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Aim: Ecological communities are structured through the interplay of deterministic assembly processes such as competition and environmental filtering. Whereas the drivers of spatial community structure are frequently studied in extant communities, little is known about the relative importance of assembly processes in response to environmental factors over evolutionary time-scales. Here, we use an integrative framework to unravel community assembly processes since the inception of a long-lived lake ecosystem. Location: Lake Ohrid. Time period: From lake formation 1.36 million years ago to the present. Major taxa studied: Planktonic diatoms. Methods: We constructed a dated phylogeny of extant and extinct diatoms and collected trait data for 380 fossil communities to quantify phylogenetic community structure and functional richness and to determine the relative importance of deterministic assembly processes over time. We then used regression analysis to correlate the phylogenetic community structure with palaeoenvironmental and intrinsic biological predictors and to identify primary drivers of assembly processes. Results: Our results suggest a dense packing of niche space with higher species richness and co-occurrence of closely related species. There are only two short episodes in the very recent past dominated by distantly related taxa. We found distinct changes in phylogenetic community structure upon speciation or extinction events and an increase in mean community relatedness over time. Main conclusions: Our finding of closely related co-occurring species implies environmental filtering as the primary assembly mechanism, with a minor but increasingly important role of competition towards the present, driven by evolutionary dynamics. Such an increase in the relative contribution of competition to the assembly of communities in relation to the aging of an insular ecosystem, together with a denser packing of morphospace in the early phase of system ontogeny is compatible with ecological predictions according to the theory of island biogeography.

Phytoplankton photosynthesis strongly relies on the operation of carbon‐concentrating mechanisms (CCMs) to accumulate CO2 around their carboxylating enzyme ribulose‐1,5‐bisphosphate carboxylase/oxygenase (RuBisCO). Earlier evolved phytoplankton groups were shown to exhibit higher CCM activities to compensate for their RuBisCO with low CO2 specificities. Here, we tested whether earlier evolved phytoplankton groups also exhibit a higher CCM plasticity. To this end, we collected data from literature and applied a Bayesian linear meta‐analytic model. Our results show that with elevated pCO2, photosynthetic CO2 affinities decreased strongest and most consistent for the earlier evolved groups, i.e., cyanobacteria and dinoflagellates, while CO2‐dependent changes in affinities for haptophytes and diatoms were smaller and less consistent. In addition, responses of maximum photosynthetic rates toward elevated pCO2 were generally small and inconsistent across species. Our results demonstrate that phytoplankton groups with an earlier origin possess a high CCM plasticity, whereas more recently evolved groups do not, which likely results from evolved differences in the CO2 specificity of RuBisCO.

Dissolved oceanic CO

Elevated pCO2 and warming may promote algal growth and toxin production, and thereby possibly support the proliferation and toxicity of harmful algal blooms (HABs). Here, we tested whether empirical data support this hypothesis using a meta‐analytic approach and investigated the responses of growth rate and toxin content or toxicity of numerous marine and estuarine HAB species to elevated pCO2 and warming. Most of the available data on HAB responses towards the two tested climate change variables concern dinoflagellates, as many members of this phytoplankton group are known to cause HAB outbreaks. Toxin content and toxicity did not reveal a consistent response towards both tested climate change variables, while growth rate increased consistently with elevated pCO2. Warming also led to higher growth rates, but only for species isolated at higher latitudes. The observed gradient in temperature growth responses shows the potential for enhanced development of HABs at higher latitudes. Increases in growth rates with more CO2 may present an additional competitive advantage for HAB species, particularly as CO2 was not shown to enhance growth rate of other non‐HAB phytoplankton species. However, this may also be related to the difference in representation of dinoflagellate and diatom species in the respective HAB and non‐HAB phytoplankton groups. Since the proliferation of HAB species may strongly depend on their growth rates, our results warn for a greater potential of dinoflagellate HAB development in future coastal waters, particularly in temperate regions., Elevated pCO2 and warming may promote algal growth and toxin production, and thereby possibly support the proliferation and toxicity of HABs. Using a meta‐analytic approach we found that elevated pCO2 increased growth rates of dinoflagellate HAB species, while this was not the case for non‐HAB phytoplankton species. Warming also led to higher growth rates, but mainly for species isolated at higher latitudes. These results warn for a greater potential of dinoflagellate HAB development in future coastal waters, particularly in temperate regions.

Global climate change involves an increase in oceanic CO2 concentrations as well as thermal stratification of the water column, thereby reducing nutrient supply from deep to surface waters. Changes in inorganic carbon (C) or nitrogen (N) availability have been shown to affect marine primary production, yet little is known about their interactive effects. To test for these effects, we conducted continuous culture experiments under N limitation and exposed the bloom- forming dinoflagellate species Scrippsiella trochoidea and Alexandrium fundyense (formerly A. tamarense) to CO2 partial pressures (pCO2) ranging between 250 and 1000 μatm. Ratios of partic- ulate organic carbon (POC) to organic nitrogen (PON) were elevated under N limitation, but also showed a decreasing trend with increasing pCO2. PON production rates were highest and affini- ties for dissolved inorganic N were lowest under elevated pCO2, and our data thus demonstrate a CO2-dependent trade-off in N assimilation. In A. fundyense, quotas of paralytic shellfish poison- ing toxins were lowered under N limitation, but the offset to those obtained under N-replete con- ditions became smaller with increasing pCO2. Consequently, cellular toxicity under N limitation was highest under elevated pCO2. All in all, our observations imply reduced N stress under ele- vated pCO2, which we attribute to a reallocation of energy from C to N assimilation as a conse- quence of lowered costs in C acquisition. Such interactive effects of ocean acidification and nutri- ent limitation may favor species with adjustable carbon concentrating mechanisms and have consequences for their competitive success in a future ocean.

Alexandrium ostenfeldii is among the most intensely studied marine planktonic dinophytes and in the last few years blooms have become a recurrent phenomenon mainly in brackish coastal waters. Since 2012, A. ostenfeldii recurs annually in the Ouwerkerkse Kreek, a Dutch brackish water creek discharging into an estuary with large stocks of mussels, oysters and cockles. The creek is characterized by highly dynamic abiotic conditions, notably salinity. Here, we investigated the impacts of salinities ranging from 3 to 34 on growth and toxin content of an A. ostenfeldii isolate from the creek. Our results demonstrate a broad salinity tolerance of the Dutch A. ostenfeldii population, with growth rates from 0.13 to 0.2 d−1 over a salinity range from 6 to 34. Highest paralytic shellfish toxin and cyclic imine toxin cell quotas were observed for the lowest and highest salinities, and were associated with increases in cell size. Lytic activity was highest at the lowest salinity, and was 5-fold higher in the cell-free supernatants compared to cell extracts. Together our results demonstrate a tight coupling between salinity and A. ostenfeldii growth rate, cell size and toxin synthesis, which may have consequences for the seasonal dynamics of bloom toxicity.