Shifts in fish communities along the productivity gradient of temperate lakes—patterns and the importance of size-structured interactions

Species biomass and size composition of fish faunas along a productivity gradient were studied in south Swedish lakes. Generally, with increasing productivity (measured as chlorophyll content), Salmoniformes were replaced by percids, which in turn were replaced by cyprinids, as suggested in previous studies. However, percids showed two peaks in biomass, one in medium productive lakes due to perch and one in highly productive lakes due to zander. Benthic piscivores were present in all lakes, whereas pelagic piscivores were absent in the least productive lakes. The proportion of piscivores in the total fish biomass showed a peak in medium productive lakes, largely reflecting the importance of piscivorous perch. The median size of the dominant cyprinids (roach) in the systems studied increased as the importance of piscivores increased, which was interpreted as a size refuge response to increased predation pressure. The same pattern was present for the dominant planktivore, vendace, in low productive systems. Although a predictable pattern of change in the fish fauna was found along the productivity gradient, other environmental factors such as structural complexity may be the ultimate cause of the observed succession pattern.     

Trophic structure, species richness and biodiversity in Danish lakes: changes along a phosphorus gradient

1. Using data from 71, mainly shallow (an average mean depth of 3 m), Danish lakes with contrasting total phosphorus concentrations (summer mean 0.02–1.0 mg P L?l), we describe how species richness, biodiversity and trophic structure change along a total phosphorus (TP) gradient divided into five TP classes (class 1–5: <0.05, 0.05–0.1, 0.1–0.2, 0.2–0.4,> 0.4 mg P L?1).
2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floating‐leaved macrophytes, species richness was unimodally related to TP, all peaking at 0.1–0.4 mg P L?1. The Shannon–Wiener and the Hurlbert probability of inter‐specific encounter (PIE) diversity indices showed significant unimodal relationships to TP for zooplankton, phytoplankton and fish. Mean depth also contributed positively to the relationship for rotifers, phytoplankton and fish.
3. At low nutrient concentrations, piscivorous fish (particularly perch, Perca fluviatilis) were abundant and the biomass ratio of piscivores to plankti‐benthivorous cyprinids was high and the density of cyprinids low. Concurrently, the zooplankton was dominated by large‐bodied forms and the biomass ratio of zooplankton to phytoplankton and the calculated grazing pressure on phytoplankton were high. Phytoplankton biomass was low and submerged macrophyte abundance high.
4. With increasing TP, a major shift occurred in trophic structure. Catches of cyprinids in multiple mesh size gill nets increased 10‐fold from class 1 to class 5 and the weight ratio of piscivores to planktivores decreased from 0.6 in class 1 to 0.10–0.15 in classes 3–5. In addition, the mean body weight of dominant cyprinids (roach, Rutilus rutilus, and bream, Abramis brama) decreased two–threefold. Simultaneously, small cladocerans gradually became more important, and among copepods, a shift occurred from calanoid to cyclopoids. Mean body weight of cladocerans decreased from 5.1 μg in class 1 to 1.5 μg in class 5, and the biomass ratio of zooplankton to phytoplankton from 0.46 in class 1 to 0.08–0.15 in classes 3–5. Conversely, phytoplankton biomass and chlorophyll a increased 15‐fold from class 1 to 5 and submerged macrophytes disappeared from most lakes.
5. The suggestion that fish have a significant structuring role in eutrophic lakes is supported by data from three lakes in which major changes in the abundance of planktivorous fish occurred following fish kill or fish manipulation. In these lakes, studied for 8 years, a reduction in planktivores resulted in a major increase in cladoceran mean size and in the biomass ratio of zooplankton to phytoplankton, while chlorophyll a declined substantially. In comparison, no significant changes were observed in 33 ‘control’ lakes studied during the same period.     

The distribution and growth of coarse fish in gravel-pit lakes in south-east England

SUMMARY. Age and growth rates of the coarse fish of thirty-nine gravel-pit lakes in south-east England are compared. Species composition was very variable, the commonest species being roach ( Rutilus rutilus ), bream ( Abramis brama ), tench ( Tinca tinca ), perch ( Perca fluviatilis ) and pike ( Esox lucius ). Two main kinds of fish community were identifiable from the survey: (1) those consisting of a mixture of cyprinids, usually roach and bream; and (2) those in which cyprinids were rare or absent and the fish community was composed of perch and pike. The majority of the lakes fell into the first category. Comparison of growth curves derived by back-calculation showed that, in all species, variation occurred from lake to lake and from year to year. The majority of gravel-pit lake fishes had 'medium' growth rates when compared with published data by other workers on different waters. Seasonal growth of roach infected with plerocercoids of Ligula intestinalis was observed at monthly intervals. Annulus formation on the scales of these fish occurred at the onset of growth in May. The heavy Ligula burden in 89% of these fish had not caused the formation of any additional or false rings.     

Development of fish communities in lakes after biomanipulation

Biomanipulation measures in the Netherlands are usually a combination of a drastic fish stock reduction and an introduction of pike fingerlings. In three small shallow lakes (Noorddiep, Bleiswijkse Zoom and Zwemlust) these measures resulted in a clear water state and the development of macrophytes. After the measures the fish community developed differently because of the new physical and biological conditions. Results of lake Noorddiep and lake Bleiswijkse Zoom showed that the fish community became more divers. Bream and carp became less dominant and were partly replaced by roach and perch. The importance of the main predator pike-perch was strongly reduced and replaced by pike and perch. The share of piscivorous fish in the total fish stock increased at all sites. The recruitment of young-of-the-year was similar or even higher in the clear overgrown areas than in the turbid water before the measures, but the recruitment of young-of-the-year to older fish differed between the species. Predation by pike and perch could not control the young-of-the-year cyprinids, but their predation may have contributed to the shift from bream to roach, because of selective predation on bream in the open water, while roach was hiding in the vegetation. The macrophytes provide new refugia and feeding conditions that favour roach and perch, but offer relatively poor survival conditions for bream and carp.     

The role of predation and competition in determining the distribution of common bream,roach and white bream in Dutch eutrophic lakes

Synopsis Most of the lakes in The Netherlands are turbid and without vegetation. This is regarded as the result of increasing eutrophication within the last decades. Under these conditions common bream, roach, and white bream are the most common cyprinids. In six shallow (1–3 m), wind exposed lakes the abundance of common bream and roach was linked to the abundance of pikeperch; bream dominated when pikeperch was abundant, but when the latter species was rare, roach was dominant and the biomass of bream was reduced. The biomass of white bream was always relatively low. In lake Tjeukemeer the distribution of roach < 20 cm fork length (FL) and bream was also related to the distribution of pikeperch. Only roach > 20 cm FL managed to coexist with pikeperch in the open water area, whereas roach < 20 cm was confined to the littoral zone where pikeperch was nearly absent. Bream occurred mainly in the open water and avoided the littoral zone where it competed with roach. White bream occupied an intermediate position, occurring in relatively low density both in the littoral zone and in open water. The importance of predation and competition in determining the distribution and abundance of roach and common bream in the eutrophic lakes of The Netherlands is discussed.     

The effects of food-web management on fish assemblage dynamics in a north temperate lake

There was an intensive trawl fishery in the Enonselkä basin in Lake Vesijärvi (southern Finland) in 1989–1993 and thereafter the fishing intensity declined. The fish assemblage dynamics were studied both during the years of intensive trawling and for 3 years afterwards. Roach Rutilus rutilus dominated the fish assemblage before the mass removal. The intensive trawling effectively diminished the roach stock and the present fishing intensity has been sufficient to prevent its recovery. There were significant decreasing trends in the gillnet cpue (catch per unit effort) of bream Abramis brama (L.) and bleak Alburnus alburnus (L.) while increasing trend was observed in cpue of ruffe Gymnocephalus cernuus (L.) and vendace Coregonus albula (L.). Perch Perca fluviatilis L. and pikeperch Stizostedion lucioperca (L.) showed variations in cpue but no trends existed. The proportion of cyprinids decreased while that of percids and coregonids increased in the catches. The changes in the fish assemblage were induced by the intensive fishing and by the concomitant decline in the ecosystem productivity.     

Restoration by biomanipulation of lake Bleiswijkse Zoom (The Netherlands): First results

In 1987, the Bleiswijkse Zoom, a small, shallow lake in The Netherlands, was divided into two compartments to investigate the possible use of biomanipulation as a tool for restoring the water quality of hypertrophic lakes. The density of the fish stock before restoration was about 650 kg.ha^–1, composed mainly of bream, white bream and carp. Pikeperch was the main fish predator in the lake. In April, 1987, in one compartment (Galgje) all planktivorous bream and white bream and about 85% of the benthivorous bream and carp were removed. Advanced pikeperch fry were introduced as predator during the transient period. The other compartment (Zeeltje) was used as a reference. Removal of the fish in Galgje resulted in low concentrations of chlorophyll-a, total phosphorus, nitrogen and suspended solids. The absence of bottom-stirring activity by benthivorous fish and the low chlorophyll-a concentrations led to an increase in the Secchi disk transparency from 20 to 110 cm. Within two months after removal of the fish, macrophytes, mainly Characeae, became abundant. Until July the high density of large zooplankton species caused low algal biomass. From June onwards, the zooplankton densities decreased, but the algal concentrations remained low. This is probably because of nutrient limitation or depression of algal growth by macrophytes or both. Compared with the non-treated compartment the number of fish species in the treated compartment was higher. Perch, rudd and roach, i.e. the species associated with aquatic vegetation, were found in the samples. The survival of the O⁺ pikeperch was poor. The pikeperch could not prevent the growth of young cyprinids. Within two months after the removal of the fish a habitat for northern pike was created.     

Variation in abundance and survival of fish larvae in shallow eutrophic lake Tjeukemeer

Synopsis Variations in abundance and survival of larvae of the major fish species, bream, roach, pikeperch, perch and smelt were studied in shallow eutrophic lake Tjeukemeer (1986–1988). Except for smelt, the size of the spawning stock and the abundance of larvae were not related. Both the timing of appearance of larvae and their growth rate were determined by water temperature as was shown by a growth model developed earlier for Tjeukemeer fishes. In cyprinids and percids the survival of the larvae was hypothesized to be negatively related to the duration of the earliest stages. Except in cyprinids, the abundance of 0+ juvenile fish could be explained by the number of larvae. In cyprinids 0+ juvenile abundance at the end of the year was not related to the number of larvae in May and June. The cyprinid juvenile survival rates are hypothesized to be related to the size of the main cyprinid predator, 0+ pikeperch.     

Growth and production of O+ bream (Abramis brama), O+ roach (Rutilus rutilus) and O+ carp (Cyprinus carpio) in 10 drainable 0.1 Ha ponds

In ten 0.1 ha drainable ponds O⁺ cyprinids were stocked to assess their impact on the aquatic community. Bream and roach were stocked in early June, 1986, and carp in early July, 1986. The ponds were drained mid November, 1986. The relationship between the fish community and its food resource was estimated in terms of production and consumption with use of the data on growth and mortality of the fish. The average gross production of the fish species in the ten ponds (48.4 kg during the experiment) was dominated by the carp (58.5%). A minimum estimate of the food consumption of the O⁺ cyprinids is 760 kJ.m^–2. The average production of bream and roach was much lower than that of carp due to a higher mortality and a more specialized feeding pattern. The gross productions of carp, bream and roach in the ten ponds were not significantly related, although stomach analysis showed diet overlap between the carp and the bream.     

Correlations between type‐indicator fish species and lake productivity in German lowland lakes

Morphotypes for 67 lakes in the German lowlands were derived, based on maximum depth and mixis type. A threshold of 11 m maximum depth was identified to be the best level to discriminate shallow from deep lake morphotypes. The fish communities in these two morphotypes were significantly different. Indicator species analyses based on fish biomasses found vendace Coregonus albula in deep lakes and ruffe Gymnocephalus cernuus , bream Abramis brama , white bream Abramis bjoerkna , roach Rutilus rutilus , pikeperch Sander lucioperca and small perch Perca fluviatilis in shallow lakes to be the most representative species of their communities. Lake productivity was closely related to biomass and in part abundance of the type‐indicator species, with vendace declining with increasing chlorophyll a concentration in the deep lakes, whereas biomass of pikeperch, bream, white bream and ruffe increased and biomass of small perch decreased with increasing chlorophyll a . These results indicate that assessment of ecological integrity of lakes by their fish fauna is generally possible, if lakes are initially separated according to a depth‐related morphotype before the assessment, and if eutrophication is considered to be the main anthropogenic degradation.     

Fish community structure in lowland drainage channels

Land drainage channels constitute a vast network of aquatic habitat utilized by fish. Surveys of these channels (dykes) in Broadland in eastern England reveal similar fish communities whether the dykes drain grazing marsh, fenland or arable land, though there are indications of poor recruitment in the recently deeply drained dykes of arable land. The species encountered most predictably were pike, roach, eel and tench. In terms of biomass, pike ranked top, followed by tench, bream and roach. Eels could not be sampled quantitatively but they must rank in the top five species in terms of biomass. Ordination and other analyses of community structure revealed several physicochemical and biotic factors that correlate with fish distribution and abundance. In particular, the presence of a floating plant mat in summer was associated with low oxygen concentrations at night and the absence of roach. The composition of fish communities in dykes contrasts in a number of ways with those recorded in local rivers and lakes (broads).     

Response of fish and plankton to nutrient loading reduction in eight shallow Danish lakes with special emphasis on seasonal dynamics

1. For 13 years the response of the plankton and fish community to a decline in external phosphorus loading was studied in eight lakes with a mean depth <5 m. We conducted chi‐square analyses of sign of slope (positive or negative) of bimonthly averages of plankton variables for the eight lakes versus time. For fish, we compared results from two periods, i.e. 1989–1994 versus 1994–2001 as less data were available. 2. Fish community structure tended to respond to the lowered concentration of total phosphorus (TP), although not all changes were significant. While catch per unit effort (multi‐mesh sized gill nets) of cyprinids (especially bream, Abramis brama and roach, Rutilus rutilus) was highest in the first 5‐year period, the quantitative importance particularly of perch (Perca fluviatilis), pike (Esox lucius) and rudd (Scardinius erythropthalmus), a littoral species, increased significantly after 1994. 3. No changes occurred in zooplankton biomass, except for an increase in November and December. Biomass of small cladocerans, however, declined during summer and autumn, and the proportion of Daphnia to cladoceran biomass also increased. Average body weight of Daphnia and that of all cladocerans increased. The proportion of calanoids among copepods decreased in summer and the average body weight of cyclopoids and calanoids decreased during summer and autumn/early winter. 4. Total biovolume of phytoplankton declined significantly in March to June and tended to decline in November and December as well, while no significant changes were observed during summer and autumn. Non‐heterocystous cyanobacteria showed a decreasing trend during summer and autumn, while heterocystous cyanobacteria increased significantly in late summer. An increase in late summer was also evident for cryptophytes and chrysophytes, while diatoms tended to decline during most seasons. 5. We conclude that phytoplankton, and probably also fish, responded rapidly to reduced loading, whereas the effect on zooplankton was less pronounced. However, increases in body weight of cladocerans and the zooplankton to phytoplankton biomass ratio during summer indicate reduced top‐down control on zooplankton and enhanced grazing on phytoplankton. This conclusion is supported by a tendency for fish biomass to decline and a shift towards greater dominance by piscivores and, thus, an increased likelihood of predator control of zooplanktivorous cyprinids.     

Effects of different fish species and biomass on plankton interactions in a shallow lake

Thirty-six enclosures, surface area 4 m², were placed in Little Mere, a shallow fertile lake in Cheshire, U.K. The effects of different fish species (common carp, common bream, tench and roach) of zooplanktivorous size, and their biomass (0, 200 and 700 kg ha^–1) on water chemistry, zooplankton and phytoplankton communities were investigated. Fish biomass had a strong effect on mean zooplankton size and abundance. When fish biomass rose, larger zooplankters were replaced by more numerous smaller zooplankters. Consequently phytoplankton abundance rose in the presence of higher densities of zooplanktivorous fish, as zooplankton grazing was reduced. Fish species were also significant in determining zooplankton community size structure. In enclosures with bream there were significantly greater densities of small zooplankters than in enclosures stocked with either carp, tench and, in part, roach. When carp or roach were present, the phytoplankton had a greater abundance of Cyanophyta than when bream or tench were present. Whilst top-down effects of fish predation controlled the size partitioning of the zooplankton community, this, in turn apparently controlled the bottom-up regeneration of nutrients for the phytoplankton community. At the zooplankton–phytoplankton interface, both top-down and bottom-up processes were entwined in a reciprocal feedback mechanism with the extent and direction of that relationship altered by changes in fish species. This has consequences for the way that top-down and bottom-up processes are generalised.     

Littoral age 0+ fish distribution in relation to multi-scale spatial heterogeneity of a deep-valley reservoir

Littoral age 0+ fish were studied with respect to spatio-temporal heterogeneity in the deep-valley ?ímov Reservoir (Czech Republic) from June to October 2007 using point abundance sampling by electrofishing. The abundance and diversity of age 0+ fish in different types of littoral habitats were examined along the longitudinal gradient of the reservoir during daytime. The impact of some physical attributes of the studied littoral habitats, e.g. slope steepness and substrate size, along with the season was the main factors affecting the distribution of age 0+ fish. The level of structural complexity was not a strong determinant, because the overall diversity and structural complexity of the available littoral habitats were relatively too low to have greater impact on the age 0+ fish distribution. The physical factors markedly influenced the spatial segregation between the two most important taxa in the reservoir—percids and cyprinids. Perch was the only representative of age 0+ percids in the littoral zone, which occupied steep-sloped habitats early in the season. In contrast, gently sloped habitats were mainly occupied by cyprinids later in the season. Species diversity was reflected in the occurrence of age 0+ cyprinids, achieving a maximum in gently sloped habitats in October. The effect of the longitudinal gradients on age 0+ fish distribution and diversity was generally far less significant, but was rather decisive during the period of a pronounced longitudinal trophic gradient during the summer season.     

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.     

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.     

Migration of bream between the main channel and floodplain lakes along the lower River Rhine during the connection phase

Mature bream (>30 cm L_F) dominated the fish communities in floodplain lakes along the lower River Rhine in The Netherlands, in terms of biomass, in all lakes at all times. In some lakes bream made up 99% of the total fish biomass. While immature, medium–sized bream (10-30 cm) and white bream and roach were abundant in the main channel of the river, these species were almost absent from the floodplain lakes. Inundation of the floodplains did not lead to spatial homogenization of species and length distributions. Small bream (<10 cm) left the floodplains to become resident in the main channel until maturity. Once mature (c. 30 cm), they returned to the lakes during the next inundation and stayed there. Mature bream that moved into the lakes were significantly smaller and had a lower condition than the resident bream. Turbid clay-pits were probably important sources of bream for the lower Rhine and the recruitment of the 0+ year group of bream to the riverine population was influenced by their discharge pattern.     

Top-down effects of underyearling fish on a phytoplankton community

1. The influence of underyearling (0+) fish on planktonic algal biomass and composition was examined during June–August in eight large enclosures (40 m³). Averaged for the entire period, Daphnia densities, transparency and phosphate concentrations were significantly higher ( P < 0.05) and chlorophyll a values significantly lower in fishless controls than in enclosures with fish (bleak ( Alburnus alburnus ), roach ( Rutilus rutilus ) or perch ( Perca fluviatilis )).
2. Enclosures without fish demonstrated a more heterogeneous algal community structure than enclosures with fish. Desmidiales and dinoflagellates were more abundant in controls, whereas green algae dominated the phytoplankton community in enclosures with fish. Larger grazing-resistant algae occurred most frequently in enclosures without fish.
3. The importance of cascading trophic interactions was demonstrated even under strong eutrophic conditions. The strongest effects on Daphnia densities, phytoplankton biomass and nutrients were observed in enclosures with perch, with weaker effects in enclosures with cyprinids. Differences in initial fish size and species accounted for the top-down effects observed. The results of this study emphasize the importance of 0+ perch as a key predator in structuring the plankton community in lakes.