Reduction of benthic macroinvertebrates due to waterfowl foraging on submerged vegetation during autumn migration

If present in large numbers, as during migration, herbivorous waterfowlmay reduce the amount of submerged vegetation. Because the vegetation is a keyfactor in shallow eutrophic lakes, removal of the green biomass can be expectedto affect also other biota that depend on the vegetation. We conducted anexperiment to determine how the abundance of chironomids andPisidium sp. were affected by intense foraging ofwaterfowlon the submerged plant Potamogeton pectinatus. This wasdone in Lake Ringsjön in southern Sweden, during the autumn migration ofthe birds. Three treatments, replicated six times, were used: (i) closed cagesthat excluded all waterfowl, (ii) semi-open cages that excluded only largewaterfowl (geese and swans), and (iii) open plots where all waterfowl couldfreely enter. Waterfowl densities were monitored during the experiment. Theresults suggest that the foraging of large waterfowl (swans) had a clearlynegative effect on macroinvertebrate abundance and aboveground biomass ofP. pectinatus. At the end of the experiment, the densityofchironomids was about 46% lower in the open than in the closed cages. Ingeneral, the density of Pisidium sp. tended to be lower inthe open plots. Small waterfowl alone did not seem to affect either thevegetation or macroinvertebrates. We suggest that thePisidium sp. was influenced at an early stage of grazing,when waterfowl foraged on aboveground biomass, whereas chironomids wereaffectedat a later stage, when swans were digging for below-ground tubers.     

Differences in tolerance of pondweeds and charophytes to vertebrate herbivores in a shallow Baltic estuary

It has been suggested that herbivorous waterfowl may be important in shaping aquatic plant communities in shallow wetlands. As such, a shift from canopy forming pondweeds to bottom-dwelling charophytes in a formerly turbid pondweed dominated lake has been partly attributed to waterfowl herbivory. Here we study the separate and combined effects of both belowground herbivory in spring by whooper swans and Bewick ‘s swans, and grazing in summer by waterfowl and fish on the community composition in a shallow Baltic estuary during one year. The macrophyte community was dominated by charophytes (mainly Chara aspera) with Potamogeton pectinatus and Najas marina present as subdominants. Other species were rare. Both spring and summer herbivory had no effect on total plant biomass. However, P. pectinatus was more abundant in plots that were closed to spring and summer herbivores. N. marina was more abundant in grazed plots, whereas Chara spp. biomass remained unaffected. Probably belowground propagules of both C. aspera and P. pectinatus were consumed by swans but since C. aspera bulbils were numerous it may have compensated for the losses. P. pectinatus may not have fully recovered from foraging on tubers and aboveground biomass. Our results are in line with other studies in Chara dominated lakes, which found no effect of grazing on summer aboveground Chara biomass, whereas several studies report strong effects of herbivory in lakes dominated by P. pectinatus.     

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.     

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.     

Effects of Cu,Pb and Zn on two Potamogeton species grown under field conditions

Two common macrophyte species, Potamogeton perfoliatus L. and Potamogeton pectinatus L. were grown for 12 weeks at shallow depths in sediments contaminated with 1250 or 2500 g Pb or Cu and/or Zn (gDW sediment)^-1. Control experiments were run at background levels of 4, 13, and 38 g Pb, Cu and Zn (gDW sediment)^-1, respectively. Effects of heavy metals on biomass production and metal uptake and distribution in plants are presented in relation to total amount and plant-available fraction of metals in the sediment.All three studied metals gave reduced biomass production, and the toxicity of the metals decreased in the order Zn>Cu>Pb. The root/shoot biomass ratio increased for P. pectinatus, but decreased for P. perfoliatus with metal treatment. The content of any single metal was higher in shoots than in roots of plants grown on sediments not contaminated with that specific metal, but addition of that metal increased the proportion in roots. The uptake by plants of any of the heavy metals increased with increased metal addition. The magnitude of the plant-available fraction of metals of untreated sediment was Zn>Cu>Pb, and increased in contaminated sediments. Addition of Cu decreased both the plant-available fraction and the total concentration of Zn in the sediment, while increased the uptake of Zn by the plants. The opposite was found for Cu when Zn was added. P. pectinatus accumulated about twice as much Cu as P. perfoliatus. On the other hand, the concentration of Pb was higher in P. perfoliatus than in P. pectinatus, and was negligible in P. pectinatus when cultivated in untreated sediments.     

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 CH₄. Microbial processes catalyzing nutrient recycling and CH₄ 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 CH₄ production, through a decrease in the activity of methanogenic Archaea and an increased rate of CH₄ 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 CH₄ 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 CH₄ cycling by means of both direct and indirect (i.e. plant-mediated) effects with potential consequences for CH₄ emission from wetland systems.Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Propagule type influences competition between two submersed aquatic macrophytes

Summary Glasshouse competition experiments with Hydrilla verticillata (L.f.) Royle indicate that plants grown from turions are weaker competitors than those grown from tubers, when compared to the widely distributed macrophyte, Potamogeton pectinatus L. These results support an earlier hypothesis about the importance of propagule size for predicting the outcome of plant competition (Grace 1985; Schaffer and Gadgil 1980). Results of outdoor growth experiments indicate that even though Hydrilla plants from turions are relatively weaker competitors, they are able to grow succesfully in an existing macrophyte bed composed of either, P. pectinatus or P. gramineus. During the early stages of Hydrilla invasion into an area of existing macrophytes, native plants may coexist with Hydrilla. However, once the abundance of Hydrilla tubers in the sediment increases, Hydrilla may displace existing plants.     

Overcompensation and grazing optimisation in a swan–pondweed system?

1. 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.     

Influence of migrant tundra swans (Cygnus columbianus) and Canada geese (Branta canadensis) on aquatic vegetation at Long Point, Lake Erie, Ontario

Numerous studies have shown that large, herbivorous waterfowl can reduce quantity of aquatic plants during the breeding or wintering season, but relatively few document herbivory effects at staging areas. This study was done to determine if feeding activities of tundra swans (Cygnus columbianus columbianus) and Canada geese (Branta canadensis) had a measurable additive influence on the amount of aquatic plants, primarily muskgrass (Chara vulgaris), wild celery (Vallisneria americana), and sago pondweed (Potamogeton pectinatus), removed during the fall migration period at Long Point, Lake Erie, Ontario. Exclosure experiments done in fall 1998 and 1999 showed that, as compared to ducks and abiotic factors, these two large herbivorous waterfowl did not have any additional impact on above or below ground biomass of those aquatic plants. As expected, however, there were substantial seasonal reductions in above-ground and below-ground biomass of aquatic plants in wetlands that were heavily used by all waterfowl. We suggest that differences in large- and small-scale habitat use, feeding activity, and food preferences between tundra swans and other smaller waterfowl as well as compensatory herbivory contributed to our main finding that large waterfowl did not increase fall reductions of Chara spp, V. Americana, and P. pectinatus biomass.     

Effects of mute swan grazing on a keystone macrophyte

1. This study describes the early summer foraging behaviour of mute swans (Cygnus olor) on the River Frome, a highly productive chalk stream in southern England in which Ranunculus penicillatus pseudofluitans is the dominant macrophyte. 2. A daily maximum of 41 ± 2.5 swans were present along the 1.1 km study reach during the study period (late May to the end of June). The river was the primary feeding habitat. Feeding activity on the river at dawn and dusk was much lower than during daylight, but we cannot rule out the possibility that swans fed during the hours of darkness. 3. The effects of herbivory on R. pseudofluitans biomass and morphology were quantified. Biomass was lower in grazed areas and swans grazed selectively on leaves in preference to stems. A lower proportion of stems from grazed areas possessed intact stem apices and flowering of the plant was reduced in grazed areas. 4. A model, based on the swans’ daily consumption, was used to predict the grazing pressure of swans on R. pseudofluitans. The model accurately predicted the number of bird days supported by the study site, only if grazing was assumed to severely reduce R. pseudofluitans growth. The proportion of the initial R. pseudofluitans biomass consumed by a fixed number of swans was predicted to be greater when the habitat area was smaller, initial R. pseudofluitans biomass was lower and R. pseudofluitans was of lower food value. 5. We concluded that the flux of N and P through the study reach was largely unaffected by swan activity. The quality of R. pseudofluitans mesohabitat (the plant as habitat for invertebrates and fish) was significantly reduced by grazing which also indirectly contributed to reduced roughness (Manning's n) and by inference water depth. Wetted habitat area for fish and invertebrates would also be lowered over the summer period as a consequence of the reduction in water depth. It was estimated that, while grazing, an individual swan may eat the same mass of invertebrates per day as a 300‐g trout. 6. There is a need to manage the conflict between mute swans and the keystone macrophyte, R. pseudofluitans, in chalk streams, and the modelling approach used here offers a potentially useful tool for this purpose.     

Diet of Mute Swans in Lower Great Lakes Coastal Marshes

Abstract: During the past 30 years, nonnative mute swan (Cygnus olor) populations have greatly increased, and continue to increase, in the eastern United States and within the lower Great Lakes (LGL) region. As a result, there is much concern regarding impacts of mute swan on native waterfowl, aquatic plants, and marsh habitats. There are presently only limited dietary data for mute swans in North America and none exist for birds in the LGL region. Thus, in 2001, 2002, and 2004 we collected 132 mute swans from LGL coastal marshes in Ontario, Canada, to determine dietary composition and to evaluate 1) seasonal and sex-related variation in adult diets and 2) age-related dietary differences. Adult diets did not differ among years, collection sites, or seasons, but female diets contained more pondweed spp. (Potamogeton spp.) and less slender naiad (Najas flexilis) and common waterweed (Elodea canadensis) than did diets of males. Adult males, adult females, and cygnets had similar diets during summer and autumn. Overall, mute swan diets mainly consisted of above-ground biomass of pondweed spp., muskgrass (Chara vulgaris), coontail (Ceratophyllum demersum), slender naiad, common waterweed, wild celery (Vallisneria americana), and wild rice (Zizania palustris); below-ground parts of wild celery, sago pondweed (Stuckenia pectinatus), and arrowhead spp. (Sagittaria spp.) were eaten infrequently. Comparison of our findings with those of other diet studies suggested considerable dietary overlap between mute swans and several other species of native waterfowl. Thus, we suggest that mute swans have potential to compete with native waterfowl and impact aquatic plants that are important waterfowl foods within LGL coastal marshes. Further, our results can be used to assess which aquatic plant species may be most impacted by foraging activities of mute swans at other important waterfowl stopover and wintering sites in North America. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):726–732; 2008)     

Genetic differentiation of submerged plant populations and taxa between habitats

Ceratophyllum spp., Callitriche spp., Zannichellia spp. and Potamogeton pectinatus L. are widespread submerged macrophyte species, often occurring at high abundance and forming an integral part of the vegetation of many types of shallow aquatic systems. Several species occur in both freshwater and brackish water habitats. Most have a mixed reproduction system and can reproduce sexually by seeds and propagate asexually by rhizomes, turions, root tubers or axillary tubers. It is hypothesized that sexual propagules are more important than vegetative fragments to ensure long-distance-dispersal, which in case of frequent bird or water flow-mediated dispersal should lead to lowered genetic differentiation. At a regional level, we used dominant ISSR markers in a multi-species approach and observed the largest clonal differentiation between brackish water and freshwater populations of the western European lowland (Belgium). Differentiation was pronounced at taxon level (e.g. Zannichellia), as a salinity gradient (P. pectinatus) or as a coastal-to-inland conductivity gradient (Callitriche obtusangula). These differences and trends suggested a very limited dispersal at regional level across both habitats and regions. To test the hypothesis whether vegetative reproduction and dispersal may have an important function in maintenance of the species at local scale, we investigated the microsatellite diversity and clonal distribution within and between populations of P. pectinatus from a single catchment, representing upstream forest ponds and downstream river sites along the Woluwe (Brussels, Belgium). Clonal diversity was low on average, however, with a higher number of multilocus genotypes in upstream forest ponds than in downstream river sites. A few but abundant clones were present along various stretches of the river indicating clonal spread and establishment over larger distances within the river. Clonal dispersal at a local scale was more pronounced in river than in pond habitats, indicating a higher relative importance of water flow than bird-mediated dispersal for establishment of P. pectinatus in river sites. Dispersal of seeds and establishment of seedlings were assumed more effective within ponds than in river habitats. Upstream forest ponds can be regarded as source populations and refuges of clonal diversity for recolonization of the more stressful downstream river habitat.     

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.     

Effects of grazing and inundation on pasture quality and seed production in a salt marsh

During a six-year period, changes in the composition of dominant plant species of importance to foraging birds in a salt marsh on the Swedish west coast were followed inside and outside exclosures to document effects of grazing on herbage quality and seed production. Since marshes provide an important habitat for foraging geese and ducks, it was of interest to determine how cattle grazing would affect herbage production inAgrostis stolonifera andPuccinellia maritima and seed and root-tuber production inScirpus maritimus. Measurements of cover and height in permanent plots revealed that a wetter weather type favouredAgrostis, probably through reduced salinity, at the expense ofPuccinellia, which was the most favoured food of both cattle and birds.Agrostis out-competedPuccinellia when grazing pressure was low. Seed production inScirpus maritimus was reduced by cattle grazing, particularly whenPhragmites australis formed part of the vegetation. In the absence of cattle grazing, both herbage- and seed producing plants were gradually reduced, andPhragmites increased. Since high herbage consumption and high seed production are mutually exclusive, grazing rotation in combination with mowing is suggested as a management strategy.     

基于卫星跟踪大天鹅春季迁徙时间及其影响因素

候鸟春季迁徙时间与其能否顺利完成迁徙过程,以及对繁殖地的成功选择和繁殖成效密切相关,通过对大天鹅越冬地和停歇地的春季迁徙时间选择原因及其影响因素进行分析,有助于深入理解候鸟春季迁徙时间策略和栖息地保护状况。2015年2月和12月,在河南三门峡湿地捕捉了60只越冬大天鹅并佩戴卫星跟踪器,获取了详细的大天鹅在越冬地和停歇地的春季迁徙时间等信息,并进一步分析了春季迁徙时间与气候因子的相关性。结果表明,大天鹅主要在夜间22：00-0：00和0：00-1：59迁离越冬地。大天鹅迁离越冬地的日期与温度呈显著性正相关,迁离时多选择顺风且风速较小的天气。大天鹅迁离越冬地后主要是在夜间飞行,而迁离停歇地后主要是在日间飞行。越冬地的温度越高,越有利于植物的生长,大天鹅可以快速地积累能量,提前开始春季迁徙。野外调查表明,内蒙古黄河中上游作为迁徙的重要停歇地,春季两岸捕鱼、农耕和放牧等为活动较多,因此大天鹅多选择在夜间觅食补充能量,在日间迁离。最后,针对黄河流域大天鹅栖息地的保护现状,提出了禁止经济开发项目、建立保护区和开展宣传教育等保护建议。     

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.     

Influence of livestock grazing on meadow pipit foraging behaviour in upland grassland

Changes in grazing management are believed to be responsible for declines in populations of birds breeding in grassland over the last decades. The relationships between grazing management regimes, vegetation structure and composition and the availability of invertebrate food resources to passerine birds remain poorly understood. In this study, we investigated the foraging site selection of meadow pipits (Anthus pratensis L.) breeding in high intensity sheep-grazed plots or low intensity mixed (i.e. sheep and cattle)-grazed plots. We sampled above-ground invertebrates, measured vegetation height and density and conducted a vegetation survey in areas where meadow pipits were observed to forage and areas that were randomly selected. Birds foraged in areas with a lower vegetation height and density and in areas containing a lower proportion of the dominant, tussock-forming grass species Molinia caerulea. They did not forage in areas with a total higher invertebrate biomass but at areas with preferred vegetation characteristics invertebrate biomass tended to be higher in foraging sites than random sites. The foraging distance of meadow pipits was higher in the intensively grazed plots. Our findings support the hypothesis that resource-independent factors such as food accessibility and forager mobility may determine patch selection and are of more importance as selection criteria than food abundance per se. Food accessibility seems to become an even more important selection criterion under high grazing intensity, where prey abundance and size decrease. In our upland grazing system, a low intensity, mixed grazing regime seems to provide a more suitable combination of sward height, plant diversity, structural heterogeneity and food supply for meadow pipit foraging activity compared to a more intensive grazing regime dominated by sheep.     

Mute swan (Cygnus olor) winter distribution and numerical trends over a 16-year period (1987/1988–2002/2003) in France

The numbers of mute swans (Cygnus olor) at 98 wetland locations in France were monitored monthly during the winter (December–February) for 16 years by a national network of observers as part of a broader national wildfowl monitoring scheme. Log-linear Poisson regressions with TRIM software were used to estimate missing counts and produce national numbers and indices. These corrected indices were in turn used to calculate an average annual rate of change and the associated confidence interval that subsequently enabled the computation of a cumulated (global) rate of change and associated confidence interval. The latter were interpreted to classify the numerical trends over the whole period. For the six wetlands with the largest numbers of mute swans, average numbers of swans were also compared between wetlands and months. General Linear Models were then used to test simultaneously for (1) differences between years and between wetlands for each month and (2) between years and between months separately for each of the six wetlands. The mute swan average annual rate of change was >7% for each month. The global rate of variation corresponded to a “strong increase” after TRIM analyses. This result strongly supports the necessity of maintaining both national and international monitoring schemes in order to be able to quickly detect major increases in swan numbers at specific sites, particularly where increasing numbers may result in a conflict with farmers or in inter-specific competition with other waterbirds and/or represent a possible sanitary/public health hazard due to the potential for swans to carry avian influenza viruses.     

Isozyme evidence for the parentage and multiple origins ofPotamogeton ×suecicus (P. pectinatus ×P. filiformis,Potamogetonaceae)

Evidence from isozyme analyses indicates thatPotamogeton ×suecicus is the hybrid betweenP. pectinatus andP. filiformis. The hybrid appears to have arisen on several occasions. The isozyme profiles of this hybrid from the Rivers Wharfe and Ure in Yorkshire, south of the present limit of distribution ofP. filiformis, suggest that each population is a single clone; these clones may be relics from the Weichselian glacial period. Populations of the putative hybrid from the Rivers Tweed and Till are notP. ×suecicus but probably haveP. vaginatus andP. pectinatus as parents. If so, this is a remarkable example of a pondweed hybrid persisting vegetatively in an area outside the distributional range of one of its parents.     

Coexistence and population genetic structure of the whooper swan <Emphasis Type="Italic">Cygnus cygnus</Emphasis> and mute swan <Emphasis Type="Italic">Cygnus olor</Emphasis> in Lithuania and Latvia

Two closely related swan species, the mute swan Cygnus olor and the whooper swan Cygnus cygnus, were formerly allopatric throughout their breeding ranges, but during the last decades a sympatric distribution has become characteristic of these species in the Baltic Sea region. The whooper swan has gradually replaced the mute swan in many suitable habitats in Lithuania and Latvia. Marked differences in the genetic population structure of both species may partially explain the dominance of the whooper swan, as genetic population divergence can be a major factor affecting inter-specific competition. A homogenous genetic population structure was defined for mute swans breeding in Lithuania, Latvia, Poland and Belarus. Breeding mute swans in this region are mostly of naturalised origin. A diverse population genetic structure characterizes whooper swans breeding in Lithuania and Latvia.