Size-dependent foraging capacities and intercohort competition in an ontogenetic omnivore (Arctic char)

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 (G_MRD) 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 G_mRD 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.     

Foraging capacities and effects of competitive release on ontogenetic diet shift in bream, Abramis brama

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.     

Strategies of zooplanktivory shape the dynamics and diversity of littoral plankton communities: a mesocosm approach

Planktivorous fish can exert strong top‐down control on zooplankton communities. By incorporating different feeding strategies, from selective particulate feeding to cruising filter feeding, fish species target distinct prey. In this study, we investigated the effects of two species with different feeding strategies, the three‐spined stickleback (Gasterosteus aculeatus (L.)) and roach (Rutilus rutilus (L.)), on a low‐diversity brackish water zooplankton community using a 16‐day mesocosm experiment. The experiment was conducted on a small‐bodied spring zooplankton community in high‐nutrient conditions, as well as a large‐bodied summer community in low‐nutrient conditions. Effects were highly dependent on the initial zooplankton community structure and hence seasonal variation. In a small‐bodied community with high predation pressure and no dispersal or migration, the selective particulate‐feeding stickleback depleted the zooplankton community and decreased its diversity more radically than the cruising filter‐feeding roach. Cladocerans rather than copepods were efficiently removed by predation, and their removal caused altered patterns in rotifer abundance. In a large‐bodied summer community with initial high taxonomic and functional diversity, predation pressure was lower and resource availability was high for omnivorous crustaceans preying on other zooplankton. In this community, predation maintained diversity, regardless of predator species. During both experimental periods, predation influenced the competitive relationship between the dominant calanoid copepods, and altered species composition and size structure of the zooplankton community. Changes also occurred to an extent at the level of nontarget prey, such as microzooplankton and rotifers, emphasizing the importance of subtle predation effects. We discuss our results in the context of the adaptive foraging mechanism and relate them to the natural littoral community.     

Coexistence of Fish Species in a Large Lowland River: Food Niche Partitioning between Small-Sized Percids,Cyprinids and Sticklebacks in Submersed Macrophytes

In the spring and summer of each year, large patches of submersed aquatic macrophytes overgrow the bottom of the alluvial Warta River downstream of a large dam reservoir owing to water management practices. Environmental variables, macroinvertebrates (zoobenthos and epiphytic fauna, zooplankton) and fish abundance and biomass were assessed at this biologically productive habitat to learn intraseasonal dynamics of food types, and their occurrence in the gut contents of small-sized roach, dace, perch, ruffe and three-spined stickleback. Gut fullness coefficient, niche breadth and niche overlap indicated how the fishes coexist in the macrophytes. Chironomidae dominated in the diet of the percids. However, ruffe consumed mostly benthic chironomids, while perch epiphytic chironomids and zooplankton. The diet of dace resembled that in fast flowing water although this rheophilic species occurred at unusual density there. The generalist roach displayed the lowest gut fullness coefficient values and widest niche breadth; consequently, intraspecific rather than interspecific competition decided the fate of roach. Three-spined stickleback differed from the other fishes by consuming epiphytic simuliids and fish eggs. The diet overlap between fishes reaching higher gut fullness coefficient values was rather low when the food associated with the submersed aquatic macrophytes was most abundant; this is congruent with the niche overlap hypothesis that maximal tolerable niche overlap can be higher in less intensely competitive conditions.     

Size and temperature dependent foraging capacities and metabolism: consequences for winter starvation mortality in fish

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.     

Impacts of a submerged plant (Elodea canadensis) on interactions between roach (Rutilus rutilus) and its invertebrate prey communities in a lake littoral zone

1. Using 5‐m² field enclosures, we examined the effects of Elodea canadensis on zooplankton communities and on the trophic cascade caused by 4–5 year old (approximately 16 cm) roach. We also tested the hypothesis that roach in Elodea beds use variable food resources as their diet, mainly benthic and epiphytic macroinvertebrates, and feed less efficiently on zooplankton. Switching of the prey preference stabilises the zooplankton community and, in turn, also the fluctuation of algal biomass. The factorial design of the experiment included three levels of Elodea (no‐, sparse‐ and dense‐Elodea) and two levels of fish (present and absent). 2. During the 4‐week experiment, the total biomass of euplanktonic zooplankton, especially that of the dominant cladoceran Daphnia longispina, decreased with increase in Elodea density. The Daphnia biomass was also reduced by roach in all the Elodea treatments. Thus, Elodea provided neither a favourable habitat nor a good refuge for Daphnia against predation by roach. 3. The electivity of roach for cladocerans was high in all the Elodea treatments. Roach were able to prey on cladocerans in Elodea beds, even when the abundance and size of these prey animals were low. In addition to cladocerans, the diet of roach consisted of macroinvertebrates and detrital/plant material. Although the biomass of macroinvertebrates increased during the experiment in all Elodea treatments, they were relatively unimportant in roach diets regardless of the density of Elodea beds. 4. Euplanktonic zooplankton species other than Daphnia were not affected by Elodea or fish and the treatments had no effects on the total clearance rate of euplanktonic zooplankton. However, the chlorophyll a concentration increased with fish in all the Elodea treatments, suggesting that fish enhanced algal growth through regeneration of nutrients. Thus, our results did not unequivocally show that Elodea hampered the trophic cascade of fish via lowered predation on grazing zooplankton. 5. In treatments with dense Elodea beds (750 g FW m^?2), chlorophyll a concentration was always low suggesting that phytoplankton production was controlled by Elodea. Apparently, the top‐down control of phytoplankton biomass by zooplankton was facilitated by the macrophytes and operated simultaneously with control of phytoplankton production by Elodea.     

The diet and growth of roach (Rutilus rutilus (L.)) in Lake Vesijärvi and possible changes in the course of biomanipulation

Previous studies have suggested that the roach Rutilus rutilus (L.) stock of Lake Vesijärvi is one of the main factors delaying the recovery of the lake after sewage diversion. This study is concerned with the documentation of the diet of roach in the lake. In total, 531 roach were examined. Both in the pelagial and in the littoral the roach had mixed diets in May and in September—October. The importance of zooplankton decreased and the importance of benthos and plants increased with increasing size of roach. In July, in the pelagic zone all sizes of roach fed exclusively on zooplankton (Bosmina spp.), while in the littoral zooplankton had the highest volume proportions only in the smallest (<130 mm) roach. The frequent use of plant food and slow growth rate of large roach indicate a low availability of animal prey. As the fish densities decrease due to the mass removal taking place in the lake, the percentage of plant food in the diets of roach will probably decrease and the growth of roach will increase. Additionally, the tendency of the roach to migrate into the pelagic zone in early summer may be reduced, which would decrease their predation on the zooplankton.     

Ecological consequences of food partitioning for the fish population structure in a eutrophic lake

In the highly eutrophic lake, Frederiksborg Slotssø, the diet composition of the bream (Abramis brama L.) and roach (Rutilus rutilus L.) populations was examined during three periods with different food availability. The length range of bream and roach was 9–34 cm (TL) and 5–18 cm (TL), respectively. The relative food composition was examined for 2 cm and 1 cm length intervals of bream and roach, respectively. During all three periods, bream shifted from benthic cladocerans (Alona sp.) to zooplankton and chironomids within a transitional length of 15.0–20.0 cm. These foodshifts were coupled with a change in feeding behaviour from particulate to filter feeding. The biomass of chironomids was too low to sustain the consumption of larger bream (>20.0 cm) which initiated feeding in the pelagic zone even in periods when the mean length and biomass of the preferred zooplankton, Daphnia cucullata, were low. In contrast to bream, roach fed mainly on zooplankton. With increasing size, roach progressively shifted to larger zooplankton species due to the increasing mesh size of their branchial system. The importance of benthic animals in the diet of roach was minor due to low feeding efficiency on prey buried in the sediment. Detritus appeared in the diet of bream and roach in periods of low availability of animal food items. Feeding on detritus may provide an energetic advantage to bream and roach and increase the carrying capacity for these species in lakes, where detritus is highly abundant. Especially for the larger fish due to the decrease in their relative metabolic demands. However, the ability of bream to filter feed and with increasing size to retain food items smaller than those retained by roach may be the main mechanism for the dominance of bream over roach in highly eutrophic lakes.     

Trophic dynamics in a whole lake experiment: size-structured interactions and recruitment variation

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.     

Benthic resources are the key to Daphnia middendorffiana survival in a high arctic pond

1. Shallow arctic lakes and ponds have simple and short food webs, but large uncertainties remain about benthic–pelagic links in these systems. We tested whether organic matter of benthic origin supports zooplankton biomass in a pond in NE Greenland, using stable isotope analysis of carbon and nitrogen in the pond itself and in a ¹³C‐enrichment enclosure experiment. In the latter, we manipulated the carbon isotope signature of benthic algae to enhance its isotopic discrimination from other potential food sources for zooplankton. 2. The cladoceran Daphnia middendorffiana responded to the ¹³C‐enrichment of benthic mats with progressively increasing δ¹³C values, suggesting benthic feeding. Stable isotope analysis also pointed towards a negligible contribution of terrestrial carbon to the diet of D. middendorffiana. This agreed with the apparent dominance of autochthonous dissolved organic matter in the pond revealed by analysis of coloured dissolved organic matter. 3. Daily net production by phytoplankton in the pond (18 mg C m^?2 day^?1) could satisfy only up to half of the calculated minimum energy requirements of D. middendorffiana (35 mg C m^?2 day^?1), whereas benthic primary production alone (145 mg C m^?2 day^?1) was more than sufficient. 4. Our findings highlight benthic primary production as a major dietary source for D. middendorffiana in this system and suggest that benthic organic matter may play a key role in sustaining pelagic secondary production in such nutrient‐limited high arctic ponds.     

Diet composition and prey preference of tench, Tinca tinca (L.), common bream, Abramis brama (L.), perch, Perca fluviatilis L. and roach, Rutilus rutilus (L.), in two contrasting gravel pit lakes: potential trophic overlap with wildfowl

The abundance of benthic macroinvertebrate taxa and the relative abundance of zooplankton taxa were compared with the diet of bream, tench. perch and roach from two gravel pit lakes during May-July 1986 and 1987. Significant food preferences were demonstrated between species and between lakes. Chironomid pupae dominated the macroinvertebrate diet of perch, roach, ate predominantly either Spirogyra sp. (St Peters Lake) or Daphniu hyalina (Main Lake).
The Main Lake, bream fed largely upon chironomid larvae and the bivalve Sphuerium but in St Peters Lake they positively selected a variety of less abundant benthic invertebrate taxa. Bream switched from benthos to zooplankton in the Main Lake in 1986. Tench ate large numbers of Aselhs and showed positive selection of various macroinvertebrate prey in St Peters but ate D. hjulim in Main Lake. Tropic overlap for chironomids and other macroinvertebrate prey was demonstrated between perch, bream and tench, and potentially with wildfowl which used the gravel pits for breeding and wintering.     

The role of phosphorus release by roach [Rutilus rutilus (L.)] in the water quality changes of a biomanipulated lake

SUMMARY 1. Following fish removal, the water quality in biomanipulated lakes often improves concomitant with decreased phosphorus (P) levels. Because the decrease in P concentrations derives most probably either directly or indirectly from fish, which are the main target of biomanipulation, this study examined the P release of 0+, 1+ and 2+ roach [Rutilus rutilus (L.)] and changes in the P release during summer in a shallow eutrophic lake in Finland. 2. The P release was separated into P derived from benthic and littoral food items and into recycled P derived from feeding on zooplankton, to estimate the contribution of net P additions to the water column by the fish to the increase in P concentrations of the lake water (75–110 mg P m^?3) in summer 1991–96. 3. Individual P release of roach by both egestion and excretion was estimated with a bioenergetics model. The size of the roach population was estimated with a depletion method and the proportions of different age groups from catch samples, using a programme separating mixtures of normal distributions. The sensitivity of the release estimates to variation in the growth data was estimated with the jackknife technique. 4. The biomass‐specific P release by 0+ roach (0.36–0.54 mg P g^?1 day^?1) was higher than that by older roach (0.07–0.16 mg P g^?1 day^?1) throughout the summer. The P release by the whole roach population deriving from benthic and littoral food items (0.7–2.7 mg m^?3 during July to August, representing a net addition to the water column) was 5–19 times lower in 1991–96 than the recycled P release deriving from zooplankton (8.9–25.7 mg m^?3), and too low to explain the increase in the P concentration of the lake water during the summer. Because the biomass‐specific P release and roach diet composition vary with fish age, it is important to consider the age structure of fish populations to obtain correct estimates of P release and net additions to the water column. 5. The removal of roach by fishing diminished the roach stock greatly, but the fish‐mediated P release to the water column changed little. This effect was because of the high compensation capacity of the roach population, leading to high recruitment of young fish with higher biomass‐specific P release rates. 6. External loading is very low during summer months and therefore it cannot explain the increase in the P concentration of water during that time. Internal loading from the sediment might be as high as 10.2 mg P m^?2 day^?1, i.e. 50 times higher than the maximum net P addition by the total roach population.     

Seasonal changes in the diets and relative abundances of perch and roach in the littoral and pelagic zones of a large lake

In the Enonselkä and Laitialanselkä basins of Lake Vesijärvi, perch Perca fluviatilis and roach Rutilus rutilus were abundant in the littoral and in the pelagic zones throughout the summer. In the littoral zone, roach was always more numerous than perch, while perch dominated in the open water. Intraspecific diet overlap values were higher than interspecific values. In the pelagic zone, perch <155 mm fed mainly on the cladoceran Leptodora kindtii , while small bosminids were most important food items for roach. Large perch were piscivorous, feeding mainly on smelt Osmerus eperlanus . In the littoral zone small perch foraged on zooplankton and chironomid larvae and large perch on chironomids and fish (small perch). Small roach fed mainly on bosminids and detritus, while for roach <185 mm macrophytes ( Elodea Canadensis, Lemna trisulca ) were also of importance. Detritus was more common in the food of roach in Laitialanselkä than in Enonselkä. The slower growth rate of roach in Laitialanselkä compared with Enonselkä was probably connected with this. However, considering the latitude of the lake, the growth rate of both roach and perch was relatively fast in both basins. The results indicated that in a large lake both perch and roach are able to utilize effectively the different habitats and diverse food resources. By segregation in food resource utilization they are able to co-exist in large quantities, at the same time maintaining a relatively fast growth rate.     

Effetcs of ontogenetic scaling on resource exploitation and cohort size distributions

Variation in growth rates among individuals leading to the formation of broad size distributions is commonly observed in animal cohorts. Here we use laboratory derived size–scaling relationships to identify mechanisms driving changes in size distribution patterns within cohorts during early ontogeny. We introduced young‐of‐the‐year perch Perca fluviatilis cohorts with different variation in body size distributions in pond enclosures. We kept the exploitative competitive environment constant by adjusting the number of introduced fish such that metabolic requirements were constant between different treatments. Based on modelling results we theoretically derived relative growth rates of differently sized fish when only taken exploitative competitive interactions into account. In agreement with predictions we found that initial variation in body size was negatively correlated with subsequent changes in body size variation in the pond experiment. Corresponding results were obtained in a field study covering 13 studied young‐of‐the‐year perch cohorts in a small lake. Besides having a lower maximum growth capacity, initially large fish also suffered more from resource limitation in our experiment. The results suggest that exploitation competition is a major factor behind growth patterns in young fish cohorts, generally leading to size convergence. To explain the commonly observed pattern of size divergence in animal cohorts, including fish, we suggest that differential timing of diet shifts or mechanisms not related to exploitative interactions must be taken into account. For diet shifts to lead to size divergence we suggest that individuals with an initial size advantage need access to an exclusive prey which has a high growth potential. This, in turn, allows initially larger individuals to surf on a wave of growing prey while individuals only capable to feed on a depressed initial resource experience low growth rates.     

Resource limitation during early ontogeny: constraints induced by growth capacity in larval and juvenile fish

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     

Diet-dependent body morphology and ontogenetic reaction norms in Eurasian perch

Individual morphology and performance are directly or indirectly under the influence of variation in resource levels. To study the effects of different resource conditions and their effects on morphology and ontogenetic reaction norms in young‐of‐the‐year (YOY) perch (Perca fluviatilis), we used three different approaches. First, we examined the morphological trajectories over early ontogeny in relation to lake‐specific resources in a field study. Second, one lake that lacked perch recruitment was stocked with perch eggs from a control lake in a whole‐lake experiment to study ontogenetic reaction norms. Third, we compared the development of YOY perch in the three lakes that mainly inhabited the littoral zone with YOY perch experimentally confined to enclosures in the pelagic zone of the lakes.
Overall body morphology of the YOY perch changed both as a function of size and as a function of diet. As perch increased in size they developed a deeper body morphology corresponding to an increased proportion of benthic macroinvertebrates in their diet. In pelagic enclosures where perch were constrained to feed mainly on zooplankton they had a more fusiform body morphology than perch in the lakes that fed on a mixture of zooplankton and macroinvertebrates. Similarly, the ontogenetic reaction norm of perch was related to the diet and lake‐specific zooplankton levels in the whole‐lake experiment.
In the pelagic enclosures, perch with high growth rates had a more fusiform body morphology than slow‐growing perch, whereas the opposite was found in the lakes, where perch included more macroinvertebrates in their diets. Perch in lakes with a higher proportion of macroinvertebrates in the diet also had deeper body morphology. The opposite morphology – growth rate relationship found between perch in the pelagic versus those using the whole lake suggest a morphological trade‐off between foraging on zooplankton and foraging on macroinvertebrates. Our results suggest that YOY perch show different ontogenetic reaction norms as a function of lake‐specific resource levels, which may allow YOY recruitment to later stages. Our results further suggest that diet‐related changes in morphology are a rapid process.     

Dung exploitation by the dung beetle Typhaeus typhoeus (Col., Geotrupidae)

Summary The competitive interactions between roach (Rutilus rutilus) and rudd (Scardinius erythrophthalmus) were investigated in two habitats, the open water and the waterlily zone. The growth rates of both species in enclosures were lower when confined together than when alone, demonstrating interspecific competition. Allopatric roach had the highest growth rate in both habitats although the open water was preferred. The diets of both species were dominated by zooplankton in both the open water and waterlily zones. In laboratory experiments, roach had significantly higher feeding rates than rudd when fed D. magna and Cyclops sp. The impact of roach on the mean sizes and densities of zooplankton in the enclosures, together with the results from the laboratory study, indicate that roach were competitively superior in the open water. Because of high mortality in the sympatric waterlily enclosure, no conclusions about interspecific competition in this habitat could be drawn. The observed habitat segregation between roach and rudd was at least partly interactive.     

The relationship between ontogeny, morphology, and diet in the Chinese hook snout carp (Opsariichthys bidens)

The relationship of changes of body and mouth shape with diet during ontogeny in the Chinese hook snout carp Opsariichthys bidens was examined. Shape changes were analyzed using geometric morphometrics. Body shape changed from a shallow body to a deep body as size increased. Similarly, the head of O. bidens changed from an upward-pointing direction to a downward-pointing directing as size increased. The position of the eye also changed during ontogeny, with large individuals having an eye situated higher on the head than small individuals. The conspicuous symphyseal knob on the lower jaw of O. bidens results in a typical zigzag pattern of the jaw. The zigzag pattern became more pronounced as fish grew larger. No difference in body shape or jaw shape was found between males and females. The shape changes were associated with changes in diet, suggesting that diet choice over the course of ontogeny has a strong impact on body shape. Benthic macroinvertebrates and zooplankton dominated the diet of small individuals, medium-size individuals specialized in benthic macroinvertebrates, and fish dominated the diet of large individuals.     

Temporal variation in feeding morphology and size-structured population dynamics in fishes

Considerable variation in morphology associated with resource use is a classic example of local adaptation to the environment. We demonstrate that a temporal change in feeding morphology might occur within a population. In a 5-year natural field experiment, we estimated gill raker morphology, resource density and population dynamics of the roach and its competitor, the perch. Despite a variation in density of zooplankton resources and perch across years, no change in roach density was observed. However, gill raker morphology in roach covaried with size structure of the zooplankton resource across years. A laboratory experiment confirmed that gill raker morphology has a great effect on roach foraging efficiency on zooplankton and that there is a functional trade-off with regard to zooplankton foraging. We suggest that the response in gill raker structure is an adaptation to deal with the rapid population dynamics of zooplankton, which are in turn mediated by changes in the size structure of the competing perch.