Retention and supply of zebra mussel larvae in a large river system: importance of an upstream lake

1. Persistence of zebra mussel populations in river systems probably depends upon the presence of upriver sites capable of hosting self‐recruiting adult populations that act as sources of larvae. In this paper we examine the importance of Lake Pepin, a natural riverine lake in the Upper Mississippi River, as an upriver source of larvae to the downstream populations of zebra mussels. 2. Field studies and modelling suggest that Lake Pepin plays a major role in maintaining zebra mussel populations in the Upper Mississippi River. Long water residence times in Lake Pepin allow for self‐recruitment under the right hydraulic conditions. Larval abundance was low to absent upstream of the lake but increased dramatically downriver in all 3 years of the study. Travel time estimates in the Upper Mississippi River show that newly fertilised larvae drifting out of Lake Pepin can contribute substantially to the major downstream peak in larval abundance. In contrast, backwater and other off‐channel sites are unlikely to drive main‐channel abundance patterns. Larval abundances in off‐channel sites were less than or equal to those in the main channel. 3. A key factor in assessing the importance of Lake Pepin as a source population was the abundance of early stage, unshelled larvae. Studies that consider only abundances of older shelled stages (visible by cross polarised lighting) may yield misleading results. Results of this study suggest that efforts to control zebra mussels in the Upper Mississippi River should focus on controlling adult populations within Lake Pepin and reducing or eliminating larvae exiting the lake.     

Trinucleotide microsatellite loci for the zebra mussel Dreissena polymorpha,an invasive species in Europe and North America

The high mutation rate at microsatellite loci can supply important demographic information on founder events and range expansion in an invasive species such as the zebra mussel Dreissena polymorpha, following its initial introduction. In order to facilitate studies into the colonization patterns of this species in new habitats in Europe and North America, five trinucleotide microsatellite loci were isolated from a partial DNA library. Allelic diversity at all described loci was high, ranging from 20 to 35 alleles per locus. Homologous loci were not amplified in a second related invasive species, Dreisenna bugensis, using the primers developed here.     

Genetic variability and phylogeographical patterns of a nonindigenous species invasion: a comparison of exotic vs. native zebra and quagga mussel populations

There have been few investigations of the number of founding sources and amount of genetic variability that lead to a successful nonindigenous species invasion, although genetic diversity is believed to play a central role. In the present study, population genetic structure, diversity and divergence patterns were analysed for the zebra mussel Dreissena polymorpha [n=280 samples and 63 putative randomly amplified polymorphic DNA (RAPDs) gene loci] and the quagga mussel D. bugensis (n=136 and 52 loci) from 10 nonindigenous North American and six Eurasian sampling sites, representing their present‐day ranges. Results showed that exotic populations of zebra and quagga mussels had surprisingly high genetic variability, similar to those in the Eurasian populations, suggesting large numbers of founding individuals and consistent with the hypothesis of multiple colonizations. Patterns of genetic relationships indicate that the North American populations of D. polymorpha likely were founded by multiple source populations from north‐western and northcentral Europe, but not from southcentral or eastern Europe. Sampling areas within North America also were significantly divergent, having levels of gene flow and migration about twice those separating long‐established Eurasian populations. Samples of D. bugensis in Lakes Erie and Ontario were significantly different, with the former being more closely related to a native population from the Dnieper River, Ukraine. No evidence for a founder effect was discerned for either species.     

Amplified fragment length polymorphism (AFLP) analysis of the genetic structure of the zebra mussel, Dreissena polymorpha, in the Mississippi River

1. We predicted that zebra mussel, Dreissena polymorpha (Pallas), genetic structure in the Mississippi River would follow a model of invasive species genetics, which predicts low genetic structure among populations of recently established species. This prediction was upheld in our previous genetic study using allozymes, however, one locus yielded anomalous results. 2. We employed amplified fragment length polymorphism (AFLP) analysis as a neutral marker to assess the amount of genetic structure within and among populations, and as a test of expected population structure from both invasion genetic theory, and the results from our previous study. 3. There was greater spatial differentiation, as measured by F_st, observed using AFLP's than for allozymes (P < 0.001). There was no evidence that AFLP variation conformed to an isolation by distance model, and genetic relationships of populations, as measured by AFLP markers, were not similar to those detected in our allozyme survey. 4. The lack of concordance between these two genetic marker systems probably reflects their differential responses to drift, migration, and selection occurring during this rapid invasion. Strong population structure is counter to predictions that populations of invasive species will not be differentiated, as with observations based on allozyme markers. Therefore, newly established species may require genetic surveys using multiple marker systems to evaluate population structure.     

Invasive zebra mussel (Dreissena polymorpha) threatens an exceptionally large population of the depressed river mussel (Pseudanodonta complanata) in a postglacial lake

1. Freshwater mussels are in decline worldwide, with the depressed river mussel Pseudanodonta complanata being one of the rarest and most endangered species in Europe. Invasive mussels are suspected to be an important factor of decline, but there is little information on their interaction with native species.
2. This study analyzed densities, depth distribution, and individual sizes and weights in one of the largest known populations of P. complanata in Europe in relation to the co‐occurring invasive zebra mussel Dreissena polymorpha and other mussel species, using a systematic transect analysis.
3. Pseudanodonta complanata was the dominant unionid species in Lake Siecino reaching densities of up to 26 ind/m², with half of the specimens found at a water depth of 2.0–4.0 m. Densities were highest on sandy substrates in areas of underwater currents. In contrast, 67% of native Unio tumidus were found at depths < 1 m, indicating different habitat preference.
4. In the study area, 91% of P. complanata, 92% of U. tumidus, and all Anodonta individuals were fouled by D. polymorpha. The dreissenid:unionid mass ratio (mean ± SD; maximum) was 0.43 ± 0.56; 4.22 and 0.86 ± 1.87; 8.76 in P. complanata and U. tumidus, respectively. Pseudanodonta complanata fouled with D. polymorpha were impaired in their anchoring capability and had shell deformations potentially affecting shell closing and filtration activity. Fouling intensity was negatively correlated with unionid density, potentially leading to accelerated population declines.
5. The observed adverse effects of invasive zebra mussels on the depressed river mussel and the difficulties in eradicating established populations of invasive mussels suggest that D. polymorpha should be considered a serious threat to P. complanata. Therefore, the further spread of zebra mussels into habitats with native unionids needs to be avoided by all means.

     

The pericardial glands of the zebra mussel: Ultrastructure and implication in lead detoxication process

Summary— Ultrastructural and microanalytical investigations of the pericardial gland of the freshwater mussel, Dreissena polymorpha have been performed to investigate the possible functional role of this organ in the detoxication process of lead. The cell-type of this organ exhibits the feature characteristics of podocytes, ie the typical pedicel-basal lamina complex and the well developed lysosomalvacuolar system. X-ray microanalysis demonstrated that large electron-dense granules referred to as lysosomes are the main target organelles in these cells to accumulate and sequestrate lead where the metal was associated with phosphorus and sulphur. Consequently, the pericardial gland plays an important role in the detoxication process and allows the organism to tolerate high lead concentration without suffering severe cell injury.     

Exploration of structure-antifouling relationships of capsaicin-like compounds that inhibit zebra mussel (Dreissena polymorpha) macrofouling

Abstract

Macrofouling of aquatic man-made structures by zebra mussels (Dreissena polymorpha) poses significant economic burdens on commercial freshwater shipping and facilities utilising raw water. The negative environmental impact of some current antifouling technologies has limited their use and prompted investigation of non-organometallic and non-oxidising antifoulants as possible environment-friendly alternatives. The plant-derived natural product capsaicin and 18 other compounds with one or more capsaicin-like structural features were tested for their potential to inhibit zebra mussel byssal attachment at a single high concentration of 30 μM. Of these, three compounds displaying the highest levels of attachment inhibition where selected for further concentration-response testing. This testing revealed that capsaicin (8-methyl-N-vanillyl-trans-6-nonenamide), N-vanillylnonanamide, and N-benzoylmonoethanolamine benzoate all inhibited byssal attachment with potency values (EC₅₀) in the micromolar range. None of these compounds were lethal to adult specimens of the water flea, Daphnia magna, at concentrations that inhibited mussel byssal attachment.     

The filtration rate of the zebra mussel Dreissena polymorpha used for water quality assessment in Dutch rivers

The effects of contaminated river water on the filtration rate of zebra mussels from a clean reference site were studied. After a 48-h exposure period to filtered water from the rivers Rhine, Meuse and Amstel (The Netherlands), the filtration rate was measured. It was demonstrated that water from contaminated locations inhibited the filtration rate. Inhibition was higher during low water levels in the rivers Rhine and Meuse than during high water levels, suggesting that contaminants are diluted during high water levels. It is concluded that the shortterm filtration assay with D. polymorpha can be used for assessing water quality.     

The zebra mussel (Dreissena polymorpha), a new pest in North America: reproductive mechanisms as possible targets of control strategies

Summary

The zebra mussel (Dreissena polymorpha) has spread rapidly in temperate fresh waters of North America since its introduction into the Great Lakes in 1985 or 1986. It attaches to hard substrates, forming layers, occluding water intakes, encrusting and killing native mussels, filtering algae in competition with other planktivores, and possibly interfering with fish spawning. It reproduces prolifically, suggesting that an approach to its control may be by controlling its reproduction. Previous literature suggests that spawning in bivalves is regulated by both environmental and internal chemical cues. A suggested sequence is that phytoplankton chemicals initially trigger spawning; chemicals associated with gametes provide a species-specific pheromonal positive feedback for spawning; and the response to environmental chemicals is mediated internally by serotonin (5-HT). The role of 5-HT in zebra mussels is under investigation. Both males and females can be induced to spawn by either injection or external application of 5-HT. The response can also be activated by 8-hydroxy-2-(di-n-propylamino)-tetralin, an agonist at 5-HT_1A receptors. HPLC analysis has detected 5-HT as the major biogenic amine in both male and female gonads. 5-HT immunocytochemistry demonstrates nerves containing serotonergic fibers innervating gonads of both males and females, with prominent varicosities surrounding the follicles in both sexes. A role of 5-HT in mediating spawning responses in zebra mussels is thus strongly supported. These studies have shown that reproductive behavior of zebra mussels can be modified by outside chemicals, a property that may be exploited for purposes of control.     

Indigenous microflora and opportunistic pathogens of the freshwater zebra mussel, Dreissena polymorpha

Freshwater fouling invertebrate zebra mussels (Dreissena polymorpha) harbor a diverse population of microorganisms in the Great Lakes of North America. Among the indigenous microorganisms, selective species are opportunistic pathogens to zebra mussels. Pathogenicity to zebra mussels by opportunistic bacteria isolated from the mussels was investigated in this study. Among the more than 30 bacteria isolated from temperature-stressed mussels, Aeromonas media, A. veronii, A. salmonicida subsp. salmonicida, and Shewanella putrefaciens are virulent pathogens to juvenile zebra mussels. Inoculation of a bacterial concentration of A. media, A. salmonicida subsp. salmonicida and S. putrefaciens at 10⁷ cells per zebra mussel resulted in 100% mortality within 5 days, and only 64.9% for A. veronii. In contrast, mortality was less than 12.3% following inoculation of a sterile phosphate buffer solution as a control. In addition, mortality was dependent on the size of the pathogen population used in inoculation and the incubation temperature, indicating the close relationship between the bacterial population and subsequent death. On the mussel tissue, a dense microbial population was evident from the moribund mussels viewed with Scanning Electron Microscope (SEM). Opportunistic bacteria invaded and destroyed the D. polymorpha tissue after 7 days of incubation when the bacterial inoculation was larger than 10⁵ per zebra mussel. Our results suggest that mussels are reservoirs of opportunistic pathogenic microorganisms to aquatic organisms and humans and a better understanding of the microbial ecology of the mussels will provide insights to the possible health hazards from these microorganisms.     

Specific amplification of the 18S rRNA gene as a method to detect zebra mussel (Dreissena polymorpha) larvae in plankton samples

An important issue in the management of zebra mussel (Dreissena polymorpha) populations is early, rapid, and accurate detection of the planktonic larvae (veliger) of the zebra mussel. The goal of this study was to explore the feasibility of developing a molecular approach for the detection of zebra mussel larvae in diverse environments. In this study a Dreissena polymorpha-specific 18S ribosomal RNA gene targeted oligonucleotide primer (ZEB-715a) and Polymerase Chain Reaction (PCR) assay was developed and compared with cross-polarized microscopy as a means to detect zebra mussel veligers in plankton samples. The design of the zebra mussel-specific primer was facilitated by sequencing nearly the complete 18S rRNA gene from the zebra mussel and three other closely related freshwater Veneroids including the quagga mussel (D. bugensis), the dark false mussel (Mytilopsis leucophaeata), and the Asian freshwater clam (Corbicula fluminea). The specificity of the primer for the zebra mussel was empirically tested by using the primer as a direct probe in a blot hybridization format. A single veliger in a plankton sample could be detected by PCR using this approach. Veliger detection sensitivity using the PCR approach was estimated to be over 300 times more sensitive than cross-polarized light microscopy based techniques. Cross-polarized light microscopy and the PCR technique were used to identify the presence of zebra mussel larvae in plankton samples that were collected from a variety of natural and industrial water sources. Detection results (presence or absence) were generally consistent between the two methods. Although additional studies will be required before routine application of molecular based veliger detection technology is available, a long-term goal of this work is the application of molecular technology to the development of a field device for the routine detection and quantification of zebra mussel veligers.     

A sieving method for rapid determination of size-frequency distribution of small gastropods. Example of the mud snail Hydrobia ventrosa (Gastropoda: Prosobranchia)

A sieving system using sieves of predefined mesh size has been developed for rapid size–frequency distribution analysis as a basis for the assessment of the secondary production of the hydrobiid species Hydrobia ventrosa Montagu of Lake Ichkeul in northern Tunisia (Mediterranean). The relationship between shell length (SL) and shell width (SW) SL = e^0.5005 SW^1.4306 calculated by a geometric mean regression is very highly significant allowing the application of sieves in mesh intervals of 0.1 mm for the determination of size–frequency distribution. The mean difference between sieved and measured size distributions, for different distribution patterns, only slightly exceeds 1%. The regression between shell length and ash-free dry mass (AFDM) fits an exponential model AFDM = e^{(-2.6591 + 0.9194 SL)}. Mean difference between estimated and observed biomass is in the 10% range. The elementary composition (C, H and N) of the ash-free dry mass is size-dependent. As a result, the proposed sieving method reduces by at least 20 times the data acquisition time (extraction, sorting and size-measuring), avoiding subsampling without significant loss of accuracy for secondary production studies of small gastropods.     

Microsatellite loci for dreissenid mussels (Mollusca: Bivalvia: Dreissenidae) and relatives: markers for assessing exotic and native populations

We developed and tested 14 new polymorphic microsatellite loci for dreissenid mussels, including the two species that have invaded many freshwater habitats in Eurasia and North America, where they cause serious industrial fouling damage and ecological alterations. These new loci will aid our understanding of their genetic patterns in invasive populations as well as throughout their native Ponto-Caspian distributions. Eight new loci for the zebra mussel Dreissena polymorpha polymorpha and six for the quagga mussel D. rostriformis bugensis were compared with new results from six previously published loci to generate a robust molecular toolkit for dreissenid mussels and their relatives. Taxa tested include D. p. polymorpha, D. r. bugensis, D. r. grimmi, D. presbensis, the 'living fossil'Congeria kusceri, and the dark false mussel Mytilopsis leucophaeata (the latter also is invasive). Overall, most of the 24 zebra mussel (N = 583) and 13 quagga mussel (N = 269) population samples conformed to Hardy-Weinberg equilibrium expectations for the new loci following sequential Bonferroni correction. The 11 loci (eight new, three previously published) evaluated for D. p. polymorpha averaged 35.1 alleles and 0.72 mean observed heterozygosity per locus, and 25.3 and 0.75 for the nine loci (six new, three previously published) developed for D. r. bugensis. All but three of these loci successfully amplified the other species of Dreissena, and all but one also amplified Congeria and Mytilopsis. All species and populations tested were significantly divergent using the microsatellite data, with neighbour-joining trees reflecting their evolutionary relationships; our results reveal broad utility for resolving their biogeographic, evolutionary, population and ecological patterns.     

Long-term variability and density dependence in Hudson River Dreissena populations

1. We used a 27-year record of Dreissena populations in the freshwater tidal Hudson River to describe interannual variation in population density, body size, and body condition; estimate long-term variation in recruitment, survivorship, and shell growth; and assess possible controls on the populations.
2. Dreissena populations in the Hudson have been highly variable, with interannual ranges of c. 100-fold in abundance and biomass, and 7-fold in mean body mass. This large interannual variation arises from both long-term trends and 2–5-year cycles.
3. Long-term trends include the 2008 appearance of the quagga mussel (Dreissena rostriformis), which still forms a small part (<10%) of the dreissenid community, and a decline in zebra mussel body size. The decline in body size was caused by a long-term decline in adult survivorship rather than a decline in rates of shell growth. We could detect no long-term trends in adult abundance or spread of Dreissena onto soft sediments in the Hudson.
4. We observed persistent, strong cycles in adult abundance and body size. These were driven by the appearance and decay of eight dominant year classes over the 27 years of our study, and were a result of temporal variation in recruitment rather than temporal variation in survivorship. The observed strongly irregular recruitment appears to arise from strong adult–larval interactions, and is consistent with previous simulation model results showing that interactions between adults and larvae can drive persistent cycling.
5. We found evidence that negative density dependence affects recruitment, somatic growth, and body condition of Dreissena in the Hudson. Warm summers may also cause high adult mortality.
6. We put our results into the context of a conceptual model of Dreissena population dynamics, and argue that neither the dynamics nor the controls of populations of these important invaders is known satisfactorily.

     

Mechanisms and impact of differential fouling of the zebra mussel Dreissena polymorpha on different unionid bivalves

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Diversity,habitats and size-frequency distribution of the gastropod genus Conus at Dahab in the Gulf of Aqaba,Northern Red Sea

Topographically complex subtidal reefs of the Indo-West Pacific region are characterised by a high species richness of cone snails of the genus Conus (up to 36 on some reefs) but low population densities (≤1 individual/m²), whereas Conus assemblages on reef flats usually support fewer species (5–9) and high population densities (up to 5.2 individuals/m²). Subtidal sand areas are known to be least species-rich (1–6 species). Although the diversity of this predatory gastropod genus has been described previously from different areas of the Indo-Pacific, little ecological information is available on Conus in the Northern Red Sea. Therefore, data from five habitat types were obtained along 73 line-transects (245?m²⁾, which yielded ecological data for a total of 175 individuals of 9 species. In accordance with former findings, our results demonstrate that the reef flat was the habitat with the highest observed population densities (6.15 individuals/m²) but low species diversity (H’ = 0.9; 5 species); subtidal reefs, in contrast, were characterised by low densities (0.13 individuals/m²) and a relatively high species diversity (H ‘= 1.5; 6 species). This suggests that Conus diversity and species richness in the Northern Red Sea around Dahab is lower than in other parts of the Indo-West Pacific region. Furthermore, hardand soft substrata were dominated by different Conus species in accordance to the distribution of favourable microhabitat patches, the degree of physical stress and the availability of refuges and prey organisms. The fact that these Conus were predominantly small-bodied vermivores (size range: 6–85?mm; mean shell size: 15?mm; SD = 9?mm) suggests that this size class possesses an advantage over molluscivores and piscivores. Except for subtidal reefs, which showed a highly variable species composition, the studied habitat types around Dahab were characterised by distinct assemblages of Conus.     

Biotic resistance on the increase: native predators structure invasive zebra mussel populations

1. Abundant native predators, parasites and pathogens that switch to consuming a hyper‐successful exotic species may be able to control the invasive population. Native predators may, however, need time to adapt to feed effectively on an exotic resource. In this case, mortality on an exotic population from native predators could increase over time even without a numerical increase in the predator population. 2. We measured mortality of zebra mussels (Dreissena polymorpha) in the Hudson River both in controls open to predation and in exclosures that excluded large predators to estimate mortality of zebra mussels from large predators and other causes. 3. We found that predation by the blue crab (Callinectes sapidus), and perhaps other predators, causes high mortality on zebra mussels in the Hudson River estuary. This predation apparently led to increased mortality and altered population structure in the invader over time. 4. Long‐term data from the Hudson River suggest that components of the invaded ecosystem, like rotifers, are recovering through predator‐caused release from zebra mussel grazing. Increased mortality on hyper‐successful exotic populations over time may be a common phenomenon with both ecological and management implications.     

The worldwide transfer of ants: geographical distribution and ecological invasions

Aim This is the first comprehensive account of the biogeography of ants transferred and at least temporarily established outside their native habitat. Location Using museum and literature records, I established the distributions of transferred ant species. Methods I used taxonomic and functional groups to assess how geographical spread as a transferred species is affected by taxonomy and life history. Results 147 ant species in forty-nine genera have been recorded outside of their native habitat. The proportion of transferred ants is similar to the number of genera and species in each subfamily. The species-rich subfamily Myrmicinae contains nearly 50% of all transferred species, while many of the species-poor subfamilies have absolutely no transferred species. A disproportionate high number of transferred ants originate from the Neotropical and Oriental biogeographic regions. The Pacific Islands are the recipients of the most transferred ant species. Most transferred ants belong to the CRYPTIC, OPPORTUNIST, and GENERALIZED MYRMICINE functional groups, while there are no recorded transfers of army ants or leaf-cutting ants. Both invasive and human commensal ‘tramp’ ant species are nonrandom subsets of transferred ants. Main conclusions ‘Tramp’ species and invasive species tend to have widespread geographical distributions, and share life history characteristics including queen number, nest structure, and foraging behaviour. Combining observations of functional groups and biogeography may lead to a better understanding of the factors contributing to the spread of transferred species.     

Molecular ecology of zebra mussel invasions

The invasion of the zebra mussel, Dreissena polymorpha, into North American waters has resulted in profound ecological disturbances and large monetary losses. This study examined the invasion history and patterns of genetic diversity among endemic and invading populations of zebra mussels using DNA sequences from the mitochondrial cytochrome oxidase I (COI) gene. Patterns of haplotype frequency indicate that all invasive populations of zebra mussels from North America and Europe originated from the Ponto-Caspian Sea region. The distribution of haplotypes was consistent with invasive populations arising from the Black Sea drainage, but could not exclude the possibility of an origin from the Caspian Sea drainage. Similar haplotype frequencies among North American populations of D. polymorpha suggest colonization by a single founding population. There was no evidence of invasive populations arising from tectonic lakes in Turkey, while lakes in Greece and Macedonia contained only Dreissena stankovici. Populations in Turkey might be members of a sibling species complex of D. polymorpha. Ponto-Caspian derived populations of D. polymorpha (theta = 0.0011) and Dreissena bugensis (one haplotype) exhibited low levels of genetic diversity at the COI gene, perhaps as a result of repeated population bottlenecks. In contrast, geographically isolated tectonic lake populations exhibited relatively high levels of genetic diversity (theta = 0.0032 to 0.0134). It is possible that the fluctuating environment of the Ponto-Caspian basin facilitated the colonizing habit of invasive populations of D. polymorpha and D. bugensis. Our findings were concordant with the general trend of destructive freshwater invaders in the Great Lakes arising from the Ponto-Caspian Sea basin.     

Seasonal effects of zebra mussels on littoral nitrogen transformation rates in Gull Lake, Michigan, U.S.A.

• 1 Zebra mussels (Dreissena polymorpha) are successful colonisers of lake littoral habitats and they interact strongly with littoral benthos. Previous research suggests that localised areas colonised by zebra mussels may be hotspots of nitrogen (N) cycling.
• 2 The effects of zebra mussels on nitrification and denitrification rates were examined approximately every other month for 1 year in Gull Lake, Michigan, U.S.A. Littoral sediment was collected from an area free of zebra mussels and distributed into shallow trays; rocks colonised with zebra mussels were placed in half of the trays, while uncolonised rocks were placed in the remaining trays. After an incubation period of 6–8 weeks in the lake, sediment and zebra mussels were collected from the trays, replaced with new sediment and zebra mussels, and placed in the lake for the next interval. In the laboratory, sediment nitrification and denitrification rates were measured for each tray.
• 3 Sediment nitrification rates did not increase in the presence of zebra mussels; instead nitrification rates were sensitive to changes in water temperature and increased with increasing exchangeable sediment ammonium. In contrast, denitrification rates increased in sediment trays with zebra mussels in the winter when nitrate (NO₃⁻) availability was high and when Chara did not grow in the trays.
• 4 Sediment denitrification was NO₃⁻‐limited in all seasons, regardless of zebra mussel treatment. However, sediment in the presence of zebra mussels responded less to NO₃⁻ addition, suggesting that NO₃⁻ limitation of denitrification can be reduced by zebra mussel activity. Zebra mussels have a seasonally variable impact on sediment denitrification rates, and this may translate into altered seasonal patterns of N cycling in localised areas of lakes where they are particularly abundant.