Global changes may be promoting a rise in select cyanobacteria in nutrient‐poor northern lakes

The interacting effects of global changes—including increased temperature, altered precipitation, reduced acidification and increased dissolved organic matter loads to lakes—are anticipated to create favourable environmental conditions for cyanobacteria in northern lakes. However, responses of cyanobacteria to these global changes are complex, if not contradictory. We hypothesized that absolute and relative biovolumes of cyanobacteria (both total and specific genera) are increasing in Swedish nutrient‐poor lakes and that these increases are associated with global changes. We tested these hypotheses using data from 28 nutrient‐poor Swedish lakes over 16 years (1998–2013). Increases in cyanobacteria relative biovolume were identified in 21% of the study sites, primarily in the southeastern region of Sweden, and were composed mostly of increases from three specific genera: Merismopedia, Chroococcus and Dolichospermum. Taxon‐specific changes were related to different environmental stressors; that is, increased surface water temperature favoured higher Merismopedia relative biovolume in low pH lakes with high nitrogen to phosphorus ratios, whereas acidification recovery was statistically related to increased relative biovolumes of Chroococcus and Dolichospermum. In addition, enhanced dissolved organic matter loads were identified as potential determinants of Chroococcus suppression and Dolichospermum promotion. Our findings highlight that specific genera of cyanobacteria benefit from different environmental changes. Our ability to predict the risk of cyanobacteria prevalence requires consideration of the environmental condition of a lake and the sensitivities of the cyanobacteria genera within the lake. Regional patterns may emerge due to spatial autocorrelations within and among lake history, rates and direction of environmental change and the niche space occupied by specific cyanobacteria.     

Eutrophication mediates rapid clonal evolution in Daphnia pulicaria

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Hemispheric‐scale patterns of climate‐related shifts in planktonic diatoms from North American and European lakes

A synthesis of over 200 diatom‐based paleolimnological records from nonacidified/nonenriched lakes reveals remarkably similar taxon‐specific shifts across the Northern Hemisphere since the 19th century. Our data indicate that these diatom shifts occurred in conjunction with changes in freshwater habitat structure and quality, which, in turn, we link to hemispheric warming trends. Significant increases in the relative abundances of planktonic Cyclotella taxa (P<0.01) were concurrent with sharp declines in both heavily silicified Aulacoseira taxa (P<0.01) and benthic Fragilaria taxa (P<0.01). We demonstrate that this trend is not limited to Arctic and alpine environments, but that lakes at temperate latitudes are now showing similar ecological changes. As expected, the onset of biological responses to warming occurred significantly earlier (P<0.05) in climatically sensitive Arctic regions (median age=ad 1870) compared with temperate regions (median age=ad 1970). In a detailed paleolimnological case study, we report strong relationships (P<0.005) between sedimentary diatom data from Whitefish Bay, Lake of the Woods (Ontario, Canada), and long‐term changes in air temperature and ice‐out records. Other potential environmental factors, such as atmospheric nitrogen deposition, could not explain our observations. These data provide clear evidence that unparalleled warming over the last few decades resulted in substantial increases in the length of the ice‐free period that, similar to 19th century changes in high‐latitude lakes, likely triggered a reorganization of diatom community composition. We show that many nonacidified, nutrient‐poor, freshwater ecosystems throughout the Northern Hemisphere have crossed important climatically induced ecological thresholds. These findings are worrisome, as the ecological changes that we report at both mid‐ and high‐latitude sites have occurred with increases in mean annual air temperature that are less than half of what is projected for these regions over the next half century.     

Asynchronous onset of eutrophication among shallow prairie lakes of the Northern Great Plains,Alberta, Canada

Coherent timing of agricultural expansion, fertilizer application, atmospheric nutrient deposition, and accelerated global warming is expected to promote synchronous fertilization of regional surface waters and coherent development of algal blooms and lake eutrophication. While broad‐scale cyanobacterial expansion is evident in global meta‐analyses, little is known of whether lakes in discrete catchments within a common lake district also exhibit coherent water quality degradation through anthropogenic forcing. Consequently, the primary goal of this study was to determine whether agricultural development since ca. 1900, accelerated use of fertilizer since 1960, atmospheric deposition of reactive N, or regional climate warming has resulted in coherent patterns of eutrophication of surface waters in southern Alberta, Canada. Unexpectedly, analysis of sedimentary pigments as an index of changes in total algal abundance since ca. 1850 revealed that while total algal abundance (as β‐carotene, pheophytin a) increased in nine of 10 lakes over 150 years, the onset of eutrophication varied by a century and was asynchronous across basins. Similarly, analysis of temporal sequences with least‐squares regression revealed that the relative abundance of cyanobacteria (echinenone) either decreased or did not change significantly in eight of the lakes since ca. 1850, whereas purple sulfur bacteria (as okenone) increased significantly in seven study sites. These patterns are consistent with the catchment filter hypothesis, which posits that lakes exhibit unique responses to common forcing associated with the influx of mass as water, nutrients, or particles.     

Lake responses to reduced nutrient loading – an analysis of contemporary long-term data from 35 case studies

1. 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 μg L^?1 before loading reduction), subtropical to temperate (latitude: 28–65°), and lowland to upland (altitude: 0–481 m). Shallow north‐temperate 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 μg 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 changes such as global warming. However, effects of global change are likely to run counter to reductions in nutrient loading rather than reinforcing re‐oligotrophication.     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Abundance and distribution of picoplankton in tropical, oligotrophic Lake Kivu, eastern Africa

1. We used flow cytometry to characterize freshwater photosynthetic picoplankton (PPP) and heterotrophic bacteria (HB) in Lake Kivu, one of the East‐African great lakes. Throughout three cruises run in different seasons, covering the four major basins, phycoerythrin‐rich cells dominated the PPP. Heterotrophic bacteria and PPP cell numbers were always high and spatial variations were modest. This represents an important difference from temperate and high latitude lakes that show high fluctuations in cell abundance over an annual cycle. 2. Three populations of picocyanobacteria were identified: one corresponded to single‐cells (identified as Synechococcus by epifluorescence microscopy, molecular methods and pigment content), and the two other that most probably correspond to two and four celled colonies of the same taxon. The proportion of these two subpopulations was greater under stratified conditions, with stronger nutrient limitation. 3. High PPP concentrations (c. 10⁵ cell mL^?1) relative to HB (c. 10⁶ cell mL^?1) were always found. Lake Kivu supports relatively less bacteria than phytoplankton biomass than temperate systems, probably as a consequence of factors such as temperature, oligotrophy, nutrient limitation and trophic structure. 4. A review of PPP concentration across aquatic systems suggests that the abundance of Synechococcus‐like cyanobacteria in large, oligotrophic, tropical lakes is very high. 5. Photosynthetic picoplankton cell abundances in the oligotrophic tropical lakes Kivu and Tanganyika are comparable to those of eutrophic temperate lakes. This apparently contradicts the view that PPP abundance increases with increasing eutrophy. More data on PPP in tropical lakes are needed to explore further this particular pattern.     

Warmer climates boost cyanobacterial dominance in shallow lakes

Dominance by cyanobacteria hampers human use of lakes and reservoirs worldwide. Previous studies indicate that excessive nutrient loading and warmer conditions promote dominance by cyanobacteria, but evidence from global scale field data has so far been scarce. Our analysis, based on a study of 143 lakes along a latitudinal transect ranging from subarctic Europe to southern South America, shows that although warmer climates do not result in higher overall phytoplankton biomass, the percentage of the total phytoplankton biovolume attributable to cyanobacteria increases steeply with temperature. Our results also reveal that the percent cyanobacteria is greater in lakes with high rates of light absorption. This points to a positive feedback because restriction of light availability is often a consequence of high phytoplankton biovolume, which in turn may be driven by nutrient loading. Our results indicate a synergistic effect of nutrients and climate. The implications are that in a future warmer climate, nutrient concentrations may have to be reduced substantially from present values in many lakes if cyanobacterial dominance is to be controlled.     

The fate of the Arctic seaweed Fucus distichus under climate change: an ecological niche modeling approach

Rising temperatures are predicted to melt all perennial ice cover in the Arctic by the end of this century, thus opening up suitable habitat for temperate and subarctic species. Canopy‐forming seaweeds provide an ideal system to predict the potential impact of climate‐change on rocky‐shore ecosystems, given their direct dependence on temperature and their key role in the ecological system. Our primary objective was to predict the climate‐change induced range‐shift of Fucus distichus, the dominant canopy‐forming macroalga in the Arctic and subarctic rocky intertidal. More specifically, we asked: which Arctic/subarctic and cold‐temperate shores of the northern hemisphere will display the greatest distributional change of F. distichus and how will this affect niche overlap with seaweeds from temperate regions? We used the program MAXENT to develop correlative ecological niche models with dominant range‐limiting factors and 169 occurrence records. Using three climate‐change scenarios, we projected habitat suitability of F. distichus – and its niche overlap with three dominant temperate macroalgae – until year 2200. Maximum sea surface temperature was identified as the most important factor in limiting the fundamental niche of F. distichus. Rising temperatures were predicted to have low impact on the species' southern distribution limits, but to shift its northern distribution limits poleward into the high Arctic. In cold‐temperate to subarctic regions, new areas of niche overlap were predicted between F. distichus and intertidal macroalgae immigrating from the south. While climate‐change threatens intertidal seaweeds in warm‐temperate regions, seaweed meadows will likely flourish in the Arctic intertidal. Although this enriches biodiversity and opens up new seaweed‐harvesting grounds, it will also trigger unpredictable changes in the structure and functioning of the Arctic intertidal ecosystem.     

From clear lakes to murky waters – tracing the functional response of high‐latitude lake communities to concurrent ‘greening’ and ‘browning’

Climate change and the intensification of land use practices are causing widespread eutrophication of subarctic lakes. The implications of this rapid change for lake ecosystem function remain poorly understood. To assess how freshwater communities respond to such profound changes in their habitat and resource availability, we conducted a space‐for‐time analysis of food‐web structure in 30 lakes situated across a temperature‐productivity gradient equivalent to the predicted future climate of subarctic Europe (temperature +3°C, precipitation +30% and nutrient +45 μg L^?1 total phosphorus). Along this gradient, we observed an increase in the assimilation of pelagic‐derived carbon from 25 to 75% throughout primary, secondary and tertiary consumers. This shift was overwhelmingly driven by the consumption of pelagic detritus by benthic primary consumers and was not accompanied by increased pelagic foraging by higher trophic level consumers. Our data also revealed a convergence of the carbon isotope ratios of pelagic and benthic food web endmembers in the warmest, most productive lakes indicating that the incorporation of terrestrial derived carbon into aquatic food webs increases as land use intensifies. These results, reflecting changes along a gradient characteristic of the predicted future environment throughout the subarctic, indicate that climate and land use driven eutrophication and browning are radically altering the function and fuelling of aquatic food webs in this biome.     

Impacts of forestry planting on primary production in upland lakes from north‐west Ireland

Planted forests are increasing in many upland regions worldwide, but knowledge about their potential effects on algal communities of catchment lakes is relatively unknown. Here, the effects of afforestation were investigated using palaeolimnology at six upland lake sites in the north‐west of Ireland subject to different extents of forest plantation cover (4–64% of catchment area). ²¹⁰Pb‐dated sediment cores were analysed for carotenoid pigments from algae, stable isotopes of bulk carbon (δ¹³C) and nitrogen (δ¹⁵N), and C/N ratios. In lakes with >50% of their catchment area covered by plantations, there were two‐ to sixfold increases in pigments from cryptophytes (alloxanthin) and significant but lower increases (39–116%) in those from colonial cyanobacteria (canthaxanthin), but no response from biomarkers of total algal abundance (β‐carotene). In contrast, lakes in catchments with <20% afforestation exhibited no consistent response to forestry practices, although all lakes exhibited fluctuations in pigments and geochemical variables due to peat cutting and upland grazing prior to forest plantation. Taken together, patterns suggest that increases in cyanobacteria and cryptophyte abundance reflect a combination of mineral and nutrient enrichment associated with forest fertilization and organic matter influx which may have facilitated growth of mixotrophic taxa. This study demonstrates that planted forests can alter the abundance and community structure of algae in upland humic lakes of Ireland and Northern Ireland, despite long histories of prior catchment disturbance.     

Increased risk of cyanobacterial blooms in northern high‐latitude lakes through climate warming and phosphorus enrichment

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Inverse relationship of epilithic algae and pelagic phosphorus in unproductive lakes: Roles of N2 fixers and light

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Drought-induced changes in nutrient concentrations and retention in two shallow Mediterranean lakes subjected to different degrees of management

While extensive knowledge exists on the relationship between nutrient loading and nutrient concentrations in lakes in the cold temperate region, few studies have been conducted in warm lakes, not least in warm arid lakes. This is unfortunate as a larger proportion of the world’s lakes will be situated in arid climates in the future due to climate change and a larger proportion will suffer from a higher frequency of intensive drought. We conducted a comprehensive 11–13 year mass balance study in two interconnected shallow Mediterranean lakes in Turkey, covering a period with substantial changes in climate conditions. The upstream lake was only affected by natural changes in nutrient loading, while the downstream lake was additionally influenced by sewage diversion and restoration by fish removal. Contrasting to experience from north temperate lakes we found an increase in in-lake concentrations of total phosphorus and inorganic nitrogen (ammonia as well as nitrate) in dry years despite lower external nutrient loading, and submerged macrophytes did not increase the nitrogen retention capacity of the lakes. In contrast, fish removal modulated the nitrogen concentration as in north temperate lakes, but the effect was not long-lasting. Our results suggest that climate warming reduces the nutrient retention capacity of shallow lakes in the Mediterranean and exacerbates eutrophication. Lower thresholds of nutrient loading for shifting turbid shallow lakes to a clear water state are therefore to be expected in arid zones in a future warmer climate, with important management implications.     

Regional versus local drivers of water quality in the Windermere catchment,Lake District,United Kingdom: The dominant influence of wastewater pollution over the past 200 years

Freshwater ecosystems are threatened by multiple anthropogenic stressors acting over different spatial and temporal scales, resulting in toxic algal blooms, reduced water quality and hypoxia. However, while catchment characteristics act as a ‘filter’ modifying lake response to disturbance, little is known of the relative importance of different drivers and possible differentiation in the response of upland remote lakes in comparison to lowland, impacted lakes. Moreover, many studies have focussed on single lakes rather than looking at responses across a set of individual, yet connected lake basins. Here we used sedimentary algal pigments as an index of changes in primary producer assemblages over the last ~200 years in a northern temperate watershed consisting of 11 upland and lowland lakes within the Lake District, United Kingdom, to test our hypotheses about landscape drivers. Specifically, we expected that the magnitude of change in phototrophic assemblages would be greatest in lowland rather than upland lakes due to more intensive human activities in the watersheds of the former (agriculture, urbanization). Regional parameters, such as climate dynamics, would be the predominant factors regulating lake primary producers in remote upland lakes and thus, synchronize the dynamic of primary producer assemblages in these basins. We found broad support for the hypotheses pertaining to lowland sites as wastewater treatment was the main predictor of changes to primary producer assemblages in lowland lakes. In contrast, upland headwaters responded weakly to variation in atmospheric temperature, and dynamics in primary producers across upland lakes were asynchronous. Collectively, these findings show that nutrient inputs from point sources overwhelm climatic controls of algae and nuisance cyanobacteria, but highlights that large‐scale stressors do not always initiate coherent regional lake response. Furthermore, a lake's position in its landscape, its connectivity and proximity to point nutrients are important determinants of changes in production and composition of phototrophic assemblages.     

Effects of nutrient loading,temperature regime and grazing pressure on nutrient limitation of periphyton in experimental ponds

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Small changes in timing of breeding among subarctic passerines over a 32‐year period

Many bird populations in temperate regions have advanced their timing of breeding in response to a warming climate in recent decades. However, long‐term trends in temperature differ geographically and between seasons, and so do responses of local breeding populations. Data on breeding bird phenology from subarctic and arctic passerine populations are scarce, and relatively little data has been recorded in open‐nesting species. We investigated the timing of breeding and its relationship to spring temperature of 14 mainly open‐nesting passerine species in subarctic Swedish Lapland over a period of 32 years (1984–2015). We estimated timing of breeding from the progress of post‐juvenile moult in mist‐netted birds, a new method exploring the fact that the progress of post‐juvenile moult correlates with age. Although there was a numerical tendency for earlier breeding in most species (on average ?0.09 days/year), changes were statistically significant in only three species (by ?0.16 to ?0.23 days/year). These figures are relatively low compared with what has been found in other long‐term studies but are similar to a few other studies in subarctic areas. Generally, annual hatching dates were negatively correlated with mean temperature in May. This correlation was stronger in long‐distance than in short‐distance migrants. Although annual temperatures at high northern latitudes have increased over recent decades, there was no long‐term increase in mean temperature in May over the study period at this subarctic site. This is probably the main reason why there were only small long‐term changes in hatching dates.     

Expanded spatial extent of the Medieval Climate Anomaly revealed in lake‐sediment records across the boreal region in northwest Ontario

Multi‐decadal to centennial‐scale shifts in effective moisture over the past two millennia are inferred from sedimentary records from six lakes spanning a ~250 km region in northwest Ontario. This is the first regional application of a technique developed to reconstruct drought from drainage lakes (open lakes with surface outlets). This regional network of proxy drought records is based on individual within‐lake calibration models developed using diatom assemblages collected from surface sediments across a water‐depth gradient. Analysis of diatom assemblages from sediment cores collected close to the near‐shore ecological boundary between benthic and planktonic diatom taxa indicated this boundary shifted over time in all lakes. These shifts are largely dependent on climate‐driven influences, and can provide a sensitive record of past drought. Our lake‐sediment records indicate two periods of synchronous signals, suggesting a common large‐scale climate forcing. The first is a period of prolonged aridity during the Medieval Climate Anomaly (MCA, c. 900‐1400 CE). Documentation of aridity across this region expands the known spatial extent of the MCA megadrought into a region that historically has not experienced extreme droughts such as those in central and western north America. The second synchronous period is the recent signal of the past ~100 years, which indicates a change to higher effective moisture that may be related to anthropogenic forcing on climate. This approach has the potential to fill regional gaps, where many previous paleo‐lake depth methods (based on deeper centrally located cores) were relatively insensitive. By filling regional gaps, a better understanding of past spatial patterns in drought can be used to assess the sensitivity and realism of climate model projections of future climate change. This type of data is especially important for validating high spatial resolution, regional climate models.     

Climate and productivity shape fish and invertebrate community structure in subarctic lakes

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Genome evolution and host‐microbiome shifts correspond with intraspecific niche divergence within harmful algal bloom‐forming Microcystis aeruginosa

Intraspecific niche divergence is an important driver of species range, population abundance and impacts on ecosystem functions. Genetic changes are the primary focus when studying intraspecific divergence; however, the role of ecological interactions, particularly host‐microbiome symbioses, is receiving increased attention. The relative importance of these evolutionary and ecological mechanisms has seen only limited evaluation. To address this question, we used Microcystis aeruginosa, the globally distributed cyanobacterium that dominates freshwater harmful algal blooms. These blooms have been increasing in occurrence and intensity worldwide, causing major economic and ecological damages. We evaluated 46 isolates of M. aeruginosa and their microbiomes, collected from 14 lakes in Michigan, USA, that vary over 20‐fold in phosphorus levels, the primary limiting nutrient in freshwater systems. Genomes of M. aeruginosa diverged along this phosphorus gradient in genomic architecture and protein functions. Fitness in low‐phosphorus lakes corresponded with additional shifts within M. aeruginosa including genome‐wide reductions in nitrogen use, an expansion of phosphorus assimilation genes and an alternative life history strategy of nonclonal colony formation. In addition to host shifts, despite culturing in common‐garden conditions, host‐microbiomes diverged along the gradient in taxonomy, but converged in function with evidence of metabolic interdependence between the host and its microbiome. Divergence corresponded with a physiological trade‐off between fitness in low‐phosphorus environments and growth rate in phosphorus‐rich conditions. Co‐occurrence of genotypes adapted to different nutrient environments in phosphorus‐rich lakes may have critical implications for understanding how M. aeruginosa blooms persist after initial nutrient depletion. Ultimately, we demonstrate that the intertwined effects of genome evolution, host life history strategy and ecological interactions between a host and its microbiome correspond with an intraspecific niche shift with important implications for whole ecosystem function.