In many lakes, the most conspicuous seasonal events are the phytoplankton spring bloom and the subsequent clear-water phase, a period of low-phytoplankton biomass that is frequently caused by mesozooplankton (Daphnia) grazing. In Central European lakes, the timing of the clear-water phase is linked to large-scale climatic forcing, with warmer winters being followed by an earlier onset of the clear-water phase. Mild winters may favour an early build-up of Daphnia populations, both directly through increased surface temperatures and indirectly by reducing light limitation and enhancing algal production, all being a consequence of earlier thermal stratification. We conducted a field experiment to disentangle the separate impacts of stratification depth (affecting light supply) and temperature on the magnitude and timing of successional events in the plankton. We followed the dynamics of the phytoplankton spring bloom, the clear-water phase and the spring peak in Daphnia abundance in response to our experimental manipulations. Deeper mixing delayed the timing of all spring seasonal events and reduced the magnitudes of the phytoplankton bloom and the subsequent Daphnia peak. Colder temperatures retarded the timing of the clear-water phase and the subsequent Daphnia peak, whereas the timing of the phytoplankton peak was unrelated to temperature. Most effects of mixing depth (light) and temperature manipulations were independent, effects of mixing depth being more prevalent than effects of temperature. Because mixing depth governs both the light climate and the temperature regime in the mixed surface layer, we propose that climate-driven changes in the timing and depth of water column stratification may have far-reaching consequences for plankton dynamics and should receive increased attention.

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In deep temperate lakes, the beginning of the growing season is triggered by thermal stratification, which alleviates light limitation of planktonic producers in the surface layer and prevents heat loss to deeper strata. The sequence of subsequent phenological events (phytoplankton spring bloom, grazer peak, clearwater phase) results in part from coupled phytoplankton-grazer interactions. Disentangling the separate, direct effects of correlated climatic drivers (stratificationdependent underwater light climate vs. water temperature) from their indirect effects mediated through trophic feedbacks is impossible using observational field data, which challenges our understanding of global warming effects on seasonal plankton dynamics. We therefore manipulated water temperature and stratification depth independently in experimental field mesocosms containing ambient microplankton and inocula of the resident grazer Daphnia hyalina. Higher light availability in shallower surface layers accelerated primary production, warming accelerated consumption and growth of Daphnia, and both factors speeded up successional dynamics driven by trophic feedbacks. Specifically, phytoplankton peaked and decreased earlier and Daphnia populations increased and peaked earlier at both shallower stratification and higher temperature. The timing of ciliate dynamics was unrelated to both factors. Volumetric peak densities of phytoplankton, ciliates and Daphnia in the surface layer were also unaffected by temperature but declined with stratification depth in parallel with light availability. The latter relationship vanished, however, when population sizes were integrated over the entire water column. Overall our results suggest that, integrated over the entire water column of a deep lake, surface warming and shallower stratification independently speed up spring successional events, whereas the magnitudes of phytoplankton and zooplankton spring peaks are less sensitive to these factors. Therefore, accelerated dynamics under warming need not lead to a trophic mismatch (given similar grazer inocula at the time of stratification). We emphasize that entire water column dynamics must be studied to estimate global warming effects on lake ecosystems.

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[1] To settle debate on the timing of sea level fluctuations during marine isotope stage (MIS) 3, we present records of d 18 O ruber (sea level proxy) and magnetic susceptibility from the same samples within the single sediment archive (i.e., ''coregistered'') of central Red Sea core GeoTü-KL11. Core-scanning X-ray fluorescence and environmental magnetic data establish the suitability of magnetic susceptibility as a proxy for eolian dust content in Red Sea sediments. The eolian dust data record similar variability as Greenland d 18 O ice during early to middle MIS 3, in agreement with previous observations that regional Arabian Sea climate fluctuated with a timing similar to that of Greenland climate variations. In contrast, the sea level record fluctuates with a timing similar to that of Antarctic-style climate variations. The coregistered nature of the two records in core KL11 unambiguously reveals a distinct offset in the phase relationship between sea level and eolian dust fluctuations. The results confirm that sea level rises, indicated by shifts in Red Sea d 18 O ruber to lighter values, occurred during cold episodes in Greenland during early to middle MIS 3. This indicates that the amplitudes of the reconstructed MIS 3 sea level fluctuations would not be reduced by inclusion of regional climate fluctuations in the Red Sea sea level method. Our analysis comprehensively supports our earlier conclusions of large-amplitude sea level variations during MIS 3 with a timing that is strongly similar to Antarctic-style climate variations.

[1] We used an oceanic general circulation model to evaluate the sensitivity of the hydrography and circulation of the Red Sea in response to reduced sea level and modified atmospheric conditions during the Holocene. With Holocene sea level close to the modern level, the Red Sea was sensitive to changes in atmospheric conditions, and it only shows a relatively mild response to sea level change. Changes in the monsoon system influence the exchange flow through the Strait of Bab el Mandab, the meridional overturning circulation of the Red Sea, and its hydrography. Forced by humid conditions the (modeled) Red Sea temperature increased by ∼1.5°C, while when arid conditions were imposed, the temperature decreased by ∼2.5°C. Similar heating and cooling events during the early and late Holocene are seen in a sea surface temperature record from the northern Red Sea (derived from the temperature sensitive TEX 86 molecular biomarker), which suggests that humid conditions prevailed during the early Holocene and more arid conditions prevailed during the late Holocene. The gradual decline in Red Sea temperature between these two time periods suggests a gradual decline in the summer monsoon strength. This monsoon trend and the resulting changes in the Red Sea circulation are supported by the distribution of crenarchaea fossil lipids in Red Sea sediments from this period. Monsoondriven changes in the exchange flow through the Strait of Bab el Mandab affected the crenarchaea population structure, and therefore, their molecular fossil distribution in the sediments of the Red Sea potentially provides an index for the summer monsoon strength during the Holocene.Citation: Biton, E., H. Gildor, G. Trommer, M. Siccha, M. Kucera, M. T. J. van der Meer, and S. Schouten (2010), Sensitivity of Red Sea circulation to monsoonal variability during the Holocene: An integrated data and modeling study, Paleoceanography, 25, PA4209,

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[1] In order to assess how insolation-driven climate change superimposed on sea level rise and millennial events influenced the Red Sea during the Holocene, we present new paleoceanographic records from two sediment cores to develop a comprehensive reconstruction of Holocene circulation dynamics in the basin. We show that the recovery of the planktonic foraminiferal fauna after the Younger Dryas was completed earlier in the northern than in the central Red Sea, implying significant changes in the hydrological balance of the northern Red Sea region during the deglaciation. In the early part of the Holocene, the environment of the Red Sea closely followed the development of the Indian summer monsoon and was dominated by a circulation mode similar to the current summer circulation, with low productivity throughout the central and northern Red Sea. The climatic signal during the late Holocene is dominated by a faunal transient event centered around 2.4 ka BP. Its timing corresponds to that of North Atlantic Bond event 2 and to a widespread regionally recorded dry period. This faunal transient is characterized by a more productive foraminiferal fauna and can be explained by an intensification of the winter circulation mode and high evaporation. The modern distribution pattern of planktonic foraminifera, reflecting the prevailing circulation system, was established after 1.7 ka BP.