Climate-associated variation in the drivers of benthic macroinvertebrate species-area relationships across shallow freshwater lakes.

The island species-area relationship (ISAR) describes how species richness increases with increasing area of a given island or island-like habitat, such as freshwater lakes. While the ISAR is one of the most common phenomena observed in ecology, there is variation in both the form of the relationship and its underlying mechanisms. We compiled a global data set of benthic macroinvertebrates from 524 shallow freshwater lakes, ranging from 1 to 293,300 ha in area. We used individual-based rarefaction to determine the degree to which ISAR was influenced by mechanisms other than passive sampling (larger islands passively sample more individuals from the regional pool and, therefore, have more species than smaller islands), which would bias results away from expected relationships between rarefied species richness (and other measures that capture relative abundances) and lake area. We also examined how climate may alter the shape of the ISARs. We found that both rarefied species richness (the number of species standardized by area or number of individuals) and a measure of evenness emphasizing common species exhibit shallow slopes in relationships with lake area, suggesting that the expected ISARs in these lakes most likely result from passive sampling. While there was considerable variation among ISARs across the investigated lakes, we found an overall positive rarefied ISAR for lakes in warm (i.e. tropical/subtropical) regions (n = 195), and in contrast, an overall negative rarefied ISAR in cool (i.e. north temperate) lakes (n = 329). This suggested that mechanisms beyond passive sampling (e.g. colonization-extinction dynamics and/or heterogeneity) were more likely to operate in warm lakes. One possible reason for this difference is that the area-dependent intensity of fish predation, which can lead to flatter ISARs, is weaker in warmer relative to cooler lakes. Our study illustrates the importance of understanding both the pattern and potential processes underlying the ISARs of freshwater lakes in different climatic regions. Furthermore, it provides a baseline for understanding how further changes to the ecosystem (i.e. in lake area or climate) might influence biodiversity patterns.

Predicting ecosystem state changes in shallow lakes using an aquatic ecosystem model: Lake Hinge, Denmark, an example.

In recent years, considerable efforts have been made to restore turbid, phytoplankton-dominated shallow lakes to a clear-water state with high coverage of submerged macrophytes. Various dynamic lake models with simplified physical representations of vertical gradients, such as PCLake, have been used to predict external nutrient load thresholds for such nonlinear regime shifts. However, recent observational studies have questioned the concept of regime shifts by emphasizing that gradual changes are more common than sudden shifts. We investigated if regime shifts would be more gradual if the models account for depth-dependent heterogeneity of the system by including the possibility of vertical gradients in the water column and sediment layers for the entire depth. Hence, bifurcation analysis was undertaken using the 1D hydrodynamic model GOTM, accounting for vertical gradients, coupled to the aquatic ecosystem model PCLake, which is implemented in the framework for aquatic biogeochemical modeling (FABM). First, the model was calibrated and validated against a comprehensive data set covering two consecutive 7-yr periods from Lake Hinge, a shallow, eutrophic Danish lake. The autocalibration program Auto-Calibration Python (ACPy) was applied to achieve a more comprehensive adjustment of model parameters. The model simulations showed excellent agreement with observed data for water temperature, total nitrogen, and nitrate and good agreement for ammonium, total phosphorus, phosphate, and chlorophyll a concentrations. Zooplankton and macrophyte coverage were adequately simulated for the purpose of this study, and in general the GOTM-FABM-PCLake model simulations performed well compared with other model studies. In contrast to previous model studies ignoring depth heterogeneity, our bifurcation analysis revealed that the spatial extent and depth limitation of macrophytes as well as phytoplankton chlorophyll-a responded more gradually over time to a reduction in the external phosphorus load, albeit some hysteresis effects still appeared. In a management perspective, our study emphasizes the need to include depth heterogeneity in the model structure to more correctly determine at which external nutrient load a given lake changes ecosystem state to a clear-water condition.

Temperature effects on fish production across a natural thermal gradient.

Global warming is widely predicted to reduce the biomass production of top predators, or even result in species loss. Several exceptions to this expectation have been identified, however, and it is vital that we understand the underlying mechanisms if we are to improve our ability to predict future trends. Here, we used a natural warming experiment in Iceland and quantitative theoretical predictions to investigate the success of brown trout as top predators across a stream temperature gradient (4-25 °C). Brown trout are at the northern limit of their geographic distribution in this system, with ambient stream temperatures below their optimum for maximal growth, and above it in the warmest streams. A five-month mark-recapture study revealed that population abundance, biomass, growth rate, and production of trout all increased with stream temperature. We identified two mechanisms that contributed to these responses: (1) trout became more selective in their diet as stream temperature increased, feeding higher in the food web and increasing in trophic position; and (2) trophic transfer through the food web was more efficient in the warmer streams. We found little evidence to support a third potential mechanism: that external subsidies would play a more important role in the diet of trout with increasing stream temperature. Resource availability was also amplified through the trophic levels with warming, as predicted by metabolic theory in nutrient-replete systems. These results highlight circumstances in which top predators can thrive in warmer environments and contribute to our knowledge of warming impacts on natural communities and ecosystem functioning.

Bimodality and alternative equilibria do not help explain long-term patterns in shallow lake chlorophyll-a.

Since its inception, the theory of alternative equilibria in shallow lakes has evolved and been applied to an ever wider range of ecological and socioecological systems. The theory posits the existence of two alternative stable states or equilibria, which in shallow lakes are characterised by either clear water with abundant plants or turbid water where phytoplankton dominate. Here, we used data simulations and real-world data sets from Denmark and north-eastern USA (902 lakes in total) to examine the relationship between shallow lake phytoplankton biomass (chlorophyll-a) and nutrient concentrations across a range of timescales. The data simulations demonstrated that three diagnostic tests could reliably identify the presence or absence of alternative equilibria. The real-world data accorded with data simulations where alternative equilibria were absent. Crucially, it was only as the temporal scale of observation increased (>3 years) that a predictable linear relationship between nutrient concentration and chlorophyll-a was evident. Thus, when a longer term perspective is taken, the notion of alternative equilibria is not required to explain the response of chlorophyll-a to nutrient enrichment which questions the utility of the theory for explaining shallow lake response to, and recovery from, eutrophication.

Gravel pit lakes in Denmark: Chemical and biological state.

Mining of gravel and sand for construction purposes is big business and gravel pit lakes have become increasingly common all over the world. In Denmark, hundreds of gravel pit lakes have been created during the past decades. We investigated the chemical and biological status of 33-52 gravel pit lakes and compared the results with data from similar-sized natural Danish lakes. The area of the lakes ranged from 0.2 to 13ha and their age from 0.5 to 26years. Generally, the gravel pit lakes were clear with low nutrient concentrations, the median concentrations of total phosphorus and total nitrogen being 0.023mg/l and 0.30mg/l compared with 0.115mg/l and 1.29mg/l, respectively, in natural lakes. Correspondingly, median chlorophyll a was 5µg/l in the gravel pit lakes and 36µg/l in the natural lakes. Submerged macrophytes were found in all gravel pit lakes, with particularly high cover in the shallow ones. Most gravel pit lakes were deeper than the natural lakes, which may restrict the area potentially to be covered by submerged macrophytes, with implications also for the biological quality of the lakes. Fish were found in most of the gravel pit lakes, roach (Rutilus rutilus), perch (Perca fluviatilis) and rudd (Scardinius erythrophalmus) being the most frequently observed species. Fish stocking was common and included also non-native species such as carp (Cyprinus carpio) and rainbow trout (Oncorchynchus mykiss). Compared with the natural lakes, fish species richness and catch per gillnet were overall lower in the gravel pit lakes. Groundwater-fed gravel pit lakes add importantly to the number of high-quality lakes in Denmark and with an optimised design and by avoiding negative side effects, they can be positive for both nature and society.

Influence of LAS on marine calanoid copepod population dynamics and potential reproduction.

The toxicity of linear alkyl benzene sulfonate (LAS) to marine invertebrates is well documented under laboratory conditions using single-species tests. It is less known how LAS affects natural populations of aquatic organisms. We hypothesised that LAS was more toxic to the calanoid copepod Acartia sp. under natural conditions than Acartia tonsa under cultured conditions in the laboratory. This hypothesis was checked by a direct comparison of LAS toxicity in single-species and model (mesocosm) studies. The acute and sublethal effects of LAS on the survival and egg production of laboratory reared A. tonsa were examined by standard test, i.e. incubation with LAS without food for 24-72 h. The LC(50) and EC(50) values averaged 1.23 and 0.74 mg l(-1) for survival and egg production, respectively. These values are comparable to previous reports. The effects of LAS on a natural copepod community were also investigated under in situ conditions. A series of seven mesocosms (holding approx. 3 m(3) of seawater each) was established with two mesocosms being controls without LAS and five mesocosms with increasing concentrations of LAS ranging from 0.1 to 6.5 mg l(-1) applied as a single dose. The indigenous copepod community, dominated by Acartia sp. and Centropages sp., responded clearly to LAS concentrations above 0.1 mg l(-1). The calculated no effect value was 0.14 mg LAS l(-1) (95% CI=0.08-1.82 mg LAS l(-1)) for the entire copepod community including all development stages after 24 h exposure. The increased sensitivity under in situ conditions was probably promoted by the suboptimal growth conditions, e.g. no saturated food concentration or inadequate nutritive values of the food. The amount of food expressed as chlorophyll concentration was low (around 2 microg chl. a l(-1)) but was not affected by LAS. It appeared that the naupliar stages of Acartia and Centropages were the least affected by LAS and that new cohorts were able to develop 15 days after the dosing with LAS.

Planktonic ciliate community structure in shallow lakes of lowland Western Europe.

Temperate shallow meso- to eutrophic lakes can exist in one of two alternative states with contrasting foodwebs, referred to as the clear-water and the turbid state. We describe the planktonic ciliate communities of such lakes based on a survey of 66 northwestern European lakes. Ciliates were enumerated and identified to species level according to the quantitative protargol staining technique. Ciliate biomass was on average twice as high in the turbid than in the clear-water lakes. The ciliate communities were dominated by oligotrichs and protostomatids, and no differences in functional composition or α-diversity could be detected between turbid and clear-water lakes, although ß-diversity tended to be higher in the latter. At the species level, however, community structure strongly differed between turbid and clear-water lakes, and several indicator species could be identified for the different lake categories. Variation partitioning showed that nutrient status did not explain ciliate community structure independent of the alternative states, while lake area was identified as an additional structuring factor for the ciliate communities. These results stress the importance of the ecosystem structure in shaping ciliate communities in temperate shallow lakes and suggest that nutrient status has little direct effect on ciliate community structure in such lakes.