Resource Availability and Spatial Heterogeneity Control Bacterial Community Response to Nutrient Enrichment in Lakes

The diversity and composition of ecological communities often co-vary with ecosystem productivity. However, the relative importance of productivity, or resource abundance, versus the spatial distribution of resources in shaping those ecological patterns is not well understood, particularly for the bacterial communities that underlie most important ecosystem functions. Increasing ecosystem productivity in lakes has been shown to influence the composition and ecology of bacterial communities, but existing work has only evaluated the effect of increasing resource supply and not heterogeneity in how those resources are distributed. We quantified how bacterial communities varied with the trophic status of lakes and whether community responses differed in surface and deep habitats in response to heterogeneity in nutrient resources. Using ARISA fingerprinting, we found that bacterial communities were more abundant, richer, and more distinct among habitats as lake trophic state and vertical heterogeneity in nutrients increased, and that spatial resource variation produced habitat specific responses of bacteria in response to increased productivity. Furthermore, changes in communities in high nutrient lakes were not produced by turnover in community composition but from additional taxa augmenting core bacterial communities found in lower productivity lakes. These data suggests that bacterial community responses to nutrient enrichment in lakes vary spatially and are likely influenced disproportionately by rare taxa.     

The influence of a dissolved inorganic nitrogen gradient on phytoplankton community dynamics in a chain of lakes

In a chain of lakes along which nutrient availability varies in a gradient, we performed factorial nutrient enrichment experiments to determine if nitrogen limitation was the principal factor controlling the differences in phytoplankton biomass, photosynthetic productivity, diversity, and species composition among two of the lakes in the chain. In the least productive lake, East Graham Lake, P and C enrichments (in the absence of N enrichment) had no effect on biomass and diversity, whereas within two weeks the N enrichments (alone or in any combination with P and/or C) increased the biomass and decreased the diversity of East Graham Lake phytoplankton to levels similar or identical to those in more productive Shoe Lake. Short-term ¹⁴C photosynthetic rates in East Graham Lake water also responded only to N in the third week. However, photosynthesis was stimulated by P in the first week, and a few species did increase in numbers with P enrichment, suggesting that some degree of P limitation remains in addition to the strong N limitation in East Graham Lake. A number of species responded individually to the enrichments in a manner similar to that of the overall community, and a strong overlapping of discriminant analysis scores for N-enriched East Graham Lake with those of Shoe Lake was consistent with our prediction that the community structure of N-enriched East Graham Lake water would shift toward that of Shoe Lake. However, many species did not respond consistently with these results, and the nutrients tested were clearly not a major factor in the differences in abundance of those species among the two lakes. The results support the argument that overall biomass production and diversity of the phytoplankton community in a lake can be a relatively simple function of a single most-limiting nutrient. However, many of the species responses also confirm that, while nutrient availability is an important factor in the control of the species composition of the community, other factors are likely to prevent reliable predictions of all species effects on the basis of nutrient availability alone.     

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

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Salmon‐derived nutrients drive diatom beta‐diversity patterns

1. Pacific salmon are a textbook example of migratory animals that transfer nutrients between ecosystems, but little is known about how salmon‐derived nutrients (SDN) affect the biodiversity of recipient freshwater ecosystems. We examined paleolimnological records from six Alaskan lakes to define how changes in SDN from sockeye salmon (Oncorhynchus nerka) influenced sedimentary diatom community structure and beta‐diversity among lakes and through time. 2. Using an isotopic mixing model, we showed that SDN loading could account for >80% of the lake total nitrogen budgets and strongly regulated diatom community composition. Spatial dissimilarity in diatom communities was positively related to differences in SDN among lakes (r² = 0.69, P < 0.01, n = 10). Likewise, temporal dissimilarity in diatom communities was positively related to differences in SDN in a sediment core with substantial variation in salmon spawner dynamics between 1700 and 1950 AD (r² = 0.34, P < 0.01, n = 19). Finally, beta‐diversity metrics quantifying temporal turnover within each lake’s sediment record were also positively related to the variance in SDN loading among lakes (r² = 0.88, P < 0.05, n = 5). Mean SDN was only negatively correlated to temporal diatom beta‐diversity. 3. Spatially subsidised systems often receive temporally variable resource inputs, and thus, it is not surprising that, unlike previous studies, we found that resource variability was the key driver of community composition and beta‐diversity. In habitats that receive strongly fluctuating external nutrient loads, environment heterogeneity may overweigh stochastic community processes. In addition, freshwater diatoms are characterised by great dispersal capabilities and short life cycles and therefore may be a more sensitive indicator for evaluating the role of resource variability than previously used model organisms. These results suggest that productivity–diversity relationship vary with the nature of nutrient loading and the life history of the community studied. 4. Overall, our study highlights that the transport of nutrients by sockeye salmon across ecosystem boundaries is a significant driver of algal community and biodiversity in nursery lakes, mainly through changing the magnitude of nutrient variation. As such, freshwater species diversity in regions like the U.S. Pacific Northwest may become impoverished where there have been long‐term declines in salmon populations and decreases in nutrient variability among lakes.     

Seasonal dynamics of the planktonic microbial community in a maritime Antarctic lake undergoing eutrophication

The abundance, biomass and community structure of phytoplankton,bacterioplankton and protozooplankton in a maritime Antarcticlake were determined at approximately monthly intervals fromDecember 1994 to February 1996 and compared with data from earlierstudies. Heywood Lake has become significantly eutrophic duringthe last three decades because of excreta from the expandingfur seal population in its catchment. Marked seasonal variationsin the abundance, composition and productivity of biota werecorrelated with the seasonality of both physical factors andnutrient levels. Protozooplankton were abundant, diverse andusually dominated by heterotrophic nanoflagellates (HNF), withHNF abundance peaking at 2.35 x 10⁷ l^–1 in summer. Highnumbers of naked amoebae were sometimes present, reaching amaximum of 4.8 x 10³ l^–1 in March. An estimated 89 speciesof protozoa were observed during the study, indicating substantiallymore diversity than is found in continental Antarctic lakes.Diversity was highest in spring and lowest in winter, when theentire water column became anoxic and the plankton were dominatedby bacteria and a few species of relatively large anaerobicflagellates. The current status of the lake is compared withdata for continental Antarctic and lower latitude lakes. Earlierstudies of biota and physical/chemical parameters in HeywoodLake are used to examine the effects of eutrophication overthree decades. Observed changes include increased microbialabundance and changes in both community structure and seasonalpatterns.     

Assessing lake eutrophication using chironomids: understanding the nature of community response in different lake types

1. Total phosphorus (TP) and chlorophyll a (Chl a) chironomid inference models ( Brodersen & Lindegaard, 1999 ; Brooks, Bennion & Birks, 2001 ) were used in an attempt to reconstruct changes in nutrients from three very different lake types. Both training sets were expanded, particularly at the low end of the nutrient gradient, using contemporary chironomid assemblages and environmental parameters from 12 British lakes, although this had little improvement on the model performances. 2. Dissimilarity analyses showed that the historic chironomid assemblages did not have good analogues in the original calibration or extended datasets. However, since the transfer functions are based on weighted averages of the trophic optima for the taxa present and not on community similarities, reasonable downcore inferences were produced. Ordination analyses also showed that the lakes retain their ‘identity’ over time, as the sample dissimilarities within lakes were less than the dissimilarities between lakes. 3. Analysis of the three historic lake profiles showed a range of chironomid community responses to lake development. Chironomids from a shallow lake, Slapton Ley, responded indirectly to nutrient enrichment (TP), probably through altered substrate, macrophyte and fish conditions, rather than directly to primary productivity (Chl a). A stratified lake, Old Mill Reservoir, showed a loss of the profundal chironomid fauna due to increasing primary productivity (Chl a) coupled with increasing hypoxia. A response to nutrients (TP or total nitrogen (TN)) at this site is also indirect, and the TP reconstruction therefore cannot be reliably interpreted. The third lake, March Ghyll Reservoir has little change in historic chironomid communities, suggesting that this well mixed, relatively unproductive lake has changed less than the other lakes. 4. Using chironomids to reconstruct nutrient histories does not follow a simple scheme. The response to changes in nutrients may be direct, but mediated through other ecosystem components. As alternative stable states are possible at a given level of TP it is also likely that alternative chironomid communities exist under similar nutrient conditions. Changes in biological communities can thus occur over thresholds, and it is only biological proxies that can reflect such ecosystem switches within palaeoenvironmental investigations.     

Intraspecific phenotypic variation in a fish predator affects multitrophic lake metacommunity structure

Contemporary insights from evolutionary ecology suggest that population divergence in ecologically important traits within predators can generate diversifying ecological selection on local community structure. Many studies acknowledging these effects of intraspecific variation assume that local populations are situated in communities that are unconnected to similar communities within a shared region. Recent work from metacommunity ecology suggests that species dispersal among communities can also influence species diversity and composition but can depend upon the relative importance of the local environment. Here, we study the relative effects of intraspecific phenotypic variation in a fish predator and spatial processes related to plankton species dispersal on multitrophic lake plankton metacommunity structure. Intraspecific diversification in foraging traits and residence time of the planktivorous fish alewife (Alosa pseudoharengus) among coastal lakes yields lake metacommunities supporting three lake types which differ in the phenotype and incidence of alewife: lakes with anadromous, landlocked, or no alewives. In coastal lakes, plankton community composition was attributed to dispersal versus local environmental predictors, including intraspecific variation in alewives. Local and beta diversity of zooplankton and phytoplankton was additionally measured in response to intraspecific variation in alewives. Zooplankton communities were structured by species sorting, with a strong influence of intraspecific variation in A. pseudoharengus. Intraspecific variation altered zooplankton species richness and beta diversity, where lake communities with landlocked alewives exhibited intermediate richness between lakes with anadromous alewives and without alewives, and greater community similarity. Phytoplankton diversity, in contrast, was highest in lakes with landlocked alewives. The results indicate that plankton dispersal in the region supplied a migrant pool that was strongly structured by intraspecific variation in alewives. This is one of the first studies to demonstrate that intraspecific phenotypic variation in a predator can maintain contrasting patterns of multitrophic diversity in metacommunities.     

Characterization of bacterial communities in four freshwater lakes differing in nutrient load and food web structure

The phylogenetic composition of bacterioplankton communities in the water column of four shallow eutrophic lakes was analyzed by partially sequencing cloned 16S rRNA genes and by PCR-DGGE analysis. The four lakes differed in nutrient load and food web structure: two were in a clearwater state and had dense stands of submerged macrophytes, while two others were in a turbid state characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had very high nutrient levels (total phosphorus > 100 microg/l), while the other lakes were less nutrient rich (total phosphorus < 100 microg/l). Cluster analysis, multidimensional scaling and ANOSIM (analysis of similarity) were used to investigate differences among the bacterial community composition in the four lakes. Our results show that each lake has its own distinct bacterioplankton community. The samples of lake Blankaart differed substantially from those of the other lakes; this pattern was consistent throughout the year of study. The bacterioplankton community composition in lake Blankaart seems to be less diverse and less stable than in the other three lakes. Clone library results reveal that Actinobacteria strongly dominated the bacterial community in lake Blankaart. The relative abundance of Betaproteobacteria was low, whereas this group was dominant in the other three lakes. Turbid lakes had a higher representation of Cyanobacteria, while clearwater lakes were characterized by more representatives of the Bacteroidetes. Correlating our DGGE data with environmental parameters, using the BIOENV procedure, suggests that differences are partly related to the equilibrium state of the lake.     

Sensitivity and responses of diatoms to climate warming in lakes heavily influenced by humans

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Species-specific algal responses to zooplankton: experimental and field observations in three nutrient-limited lakes

A series of 4-day manipulations of zooplankton biomass and nutrientavailability was performed in enclosures in three lakes to determinespecies-specific algal responses to herbivory and nutrient enrichment.Algal performance in enclosures was compared to the relationshipsbetween weekly algal growth rates and the zooplankton in situ.When in situ growth rates were significant functions of zooplanktonbiomass, the responses were generally consistent with responsesin the enclosure experiments. The importance of both nutrientsand zooplankton in mediating algal growth was demonstrated bynumerous observations: strong algal community response to enrichment,unimodal or positive responses of certain algal taxa to zooplanktonbiomass, differences in degree of nutrient limitation amongthe algal response types, lack of nutrient limitation of non-grazedalgal taxa and a preponderance of taxa with no net responseto increasing zooplankton biomass. Variation in the zooplanktoncommunity may be the largest source of variability in nutrientsupply rate during summer in stratified lakes, and causes substationalvariability in the algae. Algae responded more strongly to changesin zooplankton composition than to changes in zooplankton biomass.We conclude that, due to the close coupling of phytoplanktonand zooplankton communities in these nutrient-limited lakes,major compositional changes in the zooplankton have greatereffects on the algae than do changes in biomass of grazers alreadypresent. ¹Present address: Division of Environmental Studies, Universityof California, Davis, CA 95616, USA ²Present address: Division of Biological Sciences, Universityof California, Davis, CA 95616, USA     

Seasonal Dynamics of Periphyton in a Large Tropical Lake

Tropical aquatic systems are generally assumed to have little seasonality in productivity patterns. However, this study indicated that there was substantial seasonal variation in epilithic productivity and biomass in tropical Lake Tanganyika, due primarily to seasonal patterns in lake hydrodynamics that influence nutrient availability. Although they support much of the lake’s biological diversity, epilithic algae made a minor contribution to the total energy budget in Lake Tanganyika. A comparison among large, oligotrophic lakes revealed no significant latitudinal trends in periphyton productivity or biomass. However, Lake Tanganyika has relatively low benthic algal biomass and is therefore more efficient at photosynthesis than the temperate lakes. The influence of wave action and consumer density and diversity may be important in moderating productivity of the epilithic community.     

Long-term zooplankton composition data reveal impacts of invasions on community composition in the Waikato lakes,New Zealand

Data collected on zooplankton community composition over longer time periods (>?10 years) are rare. We examined among-lake spatial and temporal trends of zooplankton communities from a monitoring programme undertaken in the Waikato region, New Zealand. A total of 39 lakes were sampled over a period of 12 years, between 2007 and 2019, with varying degrees of temporal effort. We focussed particularly on eight lakes, considered here as ‘long-term lakes’, where samples were collected with greater regularity (including 5 with 12 years of data). Among lakes, suspended sediment concentrations and indicators of lake trophic state were inferred to be important in determining the zooplankton distributions; as this region is dominated by shallow lakes, the relative importance of suspended sediments was high. Among the long-term lakes, the greatest dissimilarities in zooplankton community composition among years were in Lake Waahi, where the Australian Boeckella symmetrica was first detected in 2012. That is, the greatest temporal changes to zooplankton composition during the study period were due to the invasion by non-indigenous species, rather than changes in trophic state or other environmental variables; non-native species commonly dominated the individual counts of species through much of 2014 and 2015, with most samples since 2016 being again dominated by native species. Following this lake, the largest and shallowest lakes in the dataset—Whangape and Waikare—exhibited the greatest variability in community composition among years.

     

Comparative analysis of rotifer community structure in five subtropical shallow lakes in East China: role of physical and chemical conditions

Worldwide, there have been few comparative studies on rotifer communities in subtropical lakes. We studied changes in rotifer community structure over 1 year and its relationship to several physicochemical variables in five subtropical shallow lakes in East China, covering a nutrient gradient from mesotrophy to moderate eutrophy. In these lakes, the genera Brachionus, Lecane, and Trichocerca dominated the rotifer species composition, and Polyarthra dolichoptera, Keratella cochlearis, Filinia longiseta, T. pusilla, and Anuraeopsis fissa were the dominant species. With increased nutrient loading, total rotifer abundance and species dominance increased, indicating that rotifer abundance might be a more sensitive indicator of trophic state than species composition. Comparative analyses of the six rotifer community indices calculated in this study and redundancy analysis (RDA) revealed that the two slightly eutrophic lakes and the other two moderately eutrophic lakes exhibited a high degree similarity in community structure. This suggests that the trophic state of a lake determines the rotifer community structure. In contrast, in the two moderately eutrophic lakes, the mass ratios of TN:TP and the contents of TP suggested N-limitation and cyanobacteria dominance in phytoplankton communities might be possible. In these lakes TN played a more important role in shaping the rotifer community according to stepwise multiple regression and RDA. RDA analysis also suggested that rotifer species distribution was strongly associated with trophic state and water temperature, with water temperature being the most important factor in determining seasonality.     

Analysis of the community structure of planktonic ciliated protozoa relative to trophic state in Florida lakes

The planktonic ciliate populations of 30 Florida lakes constituting a broad trophic gradient were examined to determine the response of protozoan community structure to increasing eutrophication. Both ciliate abundance and biomass were strongly related to lake trophic state. Comparison of the Florida data base with a comparable north temperate lake group indicated that subtropical lakes generally possess higher ciliate abundance and biomass at a given trophic state than temperate lakes. However the equations derived for each data base were not significantly different. Community diversity and species richness increased with increasing lake productivity. Highly acidic lakes displayed significantly reduced diversity and numbers of species when contrasted with nonacidic oligotrophic lakes. Small-bodied (< 30 um) ciliates dominated all lakes but were proportionally less important in oligotrophic lakes. Presence-absence data produced three assemblages: an ubiquitous association of primarily small ciliate taxa, a group of large ciliates mainly restricted to eutrophic-hypereutrophic lakes, and a very large ciliate,Stentor niger, which dominated the protozoan communities of acidic oligotrophic lakes.     

A Framework for Understanding Variation in Pelagic Gross Primary Production of Lake Ecosystems

Light and nutrient availability are key physiological constraints for primary production. Widespread environmental changes are causing variability in loads of terrestrial dissolved organic carbon (DOC) and nutrients from watersheds to lakes, contributing to simultaneous changes in both light and nutrient supply. Experimental evidence highlights the potential for these watershed loads to create complex and context-dependent responses of within-lake primary production; however, the field lacks a predictive model to investigate these responses. We embedded a well-established physiological model of phytoplankton growth within an ecosystem model of nutrient and DOC supply to assess how simultaneous changes in DOC and nutrient loads could impact pelagic primary production in lakes. The model generated a unimodal relationship between GPP and DOC concentration when loads of DOC and nutrients were tightly correlated across space or time. In this unimodal relationship, the magnitude of the peak GPP was primarily determined by the DOC-to-nutrient ratio of the load, and the location of the peak along the DOC axis was primarily determined by lake area. Greater nutrient supply relative to DOC load contributed to greater productivity, and larger lake area increased light limitation for primary producers at a given DOC concentration, owing to the positive relationship between lake area and epilimnion depth. When loads of DOC and nutrients were not tightly correlated in space or time, the model generated a wedge-shaped pattern between GPP and DOC, consistent with spatial surveys from a global set of lakes. Our model is thus capable of unifying the diversity of empirically observed spatial and temporal responses of lake productivity to DOC and mineral nutrient supply presented in the literature, and provides qualitative predictions for how lake pelagic primary productivity may respond to widespread environmental changes.     

Long‐term nutrient addition alters arthropod community composition but does not increase total biomass or abundance

A simple bottom–up hypothesis predicts that plant responses to nutrient addition should determine the response of consumers: more productive and less diverse plant communities, the usual result of long‐term nutrient addition, should support greater consumer abundances and biomass and less consumer diversity. We tested this hypothesis for the response of an aboveground arthropod community to an uncommonly long‐term (24‐year) nutrient addition experiment in moist acidic tundra in arctic Alaska. This experiment altered plant community composition, decreased plant diversity and increased plant production and biomass as a deciduous shrub, Betula nana, became dominant. Consistent with strong effects on the plant community, nutrient addition altered arthropod community composition, primarily through changes to herbivore taxa in the canopy‐dwelling arthropod assemblage and detritivore taxa in the ground assemblage. Surprisingly, however, the loss of more than half of plant species was accompanied by negligible changes to diversity (rarefied richness) of arthropod taxa (which were primarily identified to family). Similarly, although long‐term nutrient addition in this system roughly doubles plant production and biomass, arthropod abundance was either unchanged or decreased by nutrient addition, and total arthropod biomass was unaffected. Our findings differ markedly from the handful of terrestrial studies that have found bottom‐up diversity cascades and productivity responses by consumers to nutrient addition. This is probably because unlike grasslands and salt marshes (where such studies have historically been conducted), this arctic tundra community becomes less palatable, rather than more so, after many years of nutrient addition due to increased dominance of B. nana. Additionally, by displacing insulating mosses and increasing the cover of shrubs that cool and shade the canopy microenvironment, fertilization may displace arthropods keenly attuned to microclimate. These results indicate that terrestrial arthropod assemblages may be more constrained by producer traits (i.e. palatability, structure) than they are by total primary production or producer diversity.     

松嫩湖群鱼类群落相似性

根据2008年5月至2011年1月对松嫩湖群20个主要渔业湖泊的鱼类资源调查,分析了该湖群鱼类区系特征和群落相似性状况。松嫩湖群的鱼类区系由4目9科34属46种和亚种构成,其中土著鱼类3目8科27属39种和亚种,包括中国特有种3种,中国易危种1种,冷水种5种;由5个区系复合体构成,以东部江河平原区系复合体为主体;鲤形目31种,鲤科26种,分别占优势;鱼类区系具有南北方物种相互渗透、古北界与东洋界交汇过渡的混色类群特征。目前松嫩湖群鱼类群落种类组成的相似度总体较低,群落数量结构的相似度总体较高,鱼类群落相似性面临的主要问题是自然与人为因素导致湖泊生态环境的变化和放养、移殖与过度捕捞导致鱼类资源的减少与小型化,二者的叠加效应使鱼类群落长期处于受损状态,群落结构及其相似性处在动态变化中,群落内种间关系的协调性、种群结构的合理性和群落结构的稳定性均在下降。针对这些情况和群落相似性现状,提出未来松嫩湖群湖泊渔业的发展方向是优化调整群落结构,发展多种群湖泊渔业,合理利用土著鱼类资源。     

Rotifer communities from some Araucanian lakes of southern Chile

During the last decade much information has been produced about the zooplankton communities in southern Chile; however, most of this is related to the crustacean assemblages. The present communication examines the spatial and temporal distribution of rotifer assemblages and their relation to the environmental variables during one-year period in four Araucanian lakes. A total of 19 species was found in these oligotrophic lakes. Keratella cochlearis, Synchaeta stylata, Trichocerca porcellus, Conochilus unicornis and Collotheca pelagica were widespread, and seven species exhibited a more restricted distribution among the lakes. Species richness varied from 6 to 12; similarly, one or two dominant species usually accounted for more than 80% of the total annual abundance. Similar dominant species occurred in two lakes, but their maximum peaks of abundance differed in time; in the remaining lakes the most important species were different. Calculated rotifer diversity showed a fluctuating pattern, with low values during the year in three lakes, and high ones in Lake Llanquihue. Species diversity was significantly related to species richness in all lakes. Discriminant analysis based on the occurrence and abundances of species throughout the year revealed that the rotifer assemblage in Lake Llanquihue was different from that in the rest of the Araucanian lakes. Furthermore, the same analysis using environmental variables showed that this lake is clearly discriminated from the others on the basis of the ionic composition of the water (i.e. Cl^–, Na⁺, Mg²⁺). Rotifer abundances in these lakes were significantly influenced by a number of abiotic variables, including those related to water ionic composition. These relationships may imply that the small differences in chemical characteristics of these lakes influence the structure of the rotifer community.