Individual specialization and trophic adaptability of northern pike (Esox lucius): an isotope and dietary analysis

Northern pike (Esox lucius) are often considered to be specialist piscivores, but under some circumstances will continue to eat invertebrates as adults. To examine effects of fish assemblage composition on the trophic ecology of pike, we combined stable isotope analysis (SIA) of carbon and nitrogen and stomach content analysis (SCA) on pike from five lakes in northern Alberta, three of which contain only pike (“pike-only”) and two that also contain yellow perch (Perca flavescens) or white sucker (Catostomus commersoni) (“pike-other”). Fish were more important as prey and empty stomachs, which often characterize piscivores, were significantly more frequent in pike-other than in pike-only lakes. However, even though invertebrates were more important for pike in pike-only lakes, SIA and SCA indicated that invertebrates were also an important component of pike diets in pike-other lakes. SIA and SCA also revealed considerable intrapopulation variation in trophic ecology, with individuals in some populations differing by as much as two trophic levels. Comparisons of stomach contents and isotope signatures of the same fish suggested that within these variable populations, specialization on invertebrates or fish was a long-term trait of some individuals. SIA indicated that trophic position increased and diets shifted to a greater importance of littoral prey as pike grew in pike-only lakes, but not in lakes with other fish present. Trophic adaptability in northern pike is expressed at both the population level, where the trophic ecology is sensitive to differences in prey regimes, and at the organismal level, in the form of intrapopulation variation and individual specialization. Received: 1 July 1998 / Accepted: 3 February 1999     

Alternative equilibria in shallow lakes

The turbidity of lakes is generally considered to be a smooth function of their nutrient status. However, recent results suggest that over a range of nutrient concentrations, shallow lakes can have two alternative equilibria: a clear state dominated by aquatic vegetation, and a turbid state characterized by high algal biomass. This bi-stability has important implications for the possibilities of restoring eutrophied shallow lakes. Nutrient reduction alone may have little impact on water clarity, but an ecosystem disturbance like foodweb manipulation can bring the lake back to a stable clear state. We discuss the reasons why alternative equilibria are theoretically expected in shallow lakes, review evidence from the field and evaluate recent applications of this insight in lake management.     

Multiplicity of stable states in freshwater systems

It is shown with the use of minimal models that several ecological relationships in freshwater systems potentially give rise to the existence of alternative equilibria over a certain range of nutrient values. The existence of alternative stable states has some implications for the management of such systems. An important consequence is that signs of eutrophication are only apparent after the occurrence of changes that are very difficult to reverse. Reduction of the nutrient level as a measure to restore such systems gives poor results, but biomanipulation as an additional measure can have significant effects, provided that the nutrient level has been reduced enough to allow the existence of a stable alternative clear water equilibrium.     

Community resistance and change to nutrient enrichment and fish manipulation in a vegetated lake littoral

1. High biomass of macrophytes is considered important in the maintenance of a clear‐water state in shallow eutrophic lakes. Therefore, rehabilitation and protection of aquatic vegetation is crucial to the management of shallow lakes. 2. We conducted field mesocosm experiments in 1998 and 1999 to study community responses in the plant‐dominated littoral zone of a lake to nutrient enrichment at different fish densities. We aimed to find the threshold fish biomass for the different nutrient enrichment levels below which large herbivorous zooplankton escapes control by fish. The experiments took place in the littoral of Lake Vesijärvi in southern Finland and were part of a series of parallel studies carried out jointly at six sites across Europe. 3. In 1998, when macrophyte growth was poor, a clear‐water state with low phytoplankton biomass occurred only in unenriched mesocosms without fish or with low fish biomass (4 g fresh mass m^?2). Both nutrient enrichment and high fish biomass (20 g fresh mass m^?2) provoked a turbid water state with high planktonic and periphytic algal biomass. The zooplankton community was dominated by rotifers and failed to control the biomass of algae in nutrient enriched mesocosms. The littoral community thus had low buffer capacity against nutrient enrichment. 4. In 1999, macrophytes, especially free‐floating Lemna trisulca L., grew well and the zooplankton community was dominated by filter‐feeding cladocerans. The buffer capacity of the littoral community against nutrient enrichment was high; a clear‐water state with low phytoplankton biomass prevailed even under the highest nutrient enrichment. High grazing rates by cladocerans, together with reduced light penetration into the water caused by L. trisulca, were apparently the main mechanisms behind the low algal biomass. 5. Effects of fish manipulations were less pronounced than effects of nutrient enrichment. In 1999, clearance rates of cladocerans were similar in fish‐free and low‐fish treatments but decreased in the high‐fish treatment. This suggests that the threshold fish biomass was between the low‐ and high‐fish treatments. In 1998, such a threshold was found only between fish‐free and low‐fish treatments. 6. The pronounced difference in the observed responses to nutrient enrichment and fish additions in two successive years suggests that under similar nutrient conditions and fish feeding pressure either clear or turbid water may result depending on the initial community structure and on weather.     

The restoration of shallow eutrophic lakes,and the role of northern pike,aquatic vegetation and nutrient concentration

The feasibility of biomanipulation is related particularly to reducing the production and recruitment of planktivorous fish stocks. For assessing the level of predation needed to suppress planktivorous fish stocks, the relation between fish P and B, on one hand, and nutrient concentration on the other were analyzed. The carrying capacity of shallow lakes in terms of biomass is related to the total phosphorus concentration and nature of the lake bottom substrate. The production of planktivorous fish was 60–80% of the maximum carrying capacity. It is argued that aquatic vegetation and northern pike are effective tools to maintain water quality, but these are limited by the maximum nutrient concentration aquatic vegetation can sustain.     

Information Transfer across Fish Shoals under Predator Threat

The hypothesis that shoaling fish can obtain information about a predator's approach from changes in the behaviour of other shoal members was tested in an experiment in which receiver minnows (Phoxinus phoxinus) behind a one-way-mirror could observe the reaction of transmitter minnows threatened by the stalk of a pike (Esox lucius) model. Although the receiver minnows were out of visual contact with the pike model they decreased their foraging behaviour and started hiding when the pike model came towards the feeding patch of the transmitter fish. The presence of skittering behaviour and inspection behaviour in the transmitter but not in the receiver fish suggests that individual confirmation of the predator does play a role in determining the nature of the anti-predator response. Nevertheless there is clear evidence that receiver fish modify their own behaviour on the basis of information obtained from individuals which have seen a predator.     

The consequences of a drastic fish stock reduction in the large and shallow Lake Wolderwijd,The Netherlands. Can we understand what happened?

In 1990 an experiment started in the large and shallow lake Wolderwijd (2700 ha, mean depth 1.5 m) to improve the water quality. About 75% of the fish stock was removed (425 000 kg fish). The fish was mainly composed of bream and roach. In May 600000 young pikes (3–4 cm) were introduced.In May 1991 the water became very clear (Secchi depth 1.8 m) during a spring bloom of large Daphnia. Then the grazing by zooplankton was eight times higher than the primary production of algae and the total suspended matter concentration became very low. Compared to the situation before the fish reduction, the grazing had increased only slightly, while the primary production had decreased significantly in early spring. The fish stock reduction might have contributed to the reduction in primary production by a reduced internal nutrient load. The clear water period lasted six weeks. Daphnia disappeared in July due to food limitation, the algal biomass increased and the Secchi depth became 50 cm. Daphnia did not recover during summer, due to predation that was not caused by 0 + fish but by the mysid shrimp Neomysis integer. Neomysis could develop abundantly, because of the reduced biomass of the predator perch. The production of young fish had been low because of the cold spring weather. The cold weather was probably also responsible for the slow increase in density of macrophytes. After 1991, perch probably can control Neomysis. Due to lack of spawning places and shelter for 0 + pike, pike was probably not able to control the production of 0 + fish. In a lake of this scale, it will not be easy to get more than 50% coverage of macrophytes, which seems necessary to keep the algal biomass low by nutrient competition. Therefore, we expect also in the future a decrease in transparency in the summer. Locally, especially near Characeae, the water might stay clear.     

Restoration by biomanipulation in a small hypertrophic lake: first-year results

Biomanipulation was carried out in order to improve the water quality of the small hypertrophic Lake Zwemlust (1.5 ha; mean depth 1.5 m). In March 1987 the lake was drained to facilitate the elimination of fish. Fish populations were dominated by planktivorous and benthivorous species (total stock c. 1500 kg) and were collected by seine- and electro-fishing. The lake was subsequently re-stocked with 1500 northern pike fingerlings (Esox lucius L.) and a low density of adult rudd (Scardinius erythrophthalmus). The offspring of the rudd served as food for the predator pike. Stacks of Salix twigs, roots of Nuphar lutea and plantlets of Chara globularis were brought in as refuge and spawning grounds for the pike, as well as shelter for the zooplankton.The impact of this biomanipulation on the light penetration, phytoplankton density, macrophytes, zooplankton and fish communities and on nutrient concentrations was monitored from March 1987 onwards. This paper presents the results in the first year after biomanipulation.The abundance of phytoplankton in the first summer (1987) after this biomanipulation was very low, and consequently accompanied by increase of Secchi-disc transparency and drastic decline of chlorophyll a concentration.The submerged vegetation remained scarce, with only 5 % of the bottom covered by macrophytes at the end of the season.Zooplankters became more abundant and there was a shift from rotifers to cladocerans, comprised mainly of Daphnia and Bosmina species, the former including at least 3 species.The offspring of the stocked rudd was present in the lake from the end of August 1987. Only 19% of the stocked pike survived the first year.Bioassays and experiments with zooplankton community grazing showed that the grazing pressure imposed by the zooplankton community was able to keep chlorophyll a concentrations and algal abundance to low levels, even in the presence of very high concentrations of inorganic N and P. The total nutrient level increased after biomanipulation, probably due to increased release from the sediment by bioturbation, the biomass of chironomids being high.At the end of 1987 Lake Zwemlust was still in an unstable stage. A new fish population dominated by piscivores, intended to control the planktivorous and benthivorous fish, and the submerged macrophytes did not yet stabilize.     

Invasive pike establishment in Cook Inlet Basin lakes,Alaska: diet,native fish abundance and lake environment

Northern pike (Esox lucius) were introduced to the northern Susitna Basin of south-central Alaska in the 1950’s, and have since spread throughout the upper Cook Inlet Basin. Extirpations of several native fish populations have been documented in this area. It is hypothesized here that invasive pike remodel the ecology of lakes by removing vulnerable prey types and that these changes are reflected in the diet of invasive pike. Trends in pike diet suggest that pike switch to less desirable but more abundant macroinvertebrate prey as preferred fish prey are eliminated. The impacts of pike introduction were studied in detail for one species of resident fish, the threespine stickleback (Gasterosteus aculeatus). Stickleback abundance decreases as pike invasion progresses. Conductivity is a significant environmental predictor of stickleback abundance, with higher conductivity apparently mitigating population reduction. Higher conductivity water may lessen the physiological costs of developing more robust armor, which reduces vulnerability to predation. Maximum lake depth also appears to predict stickleback abundance, though this trend was only marginally significant. Deeper lakes may provide an open-water refuge from pike predation by allowing stickleback to exist outside of the pike inhabited littoral zone. These findings indicate the importance of diverse habitat types and certain chemical and physical characteristics to the outcome of invasion by fish predators.     

Fish–duck interactions in boreal lakes in Finland as reflected by abundance correlations

We studied the hypothesis that fish play an important role in lake use by ducks (pairs and broods) in boreal lakes. The study was based on densities of different duck and fish species in 28 boreal lakes in southern Finland. We focused on the three most common duck species (mallard Anas platyrhynchos, green-winged teal A. crecca and common goldeneye Bucephala clangula) and on the three most common fish species (perch Perca fluviatilis, roach Rutilus rutilus and pike Esox lucius) in the region. We considered both competitive and predatory interactions between ducks and fish, the perch and roach being potential competitors with ducks and the pike a potential predator of ducks. We found a negative association between green-winged teal brood density and total fish density, the other duck species having only a weak association with total fish density. When the three fish species were considered separately, a negative association, suggesting food competition, was found between perch, green-winged teal and goldeneye, whereas the role of roach as a food competitor seemed to be of minor importance. We did not find any clear signs of predatory effects of pike on ducks. Our results suggest that food competition is a more important factor than pike predation in affecting lake use by ducks in oligotrophic boreal environments in southern Finland.     

Terrestrial contributions to Afrotropical aquatic food webs: The Congo River case

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Salinity Induced Regime Shift in Shallow Brackish Lagoons

In brackish lagoons, Daphnia is replaced by calanoid copepods (Eurytemora affinis, Acartia spp.) and rotifers when a certain threshold (depending on, for instance, fish density) is reached. We hypothesize that loss of Daphnia induces a regime shift from clear to turbid at high nutrient concentrations. We conducted a factorial designed enclosure experiment with contrasting salinities (0–16‰), low fish predation (one three-spined stickleback, Gasterosteus aculeatus, m⁻²) and three levels of nutrient loading in a shallow brackish lagoon. A change point analysis suggests a strong regime shift from a clear to a turbid state at 6–8‰ salinity at low and high loading, but not for the control. From the low to the high salt regime, chlorophyll a (Chla), Chla:total phosphorus (TP) and Chla:total nitrogen (TN) ratios shifted highly significantly for all nutrient treatments, and the bacterioplankton production followed the changes in Chla. These changes occurred parallel with a shift from cladoceran and cyclopoid copepod to rotifer dominance. Monitoring data from 60 Danish brackish lagoons show increasing Chla with increasing TP and TN as well as interactive effects of TN and salinity, peaking at intermediate salinity. A relatively weak effect of salinity at low nutrient concentrations and the stronger effect at intermediate high salinity are in accordance with the experimental results. However, these data suggest a lower salinity threshold than in the experiment, which may be explained by a higher fish density. Our results have implications for the management of coastal lagoons both at present and in a future (predicted) warmer climate: (1) improved water quality can be obtained by reducing the nutrient loading or enhancing the freshwater input to a level triggering a shift to Daphnia dominance (typically <2‰), (2) fish manipulation is probably not a useful tool for brackish lagoons, unless the salinity is below the threshold for a potential shift to a clear Daphnia dominated state, and (3) more abrupt changes will expectedly occur in low-saline coastal lagoons at increasing salinity during summer in a future warmer climate.     

Population differences in the reaction of minnows to alarm substance*

Experiments showed that minnows, Phoxinus phoxinus, sympatric with pike, Esox lucius, responded more vigorously to alarm substance than minnows from a population with no experience of pike predation in the wild. Minnows from the pike-sympatric (Dorset) population were more likely to hide and less likely to risk feeding than their pike-allopatric (Gwynedd) counterparts. The reaction to alarm substance in the pike-sympatric population was further increased when it was presented along with the visual stimulus of a ‘stalking’ model pike. When the Dorset minnows experienced both alarm substance and the pike model together they reduced their inspection behaviour to a level below that of the Gwynedd minnows. Minnows from the Gwynedd (pike-allopatric) population displayed increased levels of shoaling in the treatments in which alarm substance was used.     

High levels of mercury in biota of a new Prairie irrigation reservoir with a simplified food web in Southern Alberta,Canada

This study examined mercury levels in northern pike (Esox lucius) from the Twin Valley Reservoir in southern Alberta, 2 years after construction in 2003. The hypothesis was tested that mercury concentrations in pike from the reservoir are significantly higher than in pike from the nearby Oldman River. Mercury concentrations in muscle tissue (0.37–1.54 ppm) generally exceeded the consumption guideline of 0.5 ppm total mercury (THg), and were significantly higher (3.5-fold) than northern pike mercury concentrations in the Oldman River. In addition, these levels exceeded (up to 2-fold) previously published data from other reservoirs of the northern hemisphere. Gill-netting followed by stomach contents and stable isotope analysis revealed a very simple food web. No zooplanktivorous fish species were present, and the benthic fish community contained only few white sucker (Catostomus commersoni). Thus, the reservoir almost completely lacked forage fish, and the pike fed primarily on amphipods (Gammarus lacustris and Hyalella azteca), whose average mercury concentrations were 0.21 ppm. An observed low growth trajectory of northern pike in this reservoir may reflect low growth efficiency as a result of their invertebrate diet.     

What do the empty stomachs of northern pike (<Emphasis Type="Italic">Esox lucius</Emphasis>) reveal? Insights from carbon (δ<Superscript>13</Superscript>C) and nitrogen (δ<Superscript>15</Superscript>N) stable isotopes

The frequency of individuals with empty stomachs (FES) can vary greatly among northern pike populations. However, the FES has only seldom been analyzed in this species and its meaning is still not fully understood. It has been suggested that a high FES may reflect a strongly piscivorous behaviour while low FES could reflect a higher utilization of invertebrates. We compared the stomach contents and the trophic position of northern pike in 16 populations of individuals feeding mainly on fish or benthic invertebrates. We tested the hypothesis that northern pike with empty stomachs or with fish in their stomachs have a higher trophic position than individuals feeding on invertebrates. Carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope signatures were used to estimate the trophic position of individuals. We found no significant difference in the trophic position among piscivores, invertebrate feeders, and northern pike with empty stomachs. The average trophic position of northern pike was high (mean ± SD = 4.3 ± 0.4, n = 66) and was correlated with total length. These results indicate that, although invertebrates could be an important part of the diet of northern pike in Canadian Shield lakes, fish are still the dominant prey. Hence, feeding on invertebrates in our study lakes would reflect an opportunistic rather than a specialized feeding strategy.     

Wetland macrophyte diversity, trophic status and phytoplankton： a case study of the cascading effects of an invasive herbivore

Shallow water bodies can exist in alternative stable states, a clear water state with high coverage of macrophytes or a turbid state with high phytoplankton biomass. The alternative equilibria hypothesis has been proposed to explain the occurrence of the alternative stable states (Scheffer et al., 1993)[1], which assumes that: 1),     

Evaluating stocking of YOY pike Esox lucius as a tool in the restoration of shallow lakes

1. Stocking of piscivores in shallow, eutrophicated lakes to reduce cyprinid densities is a common approach in lake restorations. Young‐of‐the‐year (YOY) pike Esox lucius are frequently used to reduce cyprinid densities, but their effectiveness is equivocal. This study uses a simple model to assess the efficacy of pike stocking as a lake restoration tool. 2. The model evaluates YOY pike consumption in relation to different survival and growth patterns, timing of stocking, degree of piscivory of pike as well as YOY cyprinid growth patterns and productivity. The parameters used in the model are obtained from field surveys in eight study lakes stocked with YOY pike as well as from the literature. 3. Our model showed that all parameters studied were important for predicting the effects of pike stocking on cyprinids. In particular, body size at stocking, cyprinid production and pike survival were good predictors of biomanipulation success. 4. However, our model showed that only under very specific circumstances will stocking of YOY pike result in a reduction of YOY cyprinid densities large enough to affect water quality. 5. The findings of this study question the efficacy of pike stocking as an appropriate and reliable tool for restoration programmes in shallow lakes, and indicate that when used efforts should be made to optimize the timing of stocking in relation to YOY cyprinid production and to increase the stocking body size of the pike.     

Modeling nutrients,oxygen and critical phosphorus loading in a shallow reservoir in China with a coupled water quality – Macrophytes model

Many macrophyte-dominated clear lakes switch to a phytoplankton-dominated turbid state when the lake becomes eutrophic. An existing Yuqiao Reservoir Water Quality Model (YRWQM) and the macrophyte submodel were coupled to simulate the effect of submerged macrophytes on nutrients and dissolve oxygen cycles in a shallow reservoir in China. The level of phosphorus loading in a transition from a clear to turbid state was addressed using the integrated model. The model runs from seedling establishment until dying out, from March 1 to July 18 in 2009. The simulations were performed for a contingent range of P loadings, starting from three different initial conditions. The results indicated that the integrated model improves accuracy of predictions compared to YRWQM. The concentrations of nutrients declined slightly during the macrophyte growth period in the reservoir and dissolved oxygen increased slightly. Although nutrient concentrations increased by submerged macrophyte release during the extinction period, the effect on the nutrients was less than that of transfer with nutrient-rich water. More released nutrients may enhance increases in substantial abundance. The critical phosphorus loading level during a switch from the clear to turbid state was estimated by these scenarios. The threshold for the switch is ∼6.1 mgP m⁻² d⁻¹ with an initial total phosphorus concentration of 160 μg l⁻¹. Moreover, the results demonstrated that the switch was also dependent on the initial total phosphorus concentration. These results suggest that the reservoir in a clear water state is at risk of a switch as nutrient levels are close to the critical levels.     

Dynamics of a Boreal Lake Ecosystem during a Long-Term Manipulation of Top Predators

We assessed the long-term (16 years) effects of introducing piscivores (northern pike) into a small, boreal lake (Lake 221, Experimental Lakes Area) containing abundant populations of two planktivorous fish species. After the introduction, pearl dace were extirpated and yellow perch abundance was greatly reduced. Daphnia species shifted from D. galeata mendota to larger bodied Daphnia catawba, but the total zooplankton biomass did not increase, nor did the biomass of large grazers such as Daphnia. Phytoplankton biomass decreased after the northern pike introduction, but increased when northern pike were partially removed from the lake. Phosphorus (P) excretion by fish was ∼0.18 mg P m⁻² d⁻¹ before pike addition, declined rapidly to approximately 0.03–0.10 as planktivorous perch and dace populations were reduced by pike, and increased back to premanipulation levels after the pike were partially removed and the perch population recovered. When perch were abundant, P excretion by fish supported about 30% of the P demand by primary producers, decreasing to 6–14% when pike were abundant. Changes in phytoplankton abundance in Lake 221 appear to be driven by changes in P cycling by yellow perch, whose abundance was controlled by the addition and removal of pike. These results confirm the role of nutrient cycling in mediating trophic cascades and are consistent with previous enclosure experiments conducted in the same lake.     

Correlation between the feeding time of the pike (Esox lucius) and the dispersion of a school of Leucaspius delineatus

Summary The illumination level at which the pike (Esox lucius) eats most Leucaspius delineatus was compared to that at which the school of Leucaspius disperses.The pike easts Leucaspius predominantly when there is little light (50–75% of its entire prey at less than 1 Lux).The school of Leucaspius gradually disperses at less than 1 Lux.A comparison of the results shows that by and large the pike captures Leucaspius at illumination levels at which the school of Leucaspius is no longer completely unified.As both the ability of the pike to catch prey and of Leucaspius to form a school depend on the optic capabilities of the animals it can be supposed that the predator is optically superior to its prey at low illumination levels and profits from this advantage when catching its prey.With many thanks to Dr. G. Rüppell for his supervision of the work reported in this paper