higher floater density. Floaters functioned to shift cost-benefit relationships for various territory sizes. They acted as food competitors to restrict territory size below a maximum through competitive interference. Although the growth rate of residents was inversely related to fish density, residents grew faster than floaters in each group. Under a given set of competitor abundances, economic defensibility determined territory size.     

Habitat context influences predator interference interactions and the strength of resource partitioning

Despite increasing evidence that habitat structure can shape predator–prey interactions, few studies have examined the impact of habitat context on interactions among multiple predators and the consequences for combined foraging rates. We investigated the individual and combined effects of stone crabs (Menippe mercenaria) and knobbed whelks (Busycon carica) when foraging on two common bivalves, the hard clam (Mercenaria mercenaria) and the ribbed mussel (Geukensia demissa) in oyster reef and sand flat habitats. Because these species co-occur across these and other estuarine habitats of varying physical complexity, this system is ideal for examining how habitat context influences foraging rates and the generality of predator interactions. Consistent with results from previous studies, consumption rates of each predator in isolation from the other were higher in the sand flat than in the more structurally complex oyster reef habitat. However, consumption by the two predators when combined surprisingly did not differ between the two habitats. This counterintuitive result probably stems from the influence of habitat structure on predator–predator interactions. In the sand-flat habitat, whelks significantly reduced their consumption of their less preferred prey when crabs were present. However, the structurally more complex oyster reef habitat appeared to reduce interference interactions among predators, such that consumption rates when the predators co-occurred did not differ from predation rates when alone. In addition, both habitat context and predator–predator interactions increased resource partitioning by strengthening predator dietary selectivity. Thus, an understanding of how habitat characteristics such as physical complexity influence interactions among predators may be critical to predicting the effects of modifying predator populations on their shared prey.     

Reef fish assemblages in north-western Sri Lanka: distribution patterns and influences of fishing practises

The distribution and abundance of reef fishes in relation to habitat structure were studied within Bar Reef Marine Sanctuary (BRMS) and on an adjacent reef, disturbed by destructive fishing techniques, in north-western Sri Lanka, by visually censusing 135 species groups using fifty metre belt-transects. Two types of continental shelf patch-reefs are found in the study area: coral reefs and sandstone reefs, which are divided into distinct habitats, four for the coral reef (shallow reef flat, shallow patch reef, deep reef flat and Porites domes) and two for the sandstone reef (structured sandstone-reef and flat sandstone-reef). Fish assemblages varied in structure between reef types and among habitats within reef types. Functional aspects of habitat structure and composition, such as available food and shelter, seemed to be important factors influencing distribution patterns. The strongest separation in the organisation of fish assemblages in BRMS was between reef types: 19% of all species were confined to the coral-reef patches while 22% were restricted to the sandstone reef patches and 59% were represented on both reef types. In terms of distribution among habitats, 21% of all species were restricted to one habitat while only 1.5% were present in all. The highest density of fish was in the coral reef habitats while highest species diversity was found in the most structurally complex habitat: the structured sandstone-reef. This habitat also had the highest proportion of species with restricted distribution. Planktivores were the most abundant trophic group in BRMS, and the species composition of the group varied among habitats. The comparison of the disturbed reef with BRMS suggested that habitat alteration caused by destructive fishing methods has strongly influenced the fish community. Within the fished area the structure of the fish assemblages was more heterogeneous, fish abundance was lower by an order of magnitude and species numbers were lower than in BRMS.     

An experimental study of the multiple effects of brown trout Salmo trutta on the bioenergetics of two Arctic charr Salvelinus alpinus morphs

This study investigated the importance of competition with brown trout Salmo trutta as a driver of the morphological and behavioural divergence of two morphs of Arctic charr Salvelinus alpinus. The morphs originated from two lakes differing in absence or presence of the competitor. The bioenergetics and behaviour of S. alpinus were quantified in replicate experimental enclosures (mean volume: 150 m(3) ) stocked with 15 S. alpinus of one morph or the other and in the absence or presence of nine S. trutta. The presence of S. trutta decreased growth rate, affected food consumption and increased activity costs in S. alpinus, but provided little support for the hypothesis that competition with S. trutta is a major driver of the divergence of the two S. alpinus morphs. Both morphs responded similarly in terms of mean growth and consumption rates per enclosure, but the association between individual morphology and growth rate reversed between allopatric and sympatric enclosures. While the activity patterns of the two morphs were unaffected by the presence of S. trutta, their swimming speed and activity rate differed. Since the profound differences in the structure of the physical habitat of the source lakes provided a more likely explanation for the difference observed among these two morphs than interspecific competition, it is hypothesized that physical habitat may sometimes be a significant driving force of the phenotypic divergence.     

The impact of an introduced predator on a threatened galaxiid fish is reduced by the availability of complex habitats

1. The availability of complex habitats such as macrophytes may be vital in determining the outcomes of interactions between introduced predators and native prey. Introduced brown trout (Salmo trutta) have impacted numerous small native freshwater fishes in the southern hemisphere, but the potential role of complex habitats in determining the direct outcomes of brown trout – native fish interactions has not been experimentally evaluated. 2. An in‐lake enclosure experiment was used to evaluate the importance of structurally complex habitats in affecting the direct impacts of brown trout on a threatened galaxiid fish. Five Galaxias auratus and a single brown trout were added to enclosures containing one of three different habitat types (artificial macrophytes, rocks and bare silt substrate). The experiment also had control enclosures without brown trout. Habitat‐dependence of predation risk was assessed by analysis of G. auratus losses to predation, and stomach contents of remaining fish were analysed to determine if brown trout directly affect the feeding of G. auratus and whether this is also habitat‐dependent. 3. Predation risk of G. auratus differed significantly between habitat types, with the highest mortality in enclosures with only bare silt substrate and the lowest in enclosures containing artificial macrophytes. This result highlights the importance of availability of complex habitats for trout – native fish interactions and suggests that increasing habitat degradation and loss in fresh waters may exacerbate the direct impacts of introduced predators. 4. Stomach contents analyses were restricted to fish in enclosures with artificial macrophytes and rocks, as most fish were consumed in enclosures with brown trout and only bare silt substrate. These analyses suggest that brown trout do not directly affect the feeding of G. auratus in complex habitats, but it is still unknown whether its feeding is reduced if complex habitats are unavailable.     

Intraspecific and interspecific competition induces density‐dependent habitat niche shifts in an endangered steppe bird

Interspecific competition is a dominant force in animal communities that induces niche shifts in ecological and evolutionary time. If competition occurs, niche expansion can be expected when the competitor disappears because resources previously inaccessible due to competitive constraints can then be exploited (i.e., ecological release). Here, we aimed to determine the potential effects of interspecific competition between the little bustard (Tetrax tetrax) and the great bustard (Otis tarda) using a multidimensional niche approach with habitat distribution data. We explored whether the degree of niche overlap between the species was a density‐dependent function of interspecific competition. We then looked for evidences of ecological release by comparing measures of niche breadth and position of the little bustard between allopatric and sympatric situations. Furthermore, we evaluated whether niche shifts could depend not only on the presence of great bustard but also on the density of little and great bustards. The habitat niches of these bustard species partially overlapped when co‐occurring, but we found no relationship between degree of overlap and great bustard density. In the presence of the competitor, little bustard's niche was displaced toward increased use of the species' primary habitat. Little bustard's niche breadth decreased proportionally with great bustard density in sympatric sites, in consistence with theory. Overall, our results suggest that density‐dependent variation in little bustard's niche is the outcome of interspecific competition with the great bustard. The use of computational tools like kernel density estimators to obtain multidimensional niches should bring novel insights on how species' ecological niches behave under the effects of interspecific competition in ecological communities.     

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.