Dispersal decreases diversity in heterogeneous metacommunities by enhancing regional competition

Experiments and models reveal that moderate dispersal rates between local communities can increase diversity by alleviating local competitive exclusion; in contrast, high dispersal rates can decrease diversity by amplifying regional competition. However, hitherto experimental tests on how dispersal affects diversity in the presence and absence of environmental heterogeneity are largely missing, although it is known that environmental heterogeneity influences diversity. For the first time we experimentally show that the interaction between dispersal rate and the presence of an environmental gradient with on-average lower resource availability than the homogeneous control treatment affects diversity. In metacommunities of nine co-occurring species of marine benthic microalgae we factorially manipulated dispersal rate and the presence and absence of a light intensity gradient across local patches to test effects on local, regional, and beta diversity and to compare results to predictions from monoculture experiments. Although species in this experiment did not show resource partitioning along the light gradient as assumed by source–sink models, dispersal limitation maintained diversity in metacommunities with light gradients but not without. Local diversity and evenness were high under low light intensities when dispersal was limited and decreased with both increasing light intensities and dispersal rates. These diversity changes can be explained by the reduction of growth of the regional superior competitor at low light intensities alleviating its competitive strength. Increasing dispersal rate in turn compensated for the superior competitor's slow growth in those local patches with rather unfavorable light conditions and thus led to decreasing diversity and evenness. In contrast, diversity in the metacommunities without a light gradient was constantly low. Here, the superior competitor contributed 90% to total community biomass in all patches. High dominance, however, likely resulted from on-average higher resource availability (i.e., higher light intensities) compared to metacommunities with light gradient and not from patch homogeneity in itself

Technical Note: Precise quantitative measurements of total dissolved inorganic carbon from small amounts of seawater using a gas chromatographic system

Total dissolved inorganic carbon (CT) is one of the most frequently measured parameters used to calculate the partial pressure of carbon dioxide in seawater. Its determination has become increasingly important because of the rising interest in the biological effects of ocean acidification. Coulometric and infrared detection methods are currently favored in order to precisely quantify CT. These methods however are not sufficiently validated for CT measurements of biological experiments manipulating seawater carbonate chemistry with an extended CT measurement range (~1250–2400 μmol kg–1) compared to natural open ocean seawater (~1950–2200 μmol kg−1). The requirement of total sample amounts between 0.1–1 L seawater in the coulometric- and infrared detection methods potentially exclude their use for experiments working with much smaller volumes. Additionally, precise CT analytics become difficult with high amounts of biomass (e.g., phytoplankton cultures) or even impossible in the presence of planktonic calcifiers without sample pre-filtration. Filtration however, can alter CT concentration through gas exchange induced by high pressure. Addressing these problems, we present precise quantification of CT using a small, basic and inexpensive gas chromatograph as a CT analyzer. Our technique is able to provide a repeatability of ±3.1 μmol kg−1, given by the pooled standard deviation over a CT range typically applied in acidification experiments. 200 μL of sample is required to perform the actual CT measurement. The total sample amount needed is 12 mL. Moreover, we show that sample filtration is applicable with only minor alteration of the CT. The method is simple, reliable and with low cumulative material costs. Hence, it is potentially attractive for all researchers experimentally manipulating the seawater carbonate system

High nitrate to phosphorus regime attenuates negative effects of rising pCO2 on total population carbon accumulation

The ongoing rise in atmospheric pCO2 and consequent increase in ocean acidification have direct effects on marine calcifying phytoplankton, which potentially alters carbon export. To date it remains unclear, firstly, how nutrient regime, in particular by coccolithophores preferred phosphate limitation, interacts with pCO2 on particulate carbon accumulation; secondly, how direct physiological responses on the cellular level translate into total population response. In this study, cultures of Emiliania huxleyi were full-factorially exposed to two different N:P regimes and three different pCO2 levels. Cellular biovolume and PIC and POC content significantly declined in response to pCO2 in both nutrient regimes. Cellular PON content significantly increased in the Redfield treatment and decreased in the high N:P regime. Cell abundance significantly declined in the Redfield and remained constant in the high N:P regime. We hypothesise that in the high N:P regime severe phosphorous limitation could be compensated either by reduced inorganic phosphorous demand and/or by enzymatic uptake of organic phosphorous. In the Redfield regime we suggest that enzymatic phosphorous uptake to supplement enhanced phosphorous demand with pCO2 was not possible and thus cell abundance declined. These hypothesised different physiological responses of E. huxleyi among the nutrient regimes significantly altered population carrying capacities along the pCO2 gradient. This ultimately led to the attenuated total population response in POC and PIC content and biovolume to increased pCO2 in the high N:P regime. Our results point to the fact that the physiological (i.e. cellular) PIC and POC response to ocean acidification cannot be linearly extrapolated to total population response and thus carbon export. It is therefore necessary to consider both effects of nutrient limitation on cell physiology and their consequences for population size when predicting the influence of coccolithophores on atmospheric pCO2 feedback and their function in carbon export mechanisms

Eco-Evolutionary Interaction in Competing Phytoplankton: Nutrient Driven Genotype Sorting Likely Explains Dominance Shift and Species Responses to CO2

How ecological and evolutionary processes interact and together determine species and community responses to climate change is poorly understood. We studied long-term dynamics (over approximately 200 asexual generations) in two phytoplankton species, a coccolithophore (Emiliania huxleyi), and a diatom (Chaetoceros affinis), to increased CO2 growing alone, or competing with one another in co-occurrence. To allow for rapid evolutionary responses, the experiment started with a standing genetic variation of nine genotypes in each of the species. Under co-occurrence of both species, we observed a dominance shift from C. affinis to E. huxleyi after about 120 generations in both CO2 treatments, but more pronounced under high CO2. Associated with this shift, we only found weak adaptation to high CO2 in the diatom and none in the coccolithophore in terms of species’ growth rates. In addition, no adaptation to interspecific competition could be observed by comparing the single to the two-species treatments in reciprocal assays, regardless of the CO2 treatment. Nevertheless, highly reproducible genotype sorting left only one genotype remaining for each of the species among all treatments. This strong evolutionary selection coincided with the dominance shift from C. affinis to E. huxleyi. Since all other conditions were kept constant over time, the most parsimonious explanation for the dominance shift is that the strong evolutionary selection was driven by the experimental nutrient conditions, and in turn potentially altered competitive ability of the two species. Thus, observed changes in the simplest possible two-species phytoplankton “community” demonstrated that eco-evolutionary interactions can be critical for predicting community responses to climate change in rapidly dividing organisms such as phytoplankton

Inter- and intraspecific phenotypic plasticity of three phytoplankton species in response to ocean acidification

Phenotypic plasticity describes the phenotypic adjustment of the same genotype to different environmental conditions and is best described by a reaction norm. We focus on the effect of ocean acidification on inter- and intraspecific reaction norms of three globally important phytoplankton species (Emiliania huxleyi, Gephyrocapsa oceanica and Chaetoceros affinis). Despite significant differences in growth rates between the species, they all showed a high potential for phenotypic buffering (similar growth rates between ambient and high CO2 conditions). Only three coccolithophore genotypes showed a reduced growth in high CO2. Diverging responses to high CO2 of single coccolithophore genotypes compared with the respective mean species responses, however, raise the question of whether an extrapolation to the population level is possible from single-genotype experiments. We therefore compared the mean response of all tested genotypes with a total species response comprising the same genotypes, which was not significantly different in the coccolithophores. Assessing species reaction norms to different environmental conditions on short time scale in a genotype-mix could thus reduce sampling effort while increasing predictive power

Maintenance of Intraspecific Diversity in Response to Species Competition and Nutrient Fluctuations

Intraspecific diversity is a substantial part of biodiversity, yet little is known about its maintenance. Understanding mechanisms of intraspecific diversity shifts provides realistic detail about how phytoplankton communities evolve to new environmental conditions, a process especially important in times of climate change. Here, we aimed to identify factors that maintain genotype diversity and link the observed diversity change to measured phytoplankton morpho-functional traits Vmax and cell size of the species and genotypes. In an experimental setup, the two phytoplankton species Emiliania huxleyi and Chaetoceros affinis, each consisting of nine genotypes, were cultivated separately and together under different fluctuation and nutrient regimes. Their genotype composition was assessed after 49 and 91 days, and Shannon’s diversity index was calculated on the genotype level. We found that a higher intraspecific diversity can be maintained in the presence of a competitor, provided it has a substantial proportion to total biovolume. Both fluctuation and nutrient regime showed species-specific effects and especially structured genotype sorting of C. affinis. While we could relate species sorting with the measured traits, genotype diversity shifts could only be partly explained. The observed context dependency of genotype maintenance suggests that the evolutionary potential could be better understood, if studied in more natural settings including fluctuations and competition

Manipulation of Non-random Species Loss in Natural Phytoplankton: Qualitative and Quantitative Evaluation of Different Approaches

Ecological research in recent decades revealed that species loss has a predominantly negative effect on ecosystem functioning and stability. Most of these studies were based on random species loss scenarios, but extinctions in nature are not random. Recent experimental studies using macroscopic communities largely advanced knowledge about the effects of non-random species loss. However, in microscopic communities like the phytoplankton, implementing realistic species loss scenarios is challenging and experimental data are scarce. Creating more realistic experiments to study the role of phytoplankton diversity for ecosystem functioning is particularly important, as they provide up to 50% of global primary productivity, form the basis of all pelagic food webs, and are important for biogeochemical cycling. In this study, we experimentally tested and evaluated three methods for non-random species loss in a natural marine phytoplankton community. Dilution, filtration, and heat stress removed the targeted rare, large, and sensitive species, respectively. All these species groups are extremely vulnerable to extinction in future climate scenarios and play important roles in the communities. Dilution and filtration with a fine mesh additionally decreased initial biomass, which increased the variability of species left in the respective replicates. The methods tested in this study can be used to non-randomly manipulate phytoplankton species diversity in communities used for experiments. However, in studies where species identities are more important than species richness, the dilution and filtration methods should be modified to eliminate the effect of decreasing initial biomass

Temporal variation in ecological and evolutionary contributions to phytoplankton functional shifts

Communities and their functioning are jointly shaped by ecological and evolutionary processes that manifest in diversity shifts of their component species and genotypes. How both processes contribute to community functional change over time is rarely studied. We here repeatedly quantified eco-evolutionary contributions to CO2-driven total abundance and mean cell size changes after short-, mid-, and longer-term (80, 168, and >168d, respectively) in experimental phytoplankton communities. While the CO2-driven changes in total abundance and mean size in the short- and mid-term could be predominantly attributed to ecological shifts, the relative contribution of evolution increased. Over the longer-term, the CO2-effect and underlying eco-evolutionary changes disappeared, while total abundance increased, and mean size decreased significantly independently of CO2. The latter could be presumably attributed to CO2-independent genotype selection which fed back to species composition. In conclusion, ecological changes largely dominated the regulation of environmentally driven phytoplankton functional shifts at first. However, evolutionary changes gained importance with time, and can ultimately feedback on species composition, and thus must be considered when predicting phytoplankton change

Mussel beds are biological power stations on intertidal flats

Intertidal flats are highly productive areas that support large numbers of invertebrates, fish, and birds. Benthic diatoms are essential for the function of tidal flats. They fuel the benthic food web by forming a thin photosynthesizing compartment in the top-layer of the sediment that stretches over the vast sediment flats during low tide. However, the abundance and function of the diatom film is not homogenously distributed. Recently, we have realized the importance of bivalve reefs for structuring intertidal ecosystems; by creating structures on the intertidal flats they provide habitat, reduce hydrodynamic stress and modify the surrounding sediment conditions, which promote the abundance of associated organisms. Accordingly, field studies show that high chlorophyll a concentration in the sediment co-vary with the presence of mussel beds. Here we present conclusive evidence by a manipulative experiment that mussels increase the local biomass of benthic microalgae; and relate this to increasing biomass of microalgae as well as productivity of the biofilm across a nearby mussel bed. Our results show that the ecosystem engineering properties of mussel beds transform them into hot spots for primary production on tidal flats, highlighting the importance of biological control of sedimentary systems. (C) 2017 Elsevier Ltd. All rights reserved.</p