SUMMARY1. This synthesis examines 35 long-term (5-35 years, mean: 16 years) lake re-oligotrophication studies. It covers lakes ranging from shallow (mean depth <5 m and/or polymictic) to deep (mean depth up to 177 m), oligotrophic to hypertrophic (summer mean total phosphorus concentration from 7.5 to 3500 lg L )1 before loading reduction), subtropical to temperate (latitude: 28-65°), and lowland to upland (altitude: 0-481 m). Shallow northtemperate lakes were most abundant. 2. Reduction of external total phosphorus (TP) loading resulted in lower in-lake TP concentration, lower chlorophyll a (chl a) concentration and higher Secchi depth in most lakes. Internal loading delayed the recovery, but in most lakes a new equilibrium for TP was reached after 10-15 years, which was only marginally influenced by the hydraulic retention time of the lakes. With decreasing TP concentration, the concentration of soluble reactive phosphorus (SRP) also declined substantially. 3. Decreases (if any) in total nitrogen (TN) loading were lower than for TP in most lakes. As a result, the TN : TP ratio in lake water increased in 80% of the lakes. In lakes where the TN loading was reduced, the annual mean in-lake TN concentration responded rapidly. Concentrations largely followed predictions derived from an empirical model developed earlier for Danish lakes, which includes external TN loading, hydraulic retention time and mean depth as explanatory variables. 4. Phytoplankton clearly responded to reduced nutrient loading, mainly reflecting declining TP concentrations. Declines in phytoplankton biomass were accompanied by shifts in community structure. In deep lakes, chrysophytes and dinophytes assumed greater importance at the expense of cyanobacteria. Diatoms, cryptophytes and chrysophytes became more dominant in shallow lakes, while no significant change was seen for cyanobacteria. 5. The observed declines in phytoplankton biomass and chl a may have been further augmented by enhanced zooplankton grazing, as indicated by increases in the zooplankton : phytoplankton biomass ratio and declines in the chl a : TP ratio at a summer mean TP concentration of <100-150 lg L )1 . This effect was strongest in shallow lakes. This implies potentially higher rates of zooplankton grazing and may be ascribed to the observed large changes in fish community structure and biomass with decreasing TP contribution. In 82% of the lakes for which data on fish are available, fish biomass declined with TP. The percentage of piscivores increased in 80% of those lakes and often a shift occurred towards dominance by fish species characteristic of less eutrophic waters. 6. Data on macrophytes were available only for a small subsample of lakes. In several of those lakes, abundance, coverage, plant volume inhabited or depth distribution of submerged macrophytes increased during oligotrophication, but in others no changes were observed despite greater water clarity. 7. Recovery of lakes after nutrient loading reduction may be confounded by concomitant environmental cha...
Niche and neutral processes drive community assembly and metacommunity dynamics, but their relative importance might vary with the spatial scale. The contribution of niche processes is generally expected to increase with increasing spatial extent at a higher rate than that of neutral processes. However, the extent to what community composition is limited by dispersal (usually considered a neutral process) over increasing spatial scales might depend on the dispersal capacity of composing species. To investigate the mechanisms underlying the distribution and diversity of species known to have great powers of dispersal (hundreds of kilometres), we analysed the relative importance of niche processes and dispersal limitation in determining beta‐diversity patterns of aquatic plants and cladocerans over regional (up to 300 km) and continental (up to 3300 km) scales. Both taxonomic groups were surveyed in five different European regions and presented extremely high levels of beta‐diversity, both within and among regions. High beta‐diversity was primarily explained by species replacement (turnover) rather than differences in species richness (i.e. nestedness). Abiotic and biotic variables were the main drivers of community composition. Within some regions, small‐scale connectivity and the spatial configuration of sampled communities explained a significant, though smaller, fraction of compositional variation, particularly for aquatic plants. At continental scale (among regions), a significant fraction of compositional variation was explained by a combination of spatial effects (exclusive contribution of regions) and regionally‐structured environmental variables. Our results suggest that, although dispersal limitation might affect species composition in some regions, aquatic plant and cladoceran communities are not generally limited by dispersal at the regional scale (up to 300 km). Species sorting mediated by environmental variation might explain the high species turnover of aquatic plants and cladocerans at regional scale, while biogeographic processes enhanced by dispersal limitation among regions might determine the composition of regional biotas.
The study of species complexes is of particular interest to understand how evolutionary young species maintain genomic integrity. The Daphnia pulex complex has been intensively studied as it includes species that dominate freshwater environments in the Northern hemisphere and as it is the sole North American complex that shows transitions to obligate parthenogenesis. Past studies using mitochondrial markers have revealed the presence of 10 distinct lineages in the complex. This study is the first to examine genetic relationships among seven species of the complex at nuclear markers (nine microsatellite loci and one protein-coding gene). Clones belonging to the seven species of the Daphnia pulex complex were characterized at the mitochondrial NADH dehydrogenase (ND5) gene and at the Lactate dehydrogenase (LDH) locus. K-means, principal coordinate analyses and phylogenetic network analyses on the microsatellite data all separated European D. pulicaria, D. tenebrosa, North American D. pulex, D. pulicaria and their hybrids into distinct clusters. The hybrid cluster was composed of diploid and polyploid hybrids with D. pulex mitochondria and some clones with D. pulicaria mitochondria. By contrast, the phylogeny of the D. pulex complex using Rab4 was not well resolved but still showed clusters consisting mostly of D. pulex alleles and others of D. pulicaria alleles. Incomplete lineage sorting and hybridization may obscure genetic relationships at this locus. This study shows that hybridization and introgression have played an important role in the evolution of this complex.
The competitive ability of hybrids, compared with their parental taxa, can cover a wide fitness range from poor to superior. For example communities of the Daphnia galeata-hyalina-cucullata species complex often show hybrid dominance. We tested whether taxa composition of 43 European lakes inhabited by this species complex can be explained by habitat characteristics (e.g. size descriptors, trophy level) or geography. We found that D. galeata occurs more frequently south of the Alps, whereas D. hyalina and D. cucullata are found more in the north. Lakes with D. galeata dominance had higher temperatures whereas D. hyalina dominance could be attributed to low phosphorus loads. The dominance of F 1 -hybrids, however, was not explainable with current environmental variables. In a subset of 28 lakes, we studied the impact of eutrophication history on F 1 -hybrid success. Lakes with the highest trophic state in the past tended to be dominated by F 1 -hybrids. Our data demonstrate that human-mediated habitat disturbance (eutrophication) has facilitated hybrid success and altered the Daphnia taxon composition across lakes. At the same time, specific habitat conditions might provide a refuge from hybridization for native genotypes.
Summary 1. Species assemblages of diatoms, rotifers, chydorids, planktonic crustaceans and chironomids were studied in 235 alpine lakes in the Alps, Pyrenees, Tatras (Western Carpathians), Retezat (Southern Carpathians) and Rila Mountains (Balkans). 2. For all taxonomic groups we found a hierarchical structure in the community assemblage using distinct scales of lake clustering (number of k‐means groups) based on species composition similarity (Hellinger distance). We determined the optimal partition in assemblage types (i.e. number of lake clusters) for each taxonomic group by maximising the sum of the taxon indicative value (IndVal) and performed discriminant analyses, using environmental variables not conditioned by geographical patterns. Relevant environmental variables differed among and within taxonomic groups. Therefore the assemblages respond to a complex environmental mosaic, with the exception of diatom assemblages, which followed an acid–base gradient. 3. The significant environmental variables could be grouped into four general factors: lake size, tropho‐dynamic status, acid–base balance and ice‐cover duration (i.e., altitudinal gradient). Lake size was significant for the highest number of assemblage types; however, the most significant factor differed among taxonomic groups: acid–base balance for diatoms, lake size for rotifers, ice‐cover duration for chydorids and planktonic crustaceans and tropho‐dynamic status for chironomids. No single environmental typology accounted for the assemblage structure of all taxonomic groups. 4. However, defining ecological thresholds as values within environmental gradients at which the rate of change in assemblages is accelerated relative to points distant from that threshold, we were able to find specific threshold values for each of the four main general environmental factors identified, which were relevant across several taxonomic groups: 3 ha for lake area; 0.6 mg L−1 for dissolved organic carbon; 190 days for ice‐cover duration and 200 μeq L−1 for acid neutralising capacity. Above and below these values ecosystem organisation change substantially. They have direct applications in establishing lake typologies for environmental quality and biodiversity conservation programmes, and in improving predictions about global change impacts.
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