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     

Feeding of fish in Lake Glubokoe

Lake Glubokoe is inhabited by 13 species of fish. The commonest are roach, bream, perch and ruff. Comparison of seine catches for 1950 and 1984 revealed a considerable decrease in the share of perch (from 31% to 2%) and ruff (from 18.5 to less than 0.1%) in 1984. The richest food resource in the lake is crustacean plankton. In summer, it is mostly consumed by roach. Estimation of the electivity shows that, unlike perch, roach selectively consumes small crustaceans (Bosmina and Polyphemus). Poor growth rate of all common species of fish points to their food limitation. In summer the diets of common species of fish are differentiated to a considerable extent. Of 36 pairs of combinations a high food overlap was found only in 2 cases: 1) O + perch and Leucaspius delineatus, 2) yearlings and adult roach. Low diet overlap in fish during intensive feeding in summer probably reflects a feedback between the volume and intensity of food competition. The populations of bream and ruff probably compete for chironomid larvae which inhabit the sublittoral, as perch, roach and Leucaspius delineatus probably do for large littoral insects. However, intraspecific competition appears to be more important than interspecific for the roach and bream populations (the two most numerous species in the lake).     

Diel vertical migrations of bathypelagic perch fry

The behaviour of young‐of‐the‐year (YOY) perch Perca fluviatilis as a dominant species in the assemblage of fry in the pelagic of Slapy Reservoir (Czech Republic), was studied during late May and mid‐June 2002 using acoustic methods and complementary net catches. During the day, perch fry were present simultaneously in littoral, epipelagic and bathypelagic habitats. Bathypelagic perch fry, forming a scattering layer, migrated vertically each day between the epilimnion and hypolimnion, with an amplitude of 11·0 m in May and 12·5 m in June. At dusk, the migratory bathypelagic fry mixed in the epilimnion with non‐migrating epipelagic fry and spent the night close to the thermocline (abundance maximum at 3–4 m in May, 0–2 m in June). In June, shoaling behaviour by some of the bathypelagic perch fry was also observed: the shoaling fry remained higher in the water column than the non‐shoaling fry. Both depths of the scattering layer and the depths of the fry shoals were strongly controlled by the light intensity. The contribution of the bathypelagic part of the population to the total numbers of pelagic perch fry decreased from 28·1% in May to 4·7% in June, while the density of all pelagic perch fry increased (c. 96 000 individuals ha^?1 in May and 142 000 individuals ha^?1 in June). In May, the bathypelagic (average total length, L_T, 11·9 mm) and epipelagic (average L_T 14·6 mm) perch fry differed in size while, in June, the epipelagic fry were divided into two distinct size groups. The more abundant group, of small epipelagic perch fry (average L_T 14·6 mm), was similar in size to the bathypelagic fry (average L_T 14·6 mm) while the less abundant group, of larger epipelagic fry (average L_T 34·4 mm), was similar in size to littoral perch fry (average L_T 35·0 mm). The results suggest that in perch fry three different survival strategies with different risks can be used in the same locality, time and year.     

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.     

Food web manipulation in a small,eutrophic Lake Wirbel,Poland: effect of stocking with juvenile pike on planktivorous fish

In a four year experiment (1988–1991) carried out in a shallow, eutrophic lake, an increase in young-of-the-year pike (Esox lucius) density was used to control planktivorous fish assemblages consisting mainly of small-sized cyprinids: roach (Rutilus rutilus), Leucaspius delineatus, and white bream (Blicca bjoerkna). Stocking with pike fry of ca 30 mm in three successive springs resulted in large-scale mortality among prey of vulnerable sizes. Rotenone treatment at the end of the fourth year showed that roach and white bream populations contained few 0+ to 2+ fish, but very high numbers of 3+ to 6+ fish. By this time, short-lived, slow-growing L. delineatus, with no size refuge from predators, had been driven almost to extinction.     

Food and habitat partitioning among juveniles of three fish species in the pelagial of a mesotrophic lake

An analysis of the diet of 0⁺ perch, smelt and roach caught at night of the pelagial of a mesotrophic lake showed that their food was composed mainly of herbivorous and predatory cladocerans, copepods and Chaoborus larvae during summer, and of herbivorous cladocerans and copepods during October. An analysis of habitat use by juveniles revealed separation among the species: roach occurred in the upper, perch and smelt in the deeper water layers. Food and habitat were alternatively partitioned among the juveniles. In early summer the lowest food overlap between perch and smelt coincided with high habitat overlap. In October high food overlap between perch and smelt corresponded with low habitat overlap. Smelt and roach fry used common food resources throughout the season, but they were segregated in habitat.     

Prey selection by the larvae of three species of littoral fishes on natural zooplankton assemblages

The prey selection of larvae of three common littoral fish species (pike, Esox lucius; roach,Rutilus rutilus; and three-spined stickleback, Gasterosteus aculeatus) was studied experimentally in the laboratory by using natural zooplankton assemblages. Zooplankton prey was offered at four different concentrations to study the functional responses of the planktivores. The diets of pike and sticklebacks were formed mainly of copepod juveniles and adults, which dominated the prey communities, although sticklebacks ate also cladocerans. The diet of roach larvae consisted of rotifers, cladocerans and copepods, without prey selection, in equal proportions indicating a more omnivorous diet. All fish larvae were able to feed selectively although in sticklebacks prey selection was less pronounced. Pike and roach larvae preferred large prey to smaller prey types. Patterns of prey selection are discussed in the context of size-selection theory and apparent vs. true selection.     

Habitat shift in roach (Rutilus rutilus) induced by pikeperch (Stizostedion lucioperca) introduction: predation risk versus pelagic behaviour

Changes in the fish community structure and habitat use were followed after the introduction of pikeperch (Stizostedion lucioperca) to the roach-dominated Lake Gjersjøen. Quantitative echosounding showed that the density of juvenile roach (Rutilus rutilus) was dramatically reduced in pelagic areas, from 12 000–15 000 fish/ha to 250 fish/ha, while total fish density remained unchanged in littoral areas. At the same time, the habitat segregation between different size groups of roach was altered as larger roach utilized the pelagic zone after pikeperch introduction. The loss of the pelagic refuge for juvenile roach increased the availability of juvenile roach to littoral predators, notably perch. In littoral areas, the fish community changed from one dominated by roach (> 95%) to one dominated by perch (> 50%).     

The fish community structure in Tjeukemeer in relation to fishery and habitat utilization

The fish community in Tjeukemeer was monitored from 1971 to 1988 by trawling, and during the summer of 1988 the distribution of the more abundant species was determined in relation to size and location. Bream, pikeperch and smelt were the most dominant fishes, whereas roach, white bream, perch and ruffe comprised <10% of the total catches. After the termination in 1977 of the intensive gillnet fishery both pikeperch >50 cm and bream >30 cm increased in biomass, but roach >15 cm and perch >15 cm virtually disappeared and pikeperch <50 cm substantially decreased. Only smelt, ruffe, white bream and bream <30 cm hardly changed in biomass. Bream, pikeperch, perch and smelt were restricted to the open water zone, but roach <15 cm and 0 + bream were confined to the littoral zone. White bream and ruffe did not show a distinct habitat preference. Because the recruitment of smelt is largely dependent on immigration from the IJsselmeer, a feed-back between the smelt and pikeperch population is lacking. Because the carrying capacity of the pikeperch population is mainly determined by smelt, the other fish <15 cm are very vulnerable to predation when the smelt population is consumed, before a new year-class of smelt is recruited, or when the smelt fails to recruit. The vulnerability to predation of the different species and their feeding habits are discussed in relation to their distribution.     

Distribution and Diet of 0+ Fish within a Canyon‐Shaped European Reservoir in Late Summer

The distribution and diet of age 0+ fish were studied in the deep canyon‐shaped Římov Reservoir (Czech Republic), which is characterized by a longitudinal trophic gradient. During late summer of two years, 0+ fish were sampled from inshore and offshore habitats along the longitudinal reservoir axis. Offshore catches of 0+ fish from the surface layer were dominated by roach (Rutilus rutilus ), bream (Abramis brama ) and perch (Perca fluviatilis ), whereas in the deeper open water perch predominated. Inshore catches of 0+ fish were constituted mainly by perch and roach. The proportions of roach in the inshore catches were highest at the upper and most eutrophic part of the reservoir, whereas the proportions of perch in the inshore catches were higher at the downstream areas. Total catches of both inshore and offshore 0+ fish increased upstream in the reservoir. Offshore 0+ perch were of consistently smaller size than inshore 0+ perch. Inshore 0+ perch had significantly smaller size at the upstream reservoir part than at the downstream, more lacustrine regions. The diet of both inshore and offshore 0+ fish consisted predominantly of crustacean zooplankton. Perch diet was generally dominated both by cladocerans and copepods, whereas roach diet consisted chiefly of cladocerans. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Impact of flood on distribution of bathypelagic perch fry layer along the longitudinal profile of large canyon-shaped reservoir

Previous studies have shown that under normal spring conditions, the layer of bathypelagic perch Perca fluviatilis fry (BPF) is equally distributed along most of the longitudinal profile of reservoirs. In the first half of June 2004, local flooding entering the large canyon‐shaped Orlík Reservoir (Czech Republic) completely flushed out the existing fry community from the 31 km long riverine part of the reservoir. The pelagic zone in this reach was then recolonized by cyprinid (mainly roach Rutilus rutilus and bream Abramis brama) fry being of either littoral or riverine origin. Subsequently, the BPF layer was recorded only in the 22 km long lacustrine part of the reservoir. In the upper reach of this part where, due to the sudden increase in volume, the water current slowed down rapidly and fry originating from both riverine and central parts of the reservoir gathered in high numbers, two distinct BPF layers were observed. During mid‐day, the upper BPF layer, created predominantly by shoaling fishes (abundance >126 000 individuals ha^?1), occurred between 6 and 10 m. A second, lower BPF layer, created by non‐shoaling fishes exclusively (30 000 individuals ha^?1), was recorded between 12 and 17 m depth. Both upper and lower BPF layer were composed of perch (69·6 and 66·8% in abundance respectively) and zander Sander lucioperca (29·8 and 28·6% in abundance respectively). In the lower BPF layer, ruffe Gymnocephalus cernuus also contributed considerably to the fry assemblage (4·0% in abundance). Perch from the upper BPF layer (mean 25·1 mm total length, L_T) did not differ in size from perch from the lower BPF layer (mean 25·0 mm L_T). Similarly, zander from the upper BPF layer (mean 27·2 mm L_T) were almost the same size as those from the lower BPF layer (mean 26·9 mm L_T). Perch from both BPF layers, however, were noticeably smaller than zander. The results from acoustic survey and complementary net catches suggest that no epipelagic perch fry were found in the reservoir where thermal stratification had been destroyed by flooding and windy weather.     

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.     

Factors affecting the early life history of yellow perch,Perca flavescens

Synopsis From 1979 to 1981 we followed the movement, diet, and growth of yellow perch,Perca flavescens, for their first 70 days after hatching in Lake Itasca, Minnesota. Perch spawned inshore during early spring; hatching occurred 10–20 days after spawning. Newly hatched perch were 5.6–6.2 mm total length (TL). Soon after hatching the larvae moved into the limnetic zone where they began feeding. This movement is probably a mechanism to escape intense predation in the littoral zone. Normally the first food of perch was immature copepods, but within a week they incorporated all common zooplankters into their diet. When the perch reached 25 mm TL (about day 40) they returned to the littoral zone, where they ate larger and more abundant prey than was present in the limnetic habitat. There is no correlation between growth rates and zooplankton abundances, which suggests that food quantity is not a limiting factor in the early life history of perch in Lake Itasca.     

Competition,predation, cannibalism: the development of young‐of‐the‐year perch populations in ponds with bream or roach

Piscivory of perch can occur within a few weeks after perch hatch, leading to the development of two size‐cohorts, with the larger perch becoming cannibals. However, the possibility of early piscivory is assumed to depend on the hatch timing of the prey and predator. Two species, bream (2006) and roach (2007), were tested as the prey fish. The bream (hatching 12 days after perch) were preyed upon by the perch, leading to the predicted development of two sizes of cohorts as well as to cannibalism. With roach (hatching simultaneously with perch), however, no piscivory or cannibalism occurred and the perch population was unimodally distributed. The results of this experimental pond study underpin recent theoretical findings that size differences between predator and prey, determined through differences in the timing of hatching as well as differences in juvenile growth rates, foster the occurrence of early piscivory in YOY perch that may lead to bimodality and finally to intra‐cohort cannibalism.     

Feeding of benthophagous fish in the Chogray Reservoir

Data on feeding of the bream Abramis brama, roach Rutilus rutilus, and white bream Blicca bjoerkna in Chogray Reservoir in 2010 and 2012 are presented. The main diet of bream consists of Chironomidae, and the diet of roach and white bream consists of the bivalve mollusk Dreissena polymorpha.     

Habitat Use by YOY in the Littoral Zone of an Artificially Heated Reservoir

Both temporal and spatial distributions of a young of the year (YOY) fish community were studied in the littoral zone of a French reservoir. The time survey indicated that littoral occupation highly depends on YOY time of appearance. During late spring and summer, the smallest white bream YOY (L_T < 20 mm) were restricted to complex habitats whereas those of pumpkinseed occupied all available habitats. Growing white bream progressively migrated to open water areas. At that time, pumpkinseed was the only species sampled in abundance in the littoral zone and was still collected in all habitats. The success of the introduction of pumpkinseed might be explained by both its late hatching and its opportunistic strategy in habitat selection enabling it to exploit physical spaces left vacant. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

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.     

River habitat homogenisation enhances trophic competition and promotes individual specialisation among young of the year fish

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Feeding habits and ontogenic changes in digestive enzyme patterns in five freshwater teleosts

Feeding habits and the activity of digestive enzymes (total alkaline proteases, α‐amylase and lipase) from dace Leuciscus leuciscus, roach Rutilus rutilus, Prussian carp Carassius auratus gibelio, perch Perca fluviatilis and pikeperch Sander lucioperca fry were studied in the Malye Chany Lake–Kargat Estuary (western Siberia, Russia). The diet of fry from all studied species was mainly composed of chironomid larvae and zooplanktonic organisms (i.e. cladocera and copepoda), whereas carnivorous species such as P. fluviatilis and S. lucioperca also preyed on fry from other fishes while detritus and microalgae were also important in the diet of ommivorous species. When comparing diet similarity (Sørensen–Dice index, Q_S) among fry at different stages of development, both omnivorous and carnivorous species showed a high level of similarity (0·67 < Q_S < 0·89 and 0·73 < Q_S < 0·89, respectively). Diet similarity values were in agreement with the overall digestive activity profile analysed by cluster analysis. Diet similarity suggested potential trophic competition when zooplanktonic and benthic prey began to decline towards autumn. The analysis of pancreatic digestive enzymes revealed a correlation among their activities and fry feeding habits with α‐amylase:total proteases (A:P) values higher than 1 in omnivorous species and lower (A:P ≤ 1) in carnivorous species.