Biological invasion by a benthivorous fish reduced the cover and species richness of aquatic plants in most lakes of a large North American ecoregion

Biological invasions are projected to be the main driver of biodiversity and ecosystem function loss in lakes in the 21st century. However, the extent of these future losses is difficult to quantify because most invasions are recent and confounded by other stressors. In this study, we quantified the outcome of a century‐old invasion, the introduction of common carp to North America, to illustrate potential consequences of introducing non‐native ecosystem engineers to lakes worldwide. We used the decline in aquatic plant richness and cover as an index of ecological impact across three ecoregions: Great Plains, Eastern Temperate Forests and Northern Forests. Using whole‐lake manipulations, we demonstrated that both submersed plant cover and richness declined exponentially as carp biomass increased such that plant cover was reduced to <10% and species richness was halved in lakes in which carp biomass exceeded 190 kg ha^?1. Using catch rates amassed from 2000+ lakes, we showed that carp exceeded this biomass level in 70.6% of Great Plains lakes and 23.3% of Eastern Temperate Forests lakes, but 0% of Northern Forests lakes. Using model selection analysis, we showed that carp was a key driver of plant species richness along with Secchi depth, lake area and human development of lake watersheds. Model parameters showed that carp reduced species richness to a similar degree across lakes of various Secchi depths and surface areas. In regions dominated by carp (e.g., Great Plains), carp had a stronger impact on plant richness than human watershed development. Overall, our analysis shows that the introduction of common carp played a key role in driving a severe reduction in plant cover and richness in a majority of Great Plains lakes and a large portion of Eastern Temperate Forests lakes in North America.     

Variation in vegetation and seed banks of freshwater lakes with contrasting intensity of aquaculture along the Yangtze River, China

We examined species composition and richness in the seed banks and established vegetation in lakes along the Yangtze River with two contrasting aquaculture types: pen-culture lakes, characterized by pen-culture of commercial fish and crab, and lake-culture lake with more intensive stocking of the entire lake. The mean live biomass and species richness of submerged vegetation was significantly reduced or absent in lake-culture lakes (0-159 g m⁻² and 0-0.3 species per sample, respectively), compared to those with pen-culture lakes (1552-2971 g m⁻² and 1.5-3.2 species per sample, respectively) in all three study years. Also mean seedling density and species richness of seed banks were significantly lower in these lake-culture lakes (133 m⁻² vs. 265 m⁻² and 0.6 species vs. 0.9 species per sample, respectively). These results suggest that intensive aquaculture in these lakes has had serious negative effects on submerged vegetation and the associated seed banks. Vegetation history was partly reflected by distribution patterns of seed banks across sediment depth strata. A principal component analysis produced a very clear separation of lakes from pen-culture and lake-culture on the basis of their species composition. Moreover, the principal component analysis also indicated that the variation in the soil seed bank corresponded poorly with vegetation data. This probably reflects species-specific strategies for seed production.     

Factors controlling vegetation structure in peatland lakes—Two conceptual models of plant zonation

Two conceptual models of plant zonation in peatland lakes are given. The first represents vegetation on slightly sloping substrate (N < 0.2) in shallow and relatively large lakes. The vegetation is not diverse (H′ = 0.0 ± 0.01). The frequency and biomass of the dominant (Sphagnum denticulatum) correlate positively with lake size, and negatively with depth and substrate slope. They are also correlated with water transparency and water color (r = −0.53), concentrations of total organic carbon (r = −0.43), Ca²⁺ (r = 0.40) and humic acids (r = −0.46), and redox potential (r = 0.44). The second model represents vegetation on steep peat walls (N > 0.3) in deep, usually small lakes. Plants occur only on the upper part of the peat wall or form a multispecies curtain hanging from the lip of peat at the top. Species diversity in this scenario is higher (H′ = 0.18 ± 0.17). The curtains usually are composed of mosses such as Warnstorfia exannulata, S. cuspidatum and S. riparium, and vascular plants are rare. The frequency and biomass of bryophytes in this type of structure are related to substrate slope (r = 0.56), lake depth (r = 0.56), Ca²⁺ concentration (r = −0.69) and water color (r = −0.51). In both models, plant biomass is correlated with temperature (r = −0.78), irradiance (r = −0.64) and water oxygenation (r = −0.54).     

Homogenization of fish assemblages in different lake depth strata at local and regional scales

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Relationships between lake macrophyte cover and lake and landscape features

We examined the ability of lake and landscape features to predict a variety of macrophyte cover metrics using 54 north temperate lakes. We quantified submersed cover, emergent cover, floating leaf cover, Eurasian watermilfoil cover and total macrophyte cover. Measured lake features included lake physio-chemical and morphometric variables and landscape features included hydrologic, catchment and land use/cover variables. Univariate regression analyses demonstrated that these macrophyte cover metrics are predicted by a wide range of predictor variables, most commonly by: Secchi disk depth, maximum or mean depth, catchment morphometry, road density and the proportion of urban or agricultural land use/cover in the riparian zone or catchment (r² = 0.06–0.46). Using a combination of lake and landscape features in multiple regressions, we were able to explain 29–55% of the variation in macrophyte cover metrics. Total macrophyte cover and submersed cover were related to Secchi disk depth and mean depth, whereas the remaining metrics were best predicted by including at least one land use/cover variable (road density, proportion local catchment agriculture land use/cover, proportion cumulative catchment urban land use/cover, or proportion riparian agriculture land use/cover). The two main conclusions from our research are: (1) that different macrophyte growth forms and species are predicted by a different suite of variables and thus should be examined separately, and (2) that anthropogenic landscape features may override patterns in natural landscape or local features and are important in predicting present-day macrophytes in lakes.     

Bird abundance and species richness on Florida lakes: influence of trophic status,lake morphology,and aquatic macrophytes

Data from 46 Florida lakes were used to examine relationships between bird abundance (numbers and biomass) and species richness, and lake trophic status, lake morphology and aquatic macrophyte abundance. Average annual bird numbers ranged from 7 to 800 birds km^–2 and bird biomass ranged from 1 to 465 kg km^–2. Total species richness ranged from 1 to 30 species per lake. Annual average bird numbers and biomass were positively correlated to lake trophic status as assessed by total phosphorus (r = 0.61), total nitrogen (r = 0.60) and chlorophyll a (r = 0.56) concentrations. Species richness was positively correlated to lake area (r = 0.86) and trophic status (r = 0.64 for total phosphorus concentrations). The percentage of the total annual phosphorus load contributed to 14 Florida lakes by bird populations was low averaging 2.4%. Bird populations using Florida lakes, therefore, do not significantly impact the trophic status of the lakes under natural situations, but lake trophic status is a major factor influencing bird abundance and species richness on lakes. Bird abundance and species richness were not significantly correlated to other lake morphology or aquatic macrophyte parameters after the effects of lake area and trophic status were accounted for using stepwise multiple regression. The lack of significant relations between annual average bird abundance and species richness and macrophyte abundance seems to be related to changes in bird species composition. Bird abundance and species richness remain relatively stable as macrophyte abundance increases, but birds that use open-water habitats (e.g., double-crested cormorant, Phalacrocorax auritus) are replaced by species that use macrophyte communities (e.g., ring-necked duck, Aythya collaris).     

Recovery of lake vegetation following reduced eutrophication and acidification

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Response of native Egeria najas Planch. and invasive Hydrilla verticillata (L.f.) Royle to altered hydroecological regime in a subtropical river

Egeria najas Planch. is the dominant native submersed macrophyte of the Upper Paraná River in Brazil, while Hydrilla verticillata (L.f.) Royle has recently invaded this area. From January 2006 to December 2007, comprising two annual flood cycles, we conducted monthly surveys at two river stations and two lakes connected to the river within this stretch of the Paraná River, aiming to understand how the hydrological regime influences the distribution and abundance of these native and invasive Hydrocharitaceae species. Hydrilla did not develop in the lakes, possibly due to the elevated proportion of organic matter in the sediment (∼10% DW). However, the exotic species dominated the river sites apparently suppressing E. najas. In the lakes E. najas reached a maximum biomass of 628 ± 82 g DW m⁻² but did not surpass 333 ± 83 g DW m⁻² in the river, where H. verticillata peaked at 1415 ± 255 g DW m⁻². Macrophyte biomass development was greatest during low-water periods, with transparent water and high temperatures. Floods probably affected submersed macrophytes (especially in 2007, when an extreme flood caused by an El Niño Southern Oscillation (ENSO) event occurred) via sediment movement and plant scouring (uprooting) effects, coupled with reduced water transparency. Macrophyte recovery started soon after the (less intense) 2006 flood but was delayed in 2007. In the river recovery started five months after the major flood, but in the lakes no significant plant regeneration was found even nine months after the disturbance. E. najas and H. verticillata started regeneration practically at the same time but H. verticillata had much higher rates of biomass increase.     

A comparison between aquatic birds of lakes and coastal rivers in Florida

Aquatic birds were counted on five Gulf coast Florida rivers to determine if these river systems supported densities, biomass and species richness similar to those found on Florida lakes. Forty-two species were identified and for the species that were found on both Florida streams and lakes similar densities and biomass were encountered. As with Florida lakes, stream bird abundance and species richness were higher in winter months than in summer months, a consequence of migratory bird populations. Total bird abundance, biomass per unit of phosphorus, and species richness per unit of area were similar to data collected on Florida lakes. Thus, Florida rivers are capable of supplying sufficient resources to maintain bird densities, biomass and species richness values similar to lakes of equal size and nutrient concentrations and are therefore important habitats for aquatic bird populations. An examination of individual habitat characteristics indicates that water depth was inversely correlated and submersed aquatic vegetation was positively correlated with bird density, biomass and species richness within the river systems. While both habitat characteristics are important they are also inversely related making it difficult to separate the individual significance of each characteristic.     

Is the island biogeography model a poor predictor of biodiversity patterns in shallow lakes?

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Study of submerged aquatic vascular plants in Northern Glacial Lakes (New York State,U.S.A.)

The submerged aquatic vascular vegetation was studied in the Finger Lakes and in two bays of Lake Ontario in northern New York State, U.S.A. The species composition, structure and biomass of the communities are related to the principal habitat factors such as trophic status of the lakes, bottom slope and substrate, current and stream effects, and depth. A record of the development of changes in the trophic status of the lakes and in the species composition of their submerged vascular vegetation is outlined. An attempt is made to assess the effects of catastrophic events (floods and cloudbursts in the watershed) on the lake vegetation.     

The role of algae in the removal of Escherichia coli in a tropical eutrophic lake

Eutrophication and its accompanying algal development in lakes is a nuisance and may be problematic for aquatic life, but limited algal development may have some beneficial consequences. Dissolved oxygen concentration and pH increases attributed to algae in algal-based treatment ponds may occur in eutrophic lakes and can result in the inactivation of faecal coliforms in eutrophic lakes. We investigated the die-off of Escherichia coli placed in dialysis tubes in a eutrophic lake at different depths and locations. The importance of E. coli attachment to algae and suspended matter was also assessed. Algal presence in the lake resulted in significant decay of E. coli. At chlorophyll a concentration ≤0.08 mg L⁻¹ in Weija Lake, decay rate of E. coli is directly proportional to the chlorophyll a concentration of the lake. Under laboratory conditions, as chlorophyll a concentration increases in light however, an optimum chlorophyll a concentration (0.24 mg/L) is reached after which the rate of decay of E. coli decreases. These results show that limited algal presence representing optimum chlorophyll a concentration in restored ecosystems may have public health benefits for rural communities in developing countries that depend on raw water for domestic activities.     

Biomass and primary production of a 8–11 m depth meadow versus <3 m depth meadows of the seagrass Cymodocea nodosa (Ucria) Ascherson

Current knowledge about the abundance, growth, and primary production of the seagrass Cymodocea nodosa (Ucria) Ascherson is biased towards shallow (depth <3 m) meadows although this species also forms extensive meadows at larger depths along the coastlines. The biomass and primary production of a C. nodosa meadow located at a depth of 8–11 m was estimated at the time of maximum annual vegetative development (summer) using reconstruction techniques, and compared with those available from shallow meadows of this species. A depth-referenced data base of values at the time of maximum annual development was compiled to that end. The vegetative development of C. nodosa at 8–11 m depth was not different from that achieved by shallow (depth <3 m) meadows of this species. Only shoot density, which decreased from 1637 to 605 shoots m⁻², and the annual rate of elongation of the horizontal rhizome, which increased from 23 to 71 cm apex⁻¹ year⁻¹, were different as depth increased from <3 to 8–11 m. Depth was a poor predictor of the vegetative development and primary production of C. nodosa. The biomass of rhizomes and roots decreased with depth (g DW m⁻² = 480 (±53, S.E.) − 32 (±15, S.E.) depth (in m); R² = 0.12, F = 4.65, d.f. = 35, P = 0.0381) which made total biomass of the meadow to show a trend of decrease with depth but the variance of biomass data explained by depth was low. The annual rate of elongation of the horizontal rhizome showed a significant positive relationship with depth (cm apex⁻¹ year⁻¹ = 18 (±5.1, S.E.) + 5.0 (±1.33, S.E.) depth (in m); R² = 0.50, F = 14.07, d.f. = 14, P = 0.0021). As shoot size and growth did not change significantly with depth, the reduction of shoot density should drive any changes of biomass and productivity of C. nodosa as depth increases. The processes by which this reduction of C. nodosa abundance with depth occur remain to be elucidated.     

Long-term changes in macrophyte vegetation after reduction of fish stock in a shallow lake

Starting in the middle of the 1970s, submerged macrophytes began to disappear from shallow Lake Warniak due to feeding pressure by grass carp (Ctenopharyngodon idella). In the middle of the 1980s, the lake was stocked with seston-feeding silver carp (Hypophthalmichthys molitrix) and bighead carp (Aristichthys nobilis). By 1993, the biomass of silverhead carp and bighead carp had declined. This allowed charophytes to recolonize the bottom of the lake. The main charophyte species at this time were Chara globularis and Chara rudis. Since then, five other stonewort species have been found: Chara contraria, C. filiformis, C. tomentosa, C. aspera and Nitellopsis obtusa. Seventeen species of aquatic angiosperms have also been found. There were distinct changes in the relative abundance and spatial distribution of particular species. C. rudis developed most intensely in the shallow parts of the lake near the southern and western shores. C. globularis gradually took over the deeper central part of the lake. In 2001, C. rudis began to retreat again. The relative abundance and spatial distribution of charophytes was correlated to water clarity (r = 0.87, p < 0.05), total phosphorus level (r = −0.78; p < 0.05), and chlorophyll a content (r = −0.79; p < 0.05).     

Influence of environmental variables on macroinvertebrate community structure in Lianhuan Lake

The structural characteristics of the macroinvertebrate community can effectively reflect the health status of lake ecosystems and the quality of the lake ecological environment. It is therefore important to identify the limiting factors of macroinvertebrate community structure for the maintenance of lake ecosystem health. In this study, the community composition of macroinvertebrate assemblages and their relationships with environmental variables were investigated in 13 small lakes within Lianhuan Lake in northern China. A self‐organizing map and K‐means clustering analysis grouped the macroinvertebrate communities into five groups, and the indicator species reflected the environmental characteristics of each group. Principal component analysis indicated that the classification of the macroinvertebrate communities was affected by environmental variables. The Kruskal–Wallis test results showed that environmental variables (pH, total phosphorus, nitrate, water temperature, dissolved oxygen, conductivity, permanganate index, and ammonium) had a significant effect on the classification of the macroinvertebrate communities. Redundancy analysis showed that mollusks were significantly negatively correlated with pH and chlorophyll a, while annelids and aquatic insects were significantly positively correlated with chlorophyll a and dissolved oxygen. Spearman correlation analysis showed that the species richness and Shannon''s diversity of macroinvertebrates were significantly negatively correlated with total phosphorus, while the biomass of macroinvertebrates was significantly negatively correlated with pH. High alkalinity and lake eutrophication have a serious impact on the macroinvertebrate community. Human disturbances, such as industrial and agricultural runoff, negatively impact the ecological environment and affect macroinvertebrate community structure. Thus, macroinvertebrate community structure should be improved by enhancing the ecological environment and controlling environmental pollution at a watershed scale.     

Genetic diversity of Potamogeton pectinatus L. in relation to species diversity in a pair of lakes of contrasting trophic levels

Understanding the correlation between genetic diversity and species diversity in freshwater communities is important to elucidate the influences of local selective forces on the genetic diversity of local aquatic plant populations across different communities. This study employed amplified fragment length polymorphism (AFLP) to assess the genetic diversity of Potamogeton pectinatus L. populations between two sister-lakes with contrasting trophic levels, eutrophic and oligotrophic, in the Yunnan Plateau in southwest of China. The results showed high genetic differentiation between eutrophic lake and oligotrophic lake. The genetic distances between P. pectinatus populations were significantly correlated with the species evenness, but not with difference in species richness of aquatic plant communities. The results underpinned that genetic diversity at inter-population levels and local species diversity in plant communities are positively correlated. In addition, our results also suggested that habitat types might play an important role in the genetic diversity of the P. pectinatus populations between these two lakes.     

Richness of aquatic macrophyte floras of soft water lakes of differing pH and trace metal content in Ontario,Canada

The richness of the aquatic macrophyte floras, i.e., the total number of species, was assessed in 39 soft water lakes in central Ontario, Canada. The Cu and Ni concentrations and pH of the lakes ranged from 1 to 360 mg m^?3, 2 to 3700 mg m^?3 and 3.9 to 7.0, respectively. Two non-exclusive subsets of the data were examined to determine firstly, if floral richness was related to lake pH in lakes with low Cu and Ni levels (Data Set I) and secondly, if floral richness in acidic (pH ? 5.3) lakes was related to levels of various trace metals (Data Set II). Charophytes were not found in lakes with pH < 5.2. In Data Set I, there was no relationship between the richness of tracheophytes and pH, and there was a negative relationship between pH and bryophyte richness. Unlike phytoplankton, zooplankton, benthic macroinvertebrates and fish, there was no decrease in total species richness in lakes of pH < 5.5, as long as trace metal levels were low. Examination of Data Set II indicated tracheophyte richness of acidic lakes was negatively correlated with Cu and Ni levels. Biological surveys of metal-contaminated acidic lakes are, therefore, not of use for predicting the effects of acid deposition alone on aquatic macrophytes.     

Do freshwater plants have adaptive responses to typhoon-impacted regimes?

In tropical regimes, cyclones exert great influence on the local aquatic habitats. The objective of our study was to investigate if aquatic plants have an adaptive response to typhoon influence. Population traits of six aquatic species in different life-forms (emergent species: Scirpus triangulatus, Eleocharis plantagineiformis, Rotala rotundifolia, Eriocaulon buergerianum; submerged: Blyxa echinosperma; floating-leaved: Nymphoides indica) were investigated to compare intraspecific variations in high and low typhoon-impacted regions on Hainan Island in southern China. In the high typhoon-impacted region, there was greater belowground biomass allocation in both emergent and floating-leaved species. The ratio of belowground to total biomass of each emergent was 41% (P = 0.028), 38% (P = 0.034), 27% (P = 0.040), 19% (P = 0.043) greater respectively, and floating-leaved N. indica was 40% (P = 0.014) greater than in the low typhoon region. The stem height of relatively tall emergent species (S. triangulatus and E. plantagineiformis) was 35% (P = 0.033), 42% (P = 0.046) lower, and floating-leaved species N. indica had decreased leaf area (49%, P < 0.001) and number (30%, P < 0.001) on water surface in the high typhoon-impacted region than in the low. These adaptations of the plants will reduce their risk of mechanical damage from strong winds or wind-induced currents. Submerged species in the study showed no variation in traits between the high and low typhoon-impacted regions.     

Effects of invasive species on plant communities: an example using submersed aquatic plants at the regional scale

Submersed aquatic plants have a key role in maintaining functioning aquatic ecosystems through their effects on the hydrological regime, sedimentation, nutrient cycling and habitat of associated fauna. Modifications of aquatic plant communities, for example through the introduction of invasive species, can alter these functions. In the Sacramento-San Joaquin River Delta, California, a major invasive submersed plant, Brazilian waterweed Egeria densa, has become widespread and greatly affected the functionality of the submersed aquatic plant community. Rapid assessments of the distribution and abundance of this species are therefore crucial to direct management actions early in the season. Given the E. densa bimodal growth pattern (late spring and fall growth peaks), summer assessments of this species may indicate which and where other submersed species may occur and fall assessments may indicate where this and other species may occur in the following spring, primarily because the Delta’s winter water temperatures are usually insufficient to kill submersed aquatic plant species. We assessed community composition and distribution in the fall of 2007 and summer of 2008 using geostatistical analysis; and measured summer biomass, temperature, pH, salinity, and turbidity. In the fall of 2007, submersed aquatic plants covered a much higher proportion of the waterways (60.7%) than in the summer of 2008 (37.4%), with a significant overlap between the seasonal distribution of native and non-native species. Most patches were monospecific, and multispecies patches had significantly higher dominance by E. densa, co-occurring especially with Ceratophyllum demersum. As species richness of non-natives increased there was a significant decrease in richness of natives, and of native biomass. Sustained E. densa summer biomass negatively affected the likelihood of presence of Myriophyllum spicatum, Potamogeton crispus, and Elodea canadensis but not their biomass within patches. Depth, temperature and salinity were associated with biomass; however, the direction of the effect was species specific. Our results suggest that despite native and invasive non-native submersed plant species sharing available niches in the Delta, E. densa affects aquatic plant community structure and composition by facilitating persistence of some species and reducing the likelihood of establishment of other species. Successful management of this species may therefore facilitate shifts in existing non-native or native plant species.     

The submersed macrophyte communities of adirondack lakes (New York,U.S.A.) of varying degrees of acidity

The submersed macrophyte communities of nine high-altitude oligotrophic lakes in the Adirondack Mountain region of upstate New York were characterized in detail in terms of percent cover of each species. The pH of these clear-water lakes ranged from 6.9 to 4.4 and the acid neutralizing capacity ranged from 91 μeq 1^?1 to ?38 μeq l^?1. The degree of acidity covaried with lower total ion content and nutrient status. The dominant species in the circumneutral lakes and in the acidic lakes were the same, with several Utricularia species being the major component of the bottom cover. Species richness and diversity were lower in the acidic lakes (pH < 5.5). The plants found in the acidic lakes were typical of oligotrophic systems in general, regardless of pH, and similar to those found in lake surveys in Scandinavia and Eastern Canada. Potamogeton confervoides Reichenb., Scirpus subterminalis Torr., Sphagnum spp. and Utricularia geminiscapa Benj. were the only species that were primarily exclusive to the acidic lakes surveyed. In contrast to the results of Scandinavian studies, Sphagnum was found to be a major component of the plant cover in only two of the five acidic lakes.