Fennel Pondweed Research Articles

Monitoring of heavy metal burden in mute swan (Cygnus olor).

Concentrations of heavy metals (especially arsenic, cadmium, chromium, copper, mercury and lead) were measured in the contour (body) feathers of mute swans (Cygnus olor) and in its nutrients (fragile stonewort [Chara globularis], clasping leaf pondweed [Potamogeton perfoliatus], Eurasian watermilfoil [Myriophyllum spicatum], fennel pondweed [Potamogeton pectinatus]) to investigate the accumulation of metals during the food chain. The samples (17 feathers, 8 plants) were collected at Keszthely Bay of Lake Balaton, Hungary. Dry ashing procedure was used for preparing of sample and the heavy metal concentrations were analysed by inductively coupled plasma optical emission spectroscopy (ICP-OES). Copper (10.24 ± 2.25 mg/kg) and lead (1.11 ± 1.23 mg/kg) were detected the highest level in feathers, generally, the other metals were mostly under the detection limit (0.5 mg/kg). However, the concentrations of the arsenic (3.17 ± 1.87 mg/kg), cadmium (2.41 ± 0.66 mg/kg) and lead (2.42 ± 0.89 mg/kg) in the plants were low but the chromium (198.27 ± 102.21 mg/kg) was detected in high concentration.

The effect of herbivores on genotypic diversity in a clonal aquatic plant

In clonal plants, vegetative parts may outcompete seeds in the absence of disturbance, limiting the build‐up of genotypic diversity through repeated seedling recruitment (RSR). Herbivory may provide disturbance and trigger establishment of strong colonizers (seeds) at the expense of strong competitors (clonal propagules). In the clonal aquatic fennel pondweed Potamogeton pectinatus, two distinct herbivore guilds may modify the dynamics of propagation. In winter, Bewick's swans may deplete patches of tubers, promoting seedling establishment in spring. In summer, seed consumption by waterfowl can reduce the density of viable seeds but grazing may also reduce tuber production and hence facilitate seedling establishment. This study is among the first to experimentally test herbivore impact on plant genotypic diversity. We assess the separate and combined effects of both herbivore guilds on genotypic diversity and structure of fennel pondweed beds. Using microsatellites, we genotyped P. pectinatus from an exclosure experiment and assessed the contribution of herbivory, dispersal and sexual reproduction to the population genetic structure. Despite the predominance of clonal propagation in P. pectinatus, we found considerable genotypic diversity. Within the experimental blocks, kinship among genets decreased with geographic distance, clearly identifying a role for RSR in the maintenance of genotypic diversity within the fennel pondweed beds. However, over a period of five years, none of the herbivory treatments affected genotypic diversity. Hence, sexual reproduction on a local scale is important in this putatively clonal plant and possibly sufficient to ensure a relatively high genotypic diversity even in the absence of herbivores. Although we cannot preclude a role of herbivory in shaping genotypic diversity of a clonal plant, after five years of exclusion of the two investigated herbivore guilds no measurable effect on genotypic diversity was detected.

Clonal diversity and spatial genetic structure of Potamogeton pectinatus in managed pond and river populations

Higher levels of genetic diversity of river macrophytes are expected in downstream parts because of potential accumulation of various genotypes from upstream sites. We assessed the clonal diversity and spatial genetic structure of fennel pondweed (Potamogeton pectinatus or Stuckenia pectinata) populations with emphasis on the estimation of dispersal via clonal propagules along a river in connection to upstream ponds. We analysed genetic diversity of 354 plant shoots sampled in 2005 and 2006 at three pond and five river sites in the Woluwe river catchment (Belgium). Nine microsatellite DNA markers revealed 88 genets of which 89% occurred in only one site. Clonal propagule dispersal was detected up to 10 km along the river. Few multilocus genotypes were repeatedly present along a major part of the river indicating vegetative spread. Populations of ponds contained a higher amount of clonal diversity, indicating the importance of local seed recruitment. A fine-scaled spatial genetic structure indicated that most seedling recruitment occurred at a distance <5 m in pond populations whereas clones in river sites were unrelated and showed no spatial autocorrelation. The clonal diversity decreased along the river from upstream to downstream due to establishment of few large clones.

Underuse of stopover site by migratory swans

Many migratory birds use a chain of stopover sites to fuel their migration. Under time-minimizing migration, fuelling time and giving-up density at stopovers are predicted to depend on fuelling conditions. Fluctuations in food accessibility likely lead to changes in fuelling conditions, which should in turn be reflected in fuelling time and giving-up density. During their migration, Bewick’s Swans Cygnus columbianus bewickii refuel on belowground tubers of Fennel Pondweed Potamogeton pectinatus in shallow lakes. We studied giving-up density and stop-over use (expressed in bird-days) of Bewick's Swans at an autumn stopover site (Lauwersmeer, The Netherlands) during 1995–2008, as dependent on local environmental conditions. High water levels were hypothesized to restrict access to tuber stocks. High water levels at the stopover site were predicted to lead to higher giving-up densities and less bird-days spent at the stopover. Annual variation in giving-up densities and number of bird-days was strongly associated with year-to-year differences in initial tuber biomass density and number of days with high water levels. As predicted, giving-up density increased and bird-days decreased with the number of days with high water level. We conclude that, in line with time-minimizing migration, changes in fuelling conditions may lead to underuse of a stopover site. Underuse of stopovers by migratory birds has been reported before but only in the sense that more food was left at stopover sites than at wintering sites. In contrast, in our case, dealing with a given stopover site, more food is left behind in some years than in other years.

Human disturbance of Bewick's Swans is reflected in giving‐up net energy intake rate, but not in giving‐up food density

Measuring the food left in experimental trays when study organisms cease feeding on them [so‐called giving‐up densities (GUDs)] is an accepted technique for assessing predation risk and disturbance. However, in natural settings, accessibility and energetic harvest costs may vary spatially, and GUDs may be confounded. In this study, we assessed whether at a heterogeneous site, non‐experimental GUDs could reveal the effect of disturbance. We measured initial and GUDs of tubers of Fennel Pondweed Potamogeton pectinatus, which form here the exclusive food source of Bewick's Swans Cygnus columbianus bewickii during their migratory stopover. We calculated giving‐up net energy intake rates (GUNs) by correcting for biomass accessibility and foraging costs. The study area was a shallow lake consisting of nine creeks, three of which were open to the public (i.e. disturbed). GUDs in creeks open or closed to the public were not significantly different. In contrast, GUNs were generally higher in creeks open to the public, after correcting for initial net energy intake rate. The results suggest that natural GUDs may not reflect the effects of disturbance in heterogeneous habitats. When environmental differences are large within a site, GUNs may be a useful alternative as a behavioural indicator.

Aquatic plant shows flexible avoidance by escape from tuber predation by swans

Deeper burial of bulbs and tubers has been suggested as an escape against below-ground herbivory by vertebrates, but experimental evidence is lacking. As deep propagule burial can incur high costs of emergence after dormancy, burial depth may represent a trade-off between sprouting survival and herbivore avoidance. We tested whether burial depth of subterraneous tubers is a flexible trait in fennel pondweed (Potamogeton pectinatus), facing tuber predation by Bewick's swans (Cygnus columbianus bewickii) in shallow lakes in winter. In a four-year experiment involving eight exclosures, winter herbivory by swans and all vertebrate summer herbivory were excluded in a full-factorial design; we hence controlled for aboveground vertebrate herbivory in summer, possibly influencing tuber depth. Tuber depth was measured each September before swan arrival and each March before tuber sprouting. In accordance with our hypothesis, tuber depth in September decreased after excluding Bewick's swans in comparison to control plots. The summer exclosure showed an increase in tuber biomass and the number of shallow tubers, but not a significant effect on the mean burial depth of tuber mass. Our results suggest that a clonal plant like P. pectinatus can tune the tuber burial depth to predation pressure, either by phenotypic plasticity or genotype sorting, hence exhibiting flexible avoidance by escape. We suggest that a flexible propagule burial depth can be an effective herbivore avoidance strategy, which might be more widespread among tuber forming plant species than previously thought.

Reduced tuber banks of fennel pondweed due to summer grazing by waterfowl

In this study we investigated the effect of summer bird herbivory on the belowground tuber formation of fennel pondweed ( Potamogeton pectinatus L.). Cumulative grazing pressure of four waterfowl species (mute swans, mallards, gadwalls and coots) in the summer was calculated based on timing of grazing and body mass of the grazers. The resulting grazing pressures were significantly negatively correlated with mean autumn tuber biomass in three of the four years of study. Moreover, summer grazing pressures explained more of the variance in tuber densities than water depth, sediment particle size distribution or any interactions of these variables did in the same three years. We propose that herbivory early in the summer has the most substantial impact on the clonal reproduction of macrophytes. Herbivores with a large body mass and early congregation for moulting may be the key waterfowl species in diminishing propagule biomass. Hence, they may present pre-emptive, time-staggered competition to consumers of the belowground biomass in autumn, such as migratory swans and diving ducks.

Cryptic interference competition in swans foraging on cryptic prey

Interference can be defined as the reduction of intake rate caused by the presence of congeneric individuals. However, surrounding congeneric individuals may also accelerate food depletion. Therefore, it is difficult to quantify interference (contest) and exploitative (scramble) competition separately, particularly in the field. In an individual-based model, where food was immediately replenished in patches after a foraging bout, we isolated the effect of interference competition. In this way, we investigated how interference shapes the intake rates of flocking Bewick’s swans, Cygnus bewickii, consuming tubers of fennel pondweed, Potamogeton pectinatus. The model predicted that intake rate declines with increasing swan density, and that the rate of decline is greater in subordinate birds than in dominant birds. Subordinate birds suffered a large reduction in intake rate at high densities because they spent a large proportion of the time avoiding more dominant individuals. A major decline in intake rate of subordinates occurred close to the maximum swan densities observed in the field. We expected subordinates to leave high-density flocks and look for another foraging location: interference competition seemed to regulate the maximum swan flock density in the field. Hence, the mean population intake rate at realized densities was only slightly lower than in the absence of interference. As interference occurred mainly as avoidance behaviour, which is difficult to observe in the field, it might commonly remain cryptic for the observer. Our results may explain other field studies where interference competition seemed to be of lesser importance.

How a bottom‐dweller beats the canopy: inhibition of an aquatic weed (Potamogeton pectinatus) by macroalgae (Chara spp.)

Summary1. Bottom‐dwelling charophytes have been observed to replace canopy‐forming pondweeds within a few years in de‐eutrophied shallow lakes. Competition for bicarbonate (HCO3−) may explain this shift in vegetation dominance but inhibition of pondweeds by Chara spp. through direct competition has not been shown experimentally.2. We tested whether charophytes inhibited growth of fennel pondweed (Potamogeton pectinatus) in the absence of belowground competition by growing plants in pots in mesocosms following a replacement series experimental design. To further understand the role of bicarbonate, we studied main and interactive effects of Chara, light and bicarbonate on P. pectinatus growth in a laboratory study.3. Early in the mesocosm experiment, high charophyte densities had a negative effect on P. pectinatus cover at a time when bicarbonate levels were low. However, bicarbonate levels eventually converged to low levels in all treatments. At final harvest, both species exhibited lower biomasses at higher densities of conspecific pots, indicating that ultimately intraspecific competition was limiting. In a laboratory study, Chara inhibited P. pectinatus the most under a combination of high light and high bicarbonate concentrations, suggesting that Chara may negatively affect P. pectinatus by acting as a general nutrient sink.4. Our results suggest that Chara growth can reduce bicarbonate levels, delaying but not preventing a P. pectinatus growth pulse. Given the recorded inhibition under ample bicarbonate supply, Chara’s ability to act as nutrient sink may contribute to the decline of P. pectinatus under Chara recovery in shallow lakes.

Burial depth distribution of fennel pondweed tubers ( Potamogeton pectinatus) in relation to foraging by Bewick's swans

Deep burial in the sediment of tubers of fennel pondweed ( Potamogeton pectinatus) has been explained in terms of avoidance by escape against consumption by Bewick's swans ( Cygnus columbianus bewickii) in autumn. We therefore expected changes in foraging pressure to ultimately result in a change in the tuber distribution across sediment depth. A trade-off underlies this idea: deep tubers are less accessible to swans but must be larger to meet the higher energy demands of sprouting in spring. To test this prediction, we compared tuber burial depth over a gradient of foraging pressure both across space and across time. Tuber samples were obtained after aboveground plant senescence but before arrival of Bewick's swans. First, we compared the current tuber bank depth profile in a shallow lake with high foraging pressure, the Lauwersmeer, with that in two wetlands with moderate and low foraging pressure. Second, we compared the current tuber burial in the Lauwersmeer with that in the early 1980s when exploitation by swans had just started there. In accordance with our hypothesis, we found significantly deeper burial of tubers under high consumption risk compared to low consumption risk, both when comparing sites and comparing time periods. Since tubers in effect only survive to the next spring, the observed differences in burial depth among sites and over time cannot be a direct result of tuber losses due to consumption by swans. Rather, these observations suggest adaptive responses in tuber burial related to foraging pressure from Bewick's swans in the recent past. We thus propose that fennel pondweed exhibits flexible avoidance by escape, of a kind rarely described for plants, where both phenotypic plasticity and genotype sorting may contribute to the observed differences in tuber burial.

PERSISTENCE OF SPATIAL VARIANCE AND SPATIAL PATTERN IN THE ABUNDANCE OF A SUBMERGED PLANT.

Organisms usually benefit from heterogeneous conditions, but, by doing so, may reduce the degree of heterogeneity. The question therefore arises how heterogeneity is maintained. We investigated within-year spatiotemporal patterns in a monospecific stand of a submerged plant (fennel pondweed, Potamogeton pectinatus), with the novelty that we distinguished between different forms of heterogeneity: spatial variance (the frequency distribution of densities) and spatial pattern (the spatial distribution of densities). We repeatedly measured plant biomass that was affected by swan predation, winter mortality, and summer regrowth. Spatial variance was enhanced mostly by swan foraging, despite the fact that swans appear to exploit patches to the same threshold level. Spatial pattern, which had vanished after swan foraging, reestablished due to spatial pattern in winter mortality and was further enhanced by plant regrowth. We found that variance and pattern each have their own temporal dynamics and are maintained by different biological processes. We therefore advocate that it is pivotal to distinguish between variance and pattern in the study of spatial heterogeneity.

The role of herbivorous water birds in aquatic systems through interactions with aquatic macrophytes, with special reference to the Bewick’s Swan – Fennel Pondweed system

The role of aquatic macrophytes in stimulating biodiversity and maintaining clear waters is currently undisputed. The management of (eutrophic) shallow waters is therefore often directed at (re-)establishing macrophyte domination. In contrast, the role of water birds has long been considered of minor importance for the functioning of fresh water ecosystems. Indeed, in terms of biomass and production, water birds constitute only a minor part of these systems. However, water birds may graze heavily on water plants under certain circumstances, and the question arises whether herbivorous water birds have an important indirect effect on shallow fresh water systems. Mainly illustrated with the interaction between Bewick’s Swans and Fennel Pondweed, we present data on the role that water plants may play in the life of water birds and how water birds may impact water plants’ fitness in terms of survival, production, dispersal and competitive ability. It appears that water plants may be crucial for water birds during periods of high-energy requirements, such as migration. Despite the plants’ costs associated with water bird grazing, the interaction between water birds and water plants varies in nature from an apparent predator–prey relationship to a mutually beneficial interaction depending on the context and the perspective. For the case of the Bewick’s Swan–Fennel Pondweed interaction, regular bird grazing is sustainable and may actually favour the plant’s dispersal. Thus, Bewick’s Swans themselves may in fact play a crucial role in establishing and maintaining the Fennel Pondweed rich staging sites between the swans’ wintering and breeding grounds, which are vital for the swans’ successful migration.

Prediction of bird‐day carrying capacity on a staging site: a test of depletion models

1. The carrying capacity of a site for migratory water birds, expressed in bird-days, can be of particular conservation value. Several attempts have been made to model this carrying capacity using ideal free distribution models such as, for instance, depletion models, in which the distribution is fully determined by exploitative competition. 2. In the tests of depletion models carried out so far, no alternative models were compared; rather, one specific model was tested. We tested whether bird-days were more in accordance with birds depleting the food resource (a1) until a critical food density which just enabled survival or (a2) until a threshold food density which renders the site as profitable as an alternative site; and birds (b1) satisfying their daily requirements or (b2) maximizing daily intake. 3. We studied Bewick's swans feeding on below-ground tubers of fennel pondweed in one part of an autumn staging site. In most years between 1995 and 2005, we measured tuber biomass densities around September, November and March, and counted swans daily during their stopover in October. 4. The best fit between observed and predicted bird-days was obtained by assuming that the swans were maximizing their daily intake and depleting the tubers until a threshold biomass density (which yielded the same energetic return as the alternative food source after accounting for a small part of the initial tuber biomass being out of reach of the swans). Also in line with daily intake maximization, the daily feeding time did not differ from 10 h day(-1), the value predicted for Bewick's swans based on their feeding costs. 5. Our results suggests that the applicable model to calculate carrying capacity may depend strongly on whether birds use a site to stopover or to winter, because it determines whether the birds are more likely to use a threshold or critical food density, and to behave as energy maximizers or satisficers.

Animal–plant–microbe interactions: direct and indirect effects of swan foraging behaviour modulate methane cycling in temperate shallow wetlands

Wetlands are among the most important ecosystems on Earth both in terms of productivity and biodiversity, but also as a source of the greenhouse gas CH(4). Microbial processes catalyzing nutrient recycling and CH(4) production are controlled by sediment physico-chemistry, which is in turn affected by plant activity and the foraging behaviour of herbivores. We performed field and laboratory experiments to evaluate the direct effect of herbivores on soil microbial activity and their indirect effects as the consequence of reduced macrophyte density, using migratory Bewick's swans (Cygnus columbianus bewickii Yarrell) feeding on fennel pondweed (Potamogeton pectinatus L.) tubers as a model system. A controlled foraging experiment using field enclosures indicated that swan bioturbation decreases CH(4) production, through a decrease in the activity of methanogenic Archaea and an increased rate of CH(4) oxidation in the bioturbated sediment. We also found a positive correlation between tuber density (a surrogate of plant density during the previous growth season) and CH(4) production activity. A laboratory experiment showed that sediment sterilization enhances pondweed growth, probably due to elimination of the negative effects of microbial activity on plant growth. In summary, the bioturbation caused by swan grazing modulates CH(4) cycling by means of both direct and indirect (i.e. plant-mediated) effects with potential consequences for CH(4) emission from wetland systems.

The use of a flexible patch leaving rule under exploitative competition: a field test with swans

Learning animals are predicted to use a flexible patch‐leaving threshold (PLT) while foraging in a depletable environment under exploitative competition. This prediction was tested in flock‐feeding Bewick's swans (Cygnus columbianus bewickii) depleting hidden tubers of fennel pondweed (Potamogeton pectinatus) in a two‐dimensional, continuous environment. The swans' patch residence time was measured by combining recordings of the foraging behaviour and movement paths. The tuber biomass density was measured before and after the period of exploitation, using the presumable foraging window of the swans as the scale of measurement. Swan foraging was simulated in order to predict the effects of flexible and fixed PLTs, respectively, on the patch residence time and the spatial heterogeneity of the tuber biomass density. Flexible PLTs were predicted to lead to short and decreasing patch residence times and a decrease in the coefficient of variation in tuber biomass densities, whereas the reverse was generally the case for fixed PLTs. Observed patch residence times did not decrease with time and were intermediate between those predicted for swans with flexible and fixed PLTs. Furthermore, an increase of the coefficient of variation in the tuber biomass density was observed. Given the observed giving‐up biomass densities the most likely model was one with swans with a fixed rather than a flexible PLT. These results point at factors that may affect the spacing behaviour or constrain the use of a flexible PLT in swans.

Foraging costs and accessibility as determinants of giving‐up densities in a swan‐pondweed system

We measured the patch use behaviour of Bewick's swans (Cygnus columbianus bewickii) feeding on below ground tubers of fennel pondweed (Potamogeton pectinatus). We compared the swans’ attack rates, foraging costs and giving‐up densities (GUDs) in natural and experimental food patches that differed in water depth. Unlike most studies that attribute habitat‐specific differences in GUDs to predation risk, food quality or foraging substrate, we quantified the relative importance of energetic costs and accessibility. Accessibility is defined as the extent to which the animal's morphology restricts its harvest of all food items within a food patch. Patch use behaviours were measured at shallow (ca 0.4 m) and deep (ca 0.6 m) water depths on sandy sediments. In a laboratory foraging experiment, when harvesting food patches, the swan's attack rate (m3 s−1) did not differ between depths. In deep water the energetic costs of surfacing, feeding and trampling were 1.13 to 1.21 times higher than in shallow water with a tendency to spend relatively more time trampling, the most expensive activity. Taking time allocation as measured in the field into account, foraging in deep water was 1.26 times as expensive as in shallow water. In the lake the GUD in shallow water was on average 12.9 g m−2. If differences in energetic costs were the only factor determining differences in GUDs, then the deep water GUD should be 14.2 g m−2. Instead, the mean GUD in deep water was 20.2 g m−2, and therefore energetic costs explain just 18% of the difference in GUDs. At deep sites, 24% of tuber biomass was estimated to be out of reach, and we calculated a maximum accessible foraging depth of 0.86 m. This is close to the published 0.84 m based on body measurements. A laboratory experiment with food offered at a depth of 0.89 m confirmed that it was just out of reach. The agreement between calculated and observed maximum accessible foraging depths suggests that accessibility largely explains the remaining difference in GUDs with depth, and it confirms the existence of partial prey refuges in this system.

Time and energy constraints in demanding phases of the annual cycle: an example of time limitation in refuelling migratory swans

The daily metabolizable energy intake of an animal is potentially limited by either the available feeding time or by its capacity to process energy. Animals are generally considered not to be time‐limited but rather to be energy‐processing‐limited. This is concluded from the common observation that an animal's feeding time per day increases with a decrease in food density. We argue that such changes in feeding time are in theory also expected when no constraints are operating. Thus, a study of the constraints on energy intakes of free‐living animals should be performed during demanding phases of the year. As an example, we collected data on time and energy budgets of Bewick's swan (Cygnus columbianus bewickii) refuelling during migration on fennel pondweed (Potamogeton pectinatus) tubers in two years differing two‐fold in tuber biomass density. As predicted by time limitation, the feeding time (defined as the time with the head submerged) did not change in response to a change in food biomass density, both within and between years (averaging 12.2 h d−1). Contrary to energy‐processing limitation, and again in line with time limitation, the daily metabolizable energy intake varied, being greater in the year with high than in the year with low food densities. We conclude that more studies are needed of animals operating under demanding conditions before it can be assessed whether free‐living animals are generally energy‐processing‐ or time‐limited.

Digestive plasticity in Mallard ducks modulates dispersal probabilities of aquatic plants and crustaceans

Summary The consequences of plastic responses of the avian digestive tract for the potential of birds to disperse other organisms remain largely uninvestigated. To explore how a seasonal diet switch in Mallard (Anas platyrhynchos L.) influences their potential to disperse plants and invertebrates, we recorded the retention time of markers, following exposure to two diets of contrasting digestibility (trout chow vs seeds). We then recorded the retrieval and germination of Fennel Pondweed (Potamogeton pectinatus L.) seeds and Brine Shrimp (Artemia franciscana Kellogg) cysts ingested by the same birds. Gut passage rates of markers were increasingly longer in birds on the seed‐based, high‐fibre diet and shorter in birds on the animal‐based, low‐fibre one. Propagule digestibility, and thus survival to gut passage, differed between diet groups, with more seeds and fewer cysts retrieved from ducks on the animal‐based diet. Germination decreased with retention time, but was not affected by diet. Differences in passage rates of markers but not of seeds and cysts suggest no change in dispersal distances of plants and invertebrates between seasons, while differences in digestibility would affect the numbers of propagules dispersed.

Effects of waterfowl, large fish and periphyton on the spring growth of Potamogeton pectinatus L. in Lake Mogan, Turkey

It has been argued that waterfowl and fish may threaten growth of submerged macrophytes, especially in spring during the early growth phase when plant biomass is low. A small reduction of biomass at that time might delay growth or decrease subsequent productivity. We investigated the impact of waterfowl and large fish on the spring growth of fennel pondweed (Potamogeton pectinatusL.) by employing an exclosure experiment in the macrophyte-dominated clear-water Lake Mogan, Turkey. Birds and large fish were excluded from eight plots and both in situvegetation and macrophytes kept in pots were compared to eight open plots. Also, to investigate the effect of periphyton on plant growth it was removed from half of the pot plants. Exclusion of waterfowl and fish may decrease predation on macroinvertebrates, which in turn may affect periphyton, and macrophyte growth, why macroinvertebrates also were sampled. Waterfowl density was high (15–70 ind. of coot, Fulica atraL. ha−1), abundance of submerged plants was also high with a surface coverage of 70–80%, and benthivorous fish were present, mainly tench, (Tinca tincaL.) and carp, (Cyprinus carpioL.). Exclusion of waterfowl and large fish did not significantly affect the spring growth of pondweed; neither plants growing in situnor kept in pots. Removal of periphyton from the plants in the pots did not favour growth. The density of macroinvertebrates was not affected by the exclusion of waterfowl and large fish, but it was positively related to aboveground biomass of fennel pondweed. We suggest that even if waterfowl and large fish are in high densities, their effect on fennel pondweed spring growth in lakes with abundant submerged vegetation, such as Lake Mogan, is low.

Overcompensation and grazing optimisation in a swan–pondweed system?

Summary1. The general notion is that negative effects of vertebrate herbivores on water plants, which play a key role in freshwaters, prevail, and that positive feedbacks of herbivores on plants are insignificant.2. The most likely systems to find such positive feedbacks are those in which herbivores exert strong feeding pressures on plants during part of the year. Previous theoretical work has suggested that compensatory production occurs when migratory Bewick's swans forage on tubers of fennel pondweed. As a corollary, the swans can exploit the tubers down to a level that maximises their tuber yield.3. In order to test these hypotheses, I measured pondweed tuber biomass on three occasions per year (just before and after foraging, and just before tuber sprouting) in three consecutive years. The 17 sampling sites in the Lauwersmeer (the Netherlands) were classified according to their silt content and water depth. Within four silt‐depth classes, I predicted for each year tuber biomass production and, from that, the optimum foraging threshold that would result in the maximum tuber biomass yield.4. Water depth did not affect tuber production, and silt content only did in one of the 3 years. In accordance with overcompensation predictions, tuber production was higher at plots with moderate foraging pressures than at plots with little or no grazing. However, the winter and summer conditions following the swan foraging had large unpredictable effects on tuber mortality and production.5. These results indicate that overcompensation by fennel pondweed occurs and that Bewick's swans are generally able to profit from it, albeit without fine‐tuning of the foraging threshold to the yield.