Erik Jeppesen

Climate change and the future of freshwater biodiversity in Europe: a primer for policy-makers.

Abstract Earths climate is changing, and by the end of the 21st century in Europe, average temperatures are likely to have risen by at least 2 °C, and more likely 4 °C with associated effects on patterns of precipitation and the frequency of extreme weather events. Attention among policy-makers is divided about how to minimise the change, how to mitigate its effects, how to maintain the natural resources on which societies depend and how to adapt human societies to the changes. Natural systems are still seen, through a long tradition of conservation management that is largely species-based, as amenable to adaptive management, and biodiversity, mostly perceived as the richness of plant and vertebrate communities, often forms a focus for planning. We argue that prediction of particular species changes will be possible only in a minority of cases but that prediction of trends in general structure and operation of four generic freshwater ecosystems (erosive rivers, depositional floodplain rivers, shallow lakes and deep lakes) in three broad zones of Europe (Mediterranean, Central and Arctic-Boreal) is practicable. Maintenance and rehabilitation of ecological structures and operations will inevitably and incidentally embrace restoration of appropriate levels of species biodiversity. Using expert judgement, based on an extensive literature, we have outlined, primarily for lay policy makers, the pristine features of these systems, their states under current human impacts, how these states are likely to alter with a warming of 2 °C to 4 °C and what might be done to mitigate this. We have avoided technical terms in the interests of communication, and although we have included full referencing as in academic papers, we have eliminated degrees of detail that could confuse broad policy-making.

Small birds, big effects: the little auk (Alle alle) transforms high Arctic ecosystems

In some arctic areas, marine-derived nutrients (MDN) resulting from fish migrations fuel freshwater and terrestrial ecosystems, increasing primary production and biodiversity. Less is known, however, about the role of seabird-MDN in shaping ecosystems. Here, we examine how the most abundant seabird in the North Atlantic, the little auk (Alle alle), alters freshwater and terrestrial ecosystems around the North Water Polynya (NOW) in Greenland. We compare stable isotope ratios (δ15N and δ13C) of freshwater and terrestrial biota, terrestrial vegetation indices and physical–chemical properties, productivity and community structure of fresh waters in catchments with and without little auk colonies. The presence of colonies profoundly alters freshwater and terrestrial ecosystems by providing nutrients and massively enhancing primary production. Based on elevated δ15N in MDN, we estimate that MDN fuels more than 85% of terrestrial and aquatic biomass in bird influenced systems. Furthermore, by using different proxies of bird impact (colony distance, algal δ15N) it is possible to identify a gradient in ecosystem response to increasing bird impact. Little auk impact acidifies the freshwater systems, reducing taxonomic richness of macroinvertebrates and truncating food webs. These results demonstrate that the little auk acts as an ecosystem engineer, transforming ecosystems across a vast region of Northwest Greenland.

The influence of nutrient loading, climate and water depth on nitrogen and phosphorus loss in shallow lakes: a pan-European mesocosm experiment

Losses of phosphorus (P) and nitrogen (N) have important influences on in-lake concentrations and nutrient loading to downstream ecosystems. We performed a series of mesocosm experiments along a latitudinal gradient from Sweden to Greece to investigate the factors influencing N and P loss under different climatic conditions. In six countries, a standardised mesocosm experiment with two water depths and two nutrient levels was conducted concurrently between May and November 2011. Our results showed external nutrient loading to be of key importance for N and P loss in all countries. Almost all dissolved inorganic nitrogen (DIN) and soluble reactive phosphorus (SRP) were lost or taken up in biomass in all mesocosms. We found no consistent effect of temperature on DIN and SRP loss but a significant, though weak, negative effect of temperature on total nitrogen (TN) and total phosphorus (TP) loss in the deeper mesocosms, probably related to higher organic N and P accumulation in the water in the warmer countries. In shallow mesocosms, a positive trend in TN and TP loss with increasing temperature was observed, most likely related to macrophyte growth.

Successful restoration of a tropical shallow eutrophic lake: Strong bottom-up but weak top-down effects recorded

Fish manipulation has been used to restore lakes in the temperate zone. Often strong short-term cascading effects have been obtained, but the long term-perspectives are less clear. Fish manipulation methods are far less advanced for warm lakes, and it is debatable whether it is, in fact, possible to create a trophic cascade in warm lakes due to the dominance and high densities of fast-reproducing omnivorous fish. However, removal of benthic feeding fish also reduce disturbance of the sediment, which not only affects the nutrient level but also the concentration of suspended organic and inorganic matter with enhanced water clarity and potentially better growth conditions for submerged macrophytes. We conducted a biomanipulation experiment in one of the basins in Chinese Huizhou West Lake that have remained highly turbid after extensive nutrient loading reduction. Another basin was used as control (control-treatment pairing design). Removal of a substantial amount of plankti-benthivorous fish was followed by planting of submerged macrophytes and stocking of piscivorous fish. We found strong and relatively long-lasting effects of the restoration initiative in the form of substantial improvements in water clarity and major reductions in nutrient concentrations, particularly total phosphorus, phytoplankton and turbidity, while only minor effects were detected for crustacean zooplankton grazers occurring in low densities before as well as after the restoration. Our results add importantly to the existing knowledge of restoration of warm lakes and are strongly relevant, not least in Asia where natural lakes frequently are used extensively for fish production, often involving massive stocking of benthivorous fish. With a growing economy and development of more efficient fish production systems, the interest in restoring lakes is increasing world-wide. We found convincing evidence that fish removal and piscivores stocking combined with transplantation of submerged macrophytes may have significant effects on water clarity in warm shallow lakes even if the zooplankton grazing potential remains low, the latter most likely as a result of high predation on the zooplankton.

Nutrient availability limits biological production in Arctic sea ice melt ponds

Every spring and summer melt ponds form at the surface of polar sea ice and become habitats where biological production may take place. Previous studies report a large variability in the productivity, but the causes are unknown. We investigated if nutrients limit the productivity in these first-year ice melt ponds by adding nutrients to three enclosures ([1] PO43−, [2] NO3−, and [3] PO43− and NO3−) and one natural melt pond (PO43− and NO3−), while one enclosure and one natural melt pond acted as controls. After 7–13 days, Chl a concentrations and cumulative primary production were between two- and tenfold higher in the enclosures and natural melt ponds with nutrient addition compared with their respective controls, with the largest increase occurring in the enclosures. Separate additions of PO43− and NO3− in the enclosures led to intermediate increases in productivity, suggesting co-limitation of nutrients. Bacterial production and the biovolume of ciliates, which were the dominant grazers, were positively correlated with primary production, showing a tight coupling between primary production and both microbial activity and ciliate grazing. To our knowledge, this study is the first to ascertain nutrient limitation in melt ponds. We also document that the addition of nutrients, although at relative high concentrations, can stimulate biological productivity at several trophic levels. Given the projected increase in first-year ice, increased melt pond coverage during the Arctic spring and potential additional nutrient supply from, e.g. terrestrial sources imply that biological activity of melt ponds may become increasingly important for the sympagic carbon cycling in the future Arctic.

Ecological resilience in lakes and the conjunction fallacy.

There is a pressing need to apply stability and resilience theory to environmental management to restore degraded ecosystems effectively and to mitigate the effects of impending environmental change. Lakes represent excellent model case studies in this respect and have been used widely to demonstrate theories of ecological stability and resilience that are needed to underpin preventative management approaches. However, we argue that this approach is not yet fully developed because the pursuit of empirical evidence to underpin such theoretically grounded management continues in the absence of an objective probability framework. This has blurred the lines between intuitive logic (based on the elementary principles of probability) and extensional logic (based on assumption and belief) in this field.Lake ecosystems have provided much of the empirical evidence for ecological resilience theory. Here, a more rigorous logical approach is called for when translating this research into management decisions.

Impact of alternating wet and dry periods on long-term seasonal phosphorus and nitrogen budgets of two shallow Mediterranean lakes

The water balance, with large seasonal and annual water level fluctuations, has a critical influence on the nitrogen and phosphorus dynamics of shallow lakes in the semi-arid climate zone. We constructed seasonal water and nutrient budgets for two connected shallow lakes, Lakes Mogan and Eymir, located in Central Anatolia, Turkey. The study period covered 20years with alternations between dry and wet years as well as restoration efforts including sewage effluent diversion and biomanipulations in Lake Eymir. Both lakes experienced a 1-2m water level drop during a drought period and a subsequent increase during the wet period, with seasonal water level fluctuations of 0.60 to 0.70m. During wet years with high water levels, small seasonal differences were observed with a nutrient peak in spring caused by external loading and nutrient loss via retention during summer. During years with low water levels, nutrient concentrations increased due to internal and external loading, exacerbated by evaporative water loss. In Lake Eymir, a shift to eutrophic conditions with turbid water occurred under low water level conditions and consequent internal loading of P from the sediment, causing high nutrient concentrations in summer. Our results indicate a threat of lakes drying out in the semi-arid climate zone if evaporation increases and precipitation decreases as anticipated from the global climate change predictions. In addition, our results show the influence of the water balance on the eutrophication of shallow lakes in the Mediterranean climate zone and highlight the ultimate consequences for lake management.

Environment not dispersal limitation drives clonal composition of Arctic Daphnia in a recently deglaciated area

One of the most prominent manifestations of the ongoing climate warming is the retreat of glaciers and ice sheets around the world. Retreating glaciers result in the formation of new ponds and lakes, which are available for colonization. The gradual appearance of these new habitat patches allows us to determine to what extent the composition of asexual Daphnia (water flea) populations is affected by environmental drivers vs. dispersal limitation. Here, we used a landscape genetics approach to assess the processes structuring the clonal composition of species in the D. pulex species complex that have colonized periglacial habitats created by ice‐sheet retreat in western Greenland. We analysed 61 populations from a young (<50 years) and an old cluster (>150 years) of lakes and ponds. We identified 42 asexual clones that varied widely in spatial distribution. Beta‐diversity was higher among older than among younger systems. Lineage sorting by the environment explained 14% of the variation in clonal composition whereas the pure effect of geographical distance was very small and statistically insignificant ( Radj2 = 0.010, P = 0.085). Dispersal limitation did not seem important, even among young habitat patches. The observation of several tens of clones colonizing the area combined with environmentally driven clonal composition of populations illustrates that population assembly of asexual species in the Arctic is structured by environmental gradients reflecting differences in the ecology of clones.

Synergistic negative effects of small-sized benthivorous fish and nitrogen loading on the growth of submerged macrophytes – Relevance for shallow lake restoration

Rapid recruitment of small fish after biomanipulation in warm lakes may delay the reestablishment of submerged macrophytes, not least at high nutrient concentrations. Success has recently been obtained in controlling phosphorus (P) loading to many lakes, but nitrogen (N) inputs often remain high. To determine the interactive effects of N loading and the abundance of small-sized fish on the growth of the submerged macrophyte Vallisneria natans, we conducted an outdoor mesocosm experiment with a factorial design on the north shore of Lake Taihu, China. The experiment involved two densities of small crucian carp - low (10gm-2) and high (40gm-2) - crossed with two levels of N loading - present-day external nutrient loading (P: 5μgL-1day-1, N: 130μgL-1day-1) and P: 5μgL-1day-1 with a three times higher N loading (N: 390μgL-1day-1). The results showed that nitrogen-fish interactions significantly hindered the growth of V. natans, particularly at the high N loading. At low N loading, high densities of fish decreased the relative growth rate, mean leaf length, leaf mass and root mass of V. natans by 16%, 5%, 8%, and 23%, respectively, compared with these measures at low fish densities. The effect of fish was even stronger when N loading was high, with decreases of 232%, 32%, 57%, and 47% for the respective plant growth measures. The stronger effect at high N loading was attributed to higher turbidity due to enhanced phytoplankton biomass and to increased consumption or damage of plants by the fish in response to the more nutrient-enriched plant tissue. Our results indicate that high abundance of small crucian carp in warm lakes may reduce the resilience of submerged macrophytes to external N loading, thereby lowering the chances of successful restoration by biomanipulation.

Does turbidity induced by Carassius carassius limit phytoplankton growth? A mesocosm study

It is well established that benthivorous fish in shallow lakes can create turbid conditions that influence phytoplankton growth both positively, as a result of elevated nutrient concentration in the water column, and negatively, due to increased attenuation of light. The net effect depends upon the degree of turbidity induced by the benthivores. Stocked Carassius carassius dominate the benthivorous fish fauna in many nutrient-rich Chinese subtropical and tropical shallow lakes, but the role of the species as a potential limiting factor in phytoplankton growth is ambiguous. Clarification of this relationship will help determine the management strategy and cost of restoring eutrophic lakes in China and elsewhere. Our outdoor mesocosm experiment simulating the effect of high density of crucian carp on phytoplankton growth and community structure in eutrophic shallow lakes suggests that stocking with this species causes resuspension of sediment, thereby increasing light attenuation and elevating nutrient concentrations. However, the effect of light attenuation was insufficient to offset the impact of nutrient enhancement on phytoplankton growth, and significant increases in both phytoplankton biomass and chlorophyll a concentrations were recorded. Crucian carp stocking favored the dominance of diatoms and led to lower percentages (but not biomass) of buoyant cyanobacteria. The dominance of diatoms may be attributed to a competitive advantage of algal cells with high sedimentation velocity in an environment subjected to frequent crucian carp-induced resuspension and entrainment of benthic algae caused by the fish foraging activities. Our study demonstrates that turbidity induced by stocked crucian carp does not limit phytoplankton growth in eutrophic waters. Thus, removal of this species (and presumably other similar taxa) from subtropical or tropical shallow lakes, or suspension of aquaculture, is unlikely to boost phytoplankton growth, despite the resulting improvements in light availability.