Post-settlement dispersal: the neglected link in maintenance of soft-sediment biodiversity

Seafloor integrity is threatened by disturbances owing to human activities. The capacity of the system to recover from disturbances, as well as maintain resilience and function, depends on dispersal. In soft-sediment systems, dispersal continues after larval settlement, but there are very few measurements of how far the post-settlers disperse in nature. Spatial scales of post-settlement dispersal are, however, likely to be similar to pelagic larval dispersal because of continued, frequent, small-scale dispersal over longer periods. The consequences of this dispersal may be more important for the maintenance of biodiversity and metacommunity dynamics than is pelagic larval dispersal, because of the greater size and competency of the dispersers. We argue that an increased empirical understanding of post-settlement dispersal processes is key for predicting how benthic communities will respond to local disturbances and shrinking regional species pools, with implications for monitoring, managing and conserving biodiversity.     

Complex metacommunity structure for benthic invertebrates in a low‐diversity coastal system

The majority of studies in metacommunity ecology have focused on systems other than marine benthic ecosystems, thereby providing an impetus to broaden the focus of metacommunity research to comprise marine systems. These systems are more open than many other systems and may thus exhibit relatively less discrete patterns in community structure across space. Metacommunity structure of soft‐sediment benthic invertebrates was examined using a fine‐grained (285 sites) data set collected during one summer across a large spatial extent (1700 km²). We applied the elements of metacommunity structure (EMS) approach, allowing multiple hypothesis of variation in community structure to be tested. We demonstrated several patterns associated with environmental variation and associated processes that could simultaneously assemble species to occur at the sites. A quasi‐Clementsian pattern was observed frequently, suggesting interdependent ecological relationships among species or similar response to an underlying environmental gradient across sites. A quasi‐nested clumped species loss pattern was also observed, which suggests nested habitat specialization. Species richness declined with depth (from 0.5 to 44.8 m). We argue that sensitive species may survive in shallower water, which are more stable with regard to oxygen conditions and present greater habitat complexity, in contrast to deeper waters, which may experience periodic disturbance due to hypoxia. Future studies should better integrate disturbance in terms of temporal dynamics and dispersal rates in the EMS approach. We highlight that shallow water sites may act as sources of recruitment to deeper water sites that are relatively more prone to periodic disturbances due to hypoxia. However, these shallow sites are not currently monitored and should be better prioritized in future conservation strategies in marine systems.     

Determining effects of suspended sediment on condition of a suspension feeding bivalve (Atrina zelandica): results of a survey, a laboratory experiment and a field transplant experiment

The horse mussel Atrina zelandica (Gray) is a large, suspension feeding pinnid bivalve, common in coastal and estuarine areas of northern New Zealand. As a suspension feeder, Atrina is likely to be influenced by suspended sediment loads. We conducted a laboratory experiment to determine the effect of short-term elevations in turbidity levels, such as those commonly recorded during storms, on the physiological condition and clearance rates of Atrina. We also conducted a field survey and a 3-month transplant experiment at multiple sites along a gradient of increasing suspended sediment load in a New Zealand estuary. Laboratory clearance rates of Atrina declined above a threshold suspended sediment concentration, and Atrina physiological condition at the end of this experiment was lower in high cf. low turbidity treatments. Decreases in Atrina condition were detected after exposure to elevated levels for only 3 days. The field survey and transplant experiment provided empirical evidence of a strong, negative effect of increasing suspended sediment flux on the physiological condition of Atrina. We suggest that relationships between the physiological condition of suspension feeders and sediment settling flux could provide a link between sediment inputs, which commonly occur as a result of catchment runoff during rainfall events, and the ecological health of estuarine and shallow coastal areas. Our study also demonstrated that Atrina have a natural distribution limit controlled by suspended sediment load. Thus, there is potential for larger-scale functional and structural effects on benthic communities in estuarine and coastal areas with high rates of sedimentation.     

Long‐term temporal and spatial trends in eutrophication status of the Baltic Sea

Much of the Baltic Sea is currently classified as ‘affected by eutrophication’. The causes for this are twofold. First, current levels of nutrient inputs (nitrogen and phosphorus) from human activities exceed the natural processing capacity with an accumulation of nutrients in the Baltic Sea over the last 50–100 years. Secondly, the Baltic Sea is naturally susceptible to nutrient enrichment due to a combination of long retention times and stratification restricting ventilation of deep waters. Here, based on a unique data set collated from research activities and long‐term monitoring programs, we report on the temporal and spatial trends of eutrophication status for the open Baltic Sea over a 112‐year period using the HELCOM Eutrophication Assessment Tool (HEAT 3.0). Further, we analyse variation in the confidence of the eutrophication status assessment based on a systematic quantitative approach using coefficients of variation in the observations. The classifications in our assessment indicate that the first signs of eutrophication emerged in the mid‐1950s and the central parts of the Baltic Sea changed from being unaffected by eutrophication to being affected. We document improvements in eutrophication status that are direct consequences of long‐term efforts to reduce the inputs of nutrients. The reductions in both nitrogen and phosphorus loads have led to large‐scale alleviation of eutrophication and to a healthier Baltic Sea. Reduced confidence in our assessment is seen more recently due to reductions in the scope of monitoring programs. Our study sets a baseline for implementation of the ecosystem‐based management strategies and policies currently in place including the EU Marine Strategy Framework Directives and the HELCOM Baltic Sea Action Plan.     

Drifting algal mats as an alternative habitat for benthic invertebrates: Species specific responses to a transient resource

Patchy occurrences of benthic drift algae (i.e. loose lying macroalgal mats) may increase habitat complexity on normally bare soft bottoms, but at the same time, extensive amounts of drifting algal mats are known to stress the benthic fauna. This paper presents results of the first detailed study of the fauna associated with drift algal mats in the northern Baltic Sea. In order to assess the importance of drifting algae as an alternative habitat for benthic fauna, benthic drift algal mats were sampled on shallow (2-9 m) sandy soft bottoms in the outer archipelago of the ?land Islands (Finland). Species composition, abundance and biomass of the macrofauna associated with algal mats were recorded. The results show that drifting algae at times can harbour very high abundances of invertebrates (up to 1116 individuals/g algal dryweight), surpassing invertebrate densities recorded in seagrass communities. The algal fauna varied between sites and over time, and factors such as ambient benthic fauna, exposure to wind-wave disturbance, depth, and algal coverage and condition influenced the invertebrate community composition of the algal mats. Abundance increased while individual biomass of the animals decreased over time (summer season; July-October). A series of laboratory experiments were conducted in order to test the ability of a few important benthic species to move up into, and survive in a drifting algal mat. Macoma balthica, Hydrobia spp., Nereis diversicolor and Bathyporeia pilosa were used in the experiments, and significant differences in their survival and mobility within drifting algae were recorded. This study shows that benthic species differ significantly in their ability to utilise the algal mats, with mainly opportunistic and mobile taxa such as Hydrobia spp., Chironomidae and Ostracoda benefiting from the algae, whereas infaunal species such as M. balthica and B. pilosa are negatively affected. The occurrence of eutrophication induced drifting macroalgal mats has increased significantly during the last decade in the northern Baltic Sea. Hence, the importance of drifting algae as a stress factor and as an alternative habitat for benthic fauna increases.     

Macroalgal photosynthesis near the southern global limit for growth; Cape Evans,Ross Sea,Antarctica

Photosynthetic characteristics of the red macroalgae Phyllophora antarctica and Phymatolithon foecundum collected from under sea ice at Cape Evans, McMurdo Sound (Ross Sea) were determined using in situ fluorometric and lab-based oxygen exchange techniques. Only 0.16% of incident irradiance penetrated the 2.5 m thick ice cover and photosynthetic parameters for both taxa were characteristic of highly shade-adapted plants. Saturation onset parameter (E _k) did not exceed 13 mol photons m^-2 s^-1 in either taxon. For Phyllophora antarctica the light saturated photosynthetic rate at –1^°C was 10 mol O₂ g^-1 FW h^-1 and respiration averaged 3.3 mol O₂ g^-1 FW h^-1 between sampled depths of 10 and 25 m. A light meter deployed at 15 m depth for a year recorded a marked increase in underwater irradiance on the last day of January 2002 coinciding with ice-breakout, and a maximum value for irradiance of 120 mol photons m^-2 s^-1 on 9 February 2002. The 2-month ice-free period was the only time when irradiance consistently exceeded compensation (photosynthesis=respiration) and enabled Phyllophora antarctica to accumulate sufficient carbon to result in a measurable increase in thallus area equivalent to a biomass increment of 1.87 mg (DW) per frond. Near the southern global limit for marine macroalgae, conditions that dictate the availability of underwater irradiance are extremely variable from year to year. Low respiration rates enhance longevity of the Phyllophora antarctica thallus, enabling it to not only survive the winter darkness, but also to retain photosynthetic capacity and thus take advantage of windows of higher irradiance.     

Effects of Reduced pH on Macoma balthica Larvae from a System with Naturally Fluctuating pH-Dynamics

Ocean acidification is causing severe changes in the inorganic carbon balance of the oceans. The pH conditions predicted for the future oceans are, however, already regularly occurring in the Baltic Sea, and the system might thus work as an analogue for future ocean acidification scenarios. The characteristics of the Baltic Sea with low buffering capacity and large natural pH fluctuations, in combination with multiple other stressors, suggest that OA effects may be severe, but remain largely unexplored. A calcifying species potentially affected by low pH conditions is the bivalve Macoma balthica (L.). We investigated larval survival and development of M. balthica by exposing the larvae to a range of pH levels: 7.2, 7.4, 7.7 and 8.1 during 20 days in order to learn what the effects of reduced pH are on the larval biology and thus also potentially for the population dynamics of this key species. We found that even a slight pH decrease causes significant negative changes during the larval phase, both by slowing growth and by decreasing survival. The growth was slower in all reduced pH treatments compared to the control treatment. The size of 250 µm that is considered indicative to imminent settling in our system was reached by 22% of the larvae grown in control conditions after 20 days, whereas in all reduced pH treatments the size of 250 µm was reached by only 7–14%. The strong impact of ocean acidification on larvae is alarming as slowly growing individuals are exposed to higher predation risk in response to the longer time they are required to spend in the plankton, further decreasing the ecological competence of the species.