Polychaete invader enhances resource utilization in a species-poor system.

Ecosystem consequences of biodiversity change are often studied from a species loss perspective, while the effects of invasive species on ecosystem functions are rarely quantified. In this experimental study, we used isotope tracers to measure the incorporation and burial of carbon and nitrogen from a simulated spring phytoplankton bloom by communities of one to four species of deposit-feeding macrofauna found in the species-poor Baltic Sea. The recently invading polychaete Marenzelleria arctia, which has spread throughout the Baltic Sea, grows more rapidly than the native species Monoporeia affinis, Pontoporeia femorata (both amphipods) and Macoma balthica (a bivalve), resulting in higher biomass increase (biomass production) in treatments including the polychaete. Marenzelleria incorporated and buried bloom material at rates similar to the native species. Multi-species treatments generally had higher isotope incorporation, indicative of utilization of bloom material, than expected from monoculture yields of the respective species. The mechanism behind this observed over-yielding was mainly niche complementarity in utilization of the bloom input, and was more evident in communities including the invader. In contrast, multi-species treatments had generally lower biomass increase than expected. This contrasting pattern suggests that there is little overlap in resource use of freshly deposited bloom material between Marenzelleria and the native species but it is likely that interference competition acts to dampen resulting community biomass. In conclusion, an invasive species can enhance incorporation and burial of organic matter from settled phytoplankton blooms, two processes fundamental for marine productivity.

Diversity of larger consumers enhances interference competition effects on smaller competitors.

Competition between large and small species for the same food is common in a number of ecosystems including aquatic ones. How diversity of larger consumers affects the access of smaller competitors to a limiting resource is not well understood. We tested experimentally how species richness (0-3 spp.) of benthic deposit-feeding macrofauna changes meiofaunal ostracods' incorporation of fresh organic matter from a stable-isotope-labeled cyanobacterial bloom, using fauna from the species-poor Baltic Sea. Presence of macrofauna mostly decreased meiofaunal incorporation of bloom material, depending on the macrofauna species present. As expected, the species identity of macrofauna influenced the incorporation of organic matter by meiofauna. Interestingly, our results show that, in addition, species richness of the macrofauna significantly reduced meiofauna incorporation of freshly settled nitrogen and carbon. With more than one macrofauna species, the reduction was always greater than expected from the single-species treatments. Field data from the Baltic Sea showed a negative correlation between macrofauna diversity and meiofaunal ostracod abundance, as expected from the experimental results. We argue that this is caused by interference competition, due to spatial niche differentiation between macrofauna species reducing the sediment volume in which ostracods can feed undisturbed by larger competitors. Interference from macrofauna significantly reduces organic matter incorporation by meiofauna, indicating that diversity of larger consumers is an important factor controlling the access of smaller competitors to a limiting food resource.

Higher diversity of deposit-feeding macrofauna enhances phytodetritus processing.

The link between biodiversity and ecosystem functioning is an important question that remains unresolved, particularly in marine systems, in which cycling of organic matter by benthic organisms is of global significance. Direct observations of specific resource use by each species in single- and multispecies communities, as quantified by stable isotopes, facilitates a mechanistic understanding of the importance of each species for ecosystem functioning. We tested the effects of altered biodiversity (species richness) of deposit-feeding macrofauna on incorporation and burial of phytodetritus in combinations of three species representing natural communities found in the sediments of the species-poor Baltic Sea. The three species, two amphipods and a bivalve, had different rates of incorporation and burial and different needs for carbon (C) and nitrogen (N). The amphipods exhibited clear resource partitioning in sympatry, as a result of vertical separation in the sediment and consequent differential use of food. Communities of several species incorporated more C and N than expected from the respective single-species treatments, due to higher incorporation by surface feeders in multispecies treatments. Community incorporation of N in the most diverse treatment even exceeded N incorporation by a single-species treatment of the best-performing species, showing transgressive over-yielding. This over-yielding was primarily due to positive complementarity in all treatments. Diverse soft bottoms are also likely to be more productive in the long run, as species-specific traits (subsurface feeding) preserve fresh phytodetritus by burying it to depths in the sediment at which the mineralization rate is low. The more diverse sediment communities showed more efficient trophic transfer of phytodetritus, a finding of general significance for understanding biological processes driving the transformation of nutrients and energy in benthic ecosystems.

Effects of an invasive polychaete on benthic phosphorus cycling at sea basin scale: An ecosystem disservice.

Macrofaunal activities in sediments modify nutrient fluxes in different ways including the expression of species-specific functional traits and density-dependent population processes. The invasive polychaete genus Marenzelleria was first observed in the Baltic Sea in the 1980s. It has caused changes in benthic processes and affected the functioning of ecosystem services such as nutrient regulation. The large-scale effects of these changes are not known. We estimated the current Marenzelleria spp. wet weight biomass in the Baltic Sea to be 60-87 kton (95% confidence interval). We assessed the potential impact of Marenzelleria spp. on phosphorus cycling using a spatially explicit model, comparing estimates of expected sediment to water phosphorus fluxes from a biophysical model to ecologically relevant experimental measurements of benthic phosphorus flux. The estimated yearly net increases (95% CI) in phosphorous flux due to Marenzelleria spp. were 4.2-6.1 kton based on the biophysical model and 6.3-9.1 kton based on experimental data. The current biomass densities of Marenzelleria spp. in the Baltic Sea enhance the phosphorus fluxes from sediment to water on a sea basin scale. Although high densities of Marenzelleria spp. can increase phosphorus retention locally, such biomass densities are uncommon. Thus, the major effect of Marenzelleria seems to be a large-scale net decrease in the self-cleaning capacity of the Baltic Sea that counteracts human efforts to mitigate eutrophication in the region.

Hexabromocyclododecane affects benthic-pelagic coupling in an experimental ecosystem.

Hexabromocyclododecane (HBCDD) is an additive brominated flame retardant and a recognized PBT chemical. However, little is known about its effects on coastal species, and even less on ecosystem effects. We investigated the dose-response effects of HBCDD over 8 months in 1000 L experimental mesocosms assembled from coastal Baltic Sea ecosystem components. HBCDD was added via spiked plankton material and a range of structural and functional endpoints were measured during the experiment. Increasing HBCDD concentration decreased the biomass of large Macoma balthica, resulting in a decreased recirculation of nutrients to the water. Changes in plankton communities were also observed, either due to direct toxic HBCDD effects or indirect via changes in benthic-pelagic coupling of nutrients. Such complex ecosystem responses can only be quantified and understood by using realistic experimental set-ups, and including knowledge of system-specific ecological interactions. This is the first study of HBCDD effects on ecosystem level.

Meiofauna reduces bacterial mineralization of naphthalene in marine sediment.

The role of sediment-living meiofauna, benthic invertebrates smaller than 1000 µm such as nematodes and ostracods, on the mineralization of naphthalene, a common polycyclic aromatic hydrocarbon (PAH) in marine sediments, was studied in microcosms using radiorespirometry. A method to extract live meiofauna was developed and used in order to experimentally manipulate meiofauna abundance and group diversity. Higher abundances of meiofauna were found to significantly decrease naphthalene mineralization. Furthermore, a change in the bacterial community composition (studied using terminal restriction fragment length polymorphism) was also observed in presence of higher meiofauna abundance, as well as a lower number of cultivable naphthalene-degrading bacteria. The reduced mineralization of naphthalene and the altered bacterial community composition in the presence of increased meiofauna abundance is likely the result of top-down control by meiofauna. This study shows that higher abundances of meiofauna can significantly decrease the microbial mineralization of PAHs such as naphthalene and also significantly modify the bacterial community composition in natural marine sediments.

Effects of the antibiotic ciprofloxacin on the bacterial community structure and degradation of pyrene in marine sediment.

The ecological consequences of antibiotics in the aquatic environment have been an issue of concern over the past years due to the potential risk for negative effects on indigenous microorganisms. Microorganisms provide important ecosystem services, such as nutrient recycling, organic matter mineralization and degradation of pollutants. In this study, effects of exposure to the antibiotic ciprofloxacin on the bacterial diversity and pollutant degradation in natural marine sediments were studied using molecular methods (T-RFLP) in combination with radiorespirometry. In a microcosm experiment, sediment spiked with (14)C-labelled pyrene was exposed to five concentrations of ciprofloxacin (0, 20, 200, 1000 and 2000 microgL(-1)) in a single dose to the overlying water. The production of (14)CO(2) (i.e. complete mineralization of pyrene) was measured during 11 weeks. Sediment samples for bacterial community structure analysis were taken after 7 weeks. Results showed a significant dose-dependent inhibition of pyrene mineralization measured as the total (14)CO(2) production. The nominal EC(50) was calculated to 560 microgL(-1), corresponding to 0.4 microg/kg d.w. sediment. The lowest effect concentration on the bacterial community structure was 200 microgL(-1), which corresponds to 0.1 microg/kg d.w. sediment. Our results show that antibiotic pollution can be a potential threat to both bacterial diversity and an essential ecosystem service they perform in marine sediment.