Per-Olav Moksnes

Distribution of green algal mats throughout shallow soft bottoms of the Swedish Skagerrak archipelago in relation to nutrient sources and wave exposure

Abstract Distribution and biomass of green algal mats were studied in marine shallow (0–1 m) soft-bottom areas on the Swedish west coast from 1994 to 1996, by combining aerial photography surveys with ground truth sampling. Filamentous green algae, dominated by species of the genera Cladophora and Enteromorpha, were generally present throughout the study area during July and August, and largely absent in late April and early May. These algae occurred at 60 to 90% of the locations investigated during the summer, and were estimated to cover between 30 and 50% of the total area of shallow soft bottoms of the Swedish Skagerrak archipelago. The distributional patterns were similar during the three years of the investigation and appeared unrelated to annual local nutrient inputs from point sources and river discharge. We postulate that the apparent lack of such a relationship is due to an altered state of nutrient dynamics throughout the archipelago. Mechanisms are likely to involve long-term, diffuse elevations in nutrient levels in coastal waters of the Skagerrak and the Kattegat over several decades leading to current eutrophic conditions, exceeding nutrient requirements for abundant filamentous algal growth. Patterns of algal abundance in our study were largely related to physical factors such as exposure to wind, waves and water exchange under conditions where nutrient loads among embayments seemed to be unlimited. Further, our results show that sediments covered by algal mats had higher carbon and nitrogen contents than unvegetated sediments. We hypothesise that sustained high nutrient loads, manifested in extensive biomass of filamentous algae during summer months, are re-mineralised via decay and sedimentation in the benthic realm. Hence, accumulated carbon and nutrients in the sediment could, in turn, constitute the basic pool for future algal mat production overlying soft bottoms in areas where tidal exchange is limited.

The relative importance of habitat-specific settlement, predation and juvenile dispersal for distribution and abundance of young juvenile shore crabs Carcinus maenas L.

Abstract Young juveniles of many motile benthic species are concentrated in structurally complex habitats, but the proximate causes of this distribution are usually not clear. In the present study, I assessed three potentially important processes affecting distribution and abundance of early benthic stages in the shore crab (Carcinus maenas): (1) selection of habitat by megalopae (postlarvae); (2) habitat-specific predation; and (3) post-settlement movements by juveniles. These processes were assessed concurrently over 3–9 days at two spatial scales: at the scale of square meters using cage techniques within nursery areas, and at the scale of hectares using isolated populations of juvenile shore crabs in small nursery areas as mesocosms. The results were compared to habitat-specific distribution in the field. Shore crab megalopae and first instar juveniles (settlers) were distributed non-randomly among micro-habitats in the assessed nursery areas, with great densities in both mussel beds, eelgrass and filamentous algal patches (on average 114–232 settlers m−2), and significantly smaller densities on open sand habitats at all times (on average 4 settlers m−2). The same habitat-specific settlement pattern was found in cages where predators were excluded, suggesting that active habitat selection at settlement was responsible for the initial distribution. Older juveniles (second to ninth instar crabs) were also sparse on sand, but in contrast to settlers, were concentrated in mussel beds, which showed significantly greater densities than eelgrass and algal habitats. The cage experiment demonstrated a dynamic distribution of juvenile crabs. Young juveniles constantly migrated over open sand habitats (20 m or further) and colonized the experimental plots in a habitat-specific pattern that reflected the distribution in the field. This pattern was also found for very small crabs colonizing predator-exclusion cages, suggesting that selection of habitat by migrating juveniles caused the ontogenetic change in habitat use. Although post-settlement movements were great within nursery areas, juvenile dispersal at a regional scale appeared to be small, and the recruitment of juvenile shore crabs to the shallow bays occurred mainly through pelagic megalopae. Conservative estimates at the scale of whole nursery areas, based on migration trap data and field samples, indicated great mortality of settlers and early benthic stages of shore crabs. Results from the cage experiment suggest that predation by crabs and shrimp were responsible for the high settlement mortality. Both enclosed cannibalistic juvenile crabs and local predators on uncaged habitat plots caused significant losses of settlers in all habitats (on average 22% and 64% 3 day−1, respectively). The effect of predators was highly variable between trials, but differed little between habitat types, and predation had no detectable proximate effect on juvenile distribution, despite the great losses. Small settlement densities on sand habitats in combination with a refuge at low prey numbers, and an aggregation of cannibalistic juvenile crabs in nursery habitats appear to decrease the effect of habitat-specific predation rates on the distribution of juvenile shore crabs. This study demonstrates that active habitat selection at settlement followed by a dynamic redistribution of young juveniles can be the proximate processes responsible for habitat-specific distribution of epibenthic juveniles, and indicate that predation represents a major evolutionary process reinforcing this behavior.

Biodiversity mediates top–down control in eelgrass ecosystems: a global comparative‐experimental approach

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.

Recent increase of filamentous algae in shallow Swedish bays: Effects on the community structure of epibenthic fauna and fish

A summary is presented of estimates of distribution and growth of filamentous algae and its effect on the structure and functioning of epibenthic fauna and fish communities in shallow bays on the Swedish west coast. As a consequence of coastal eutrophication vegetation cover has gradually increased during the last decade, and during 1990s most bays in the Skagerrak-Kattegat area were variously covered with filamentous algae during spring and summer (May–July). In some areas filamentous algae (mainlyCladophora andEnteromorpha) completely covered the bottom. In field studies it was demonstrated that increased cover and dominance of filamentous algae result in structural changes of the epibenthic fauna community. Field studies showed that species richness and biomass of epibenthic fauna increased in a sandy bay with a moderate increase (30 to 50%) of filamentous algae cover. At higher cover (90%), biomass of epibenthic fauna was reduced, however, to the same level as for the sandy habitat, although the dominant epibenthic species were different. Heavy growth of epiphytic filamentous algae on eelgrass resulted in reduced biomass and a shift in the species composition of the epibenthic fauna community. Fish assemblage structure was also related to changes in vegetation. In eelgrass beds, fish species numbers were reduced with increasing cover of epiphytic filamentous algae, and at rocky bottoms with kelp algae (dominated byFucus), fish biomass decreased with increasing cover of attached filamentous algae. Further, foraging efficiency of juvenile cod and settling success of plaice were reduced as a response to increasing dominance of filamentous algae.

Active habitat selection by megalopae and juvenile shore crabs Carcinus maenas: a laboratory study in an annular flume

We studied megalopae (postlarvae) and young juveniles of the shore crab (Carcinus maenas L.) in laboratory experiments to examine four potentially important processes for juvenile distribution and recruitment: (1) hydrodynamic processes and passive deposition of megalopae, (2) active habitat selection of megalopae, (3) habitat specific predation rates, and (4) active habitat selection by juveniles.

Top-down control as important as nutrient enrichment for eutrophication effects in North Atlantic coastal ecosystems

Seagrass and seaweed habitats constitute hotspots for diversity and ecosystem services in coastal ecosystems. These habitats are subject to anthropogenic pressures, of which eutrophication is one major stressor. Eutrophication favours fast-growing ephemeral algae over perennial macroalgae and seagrasses, causing habitat degradation. However, changes in top-down control, caused by, for example, overfishing, may also have negative impacts on such habitats by decreasing grazer control of ephemeral algae. Meanwhile, systematic analyses estimating top-down effects of predator manipulations across a wide range of studies are missing, limiting the potential use of top-down control measures in coastal management. Here, we review the literature on experiments that test top-down and bottom-up controls in seagrass Zostera marina and seaweed Fucus spp. food webs in the North Atlantic. Using meta-analysis and meta-regression, we compare effect sizes of consumer and nutrient manipulations on primary producers, grazers and mesopredators. Presence of mesopredators on average doubled the biomass of ephemeral algae through trophic cascades, mainly mediated via negative effects on amphipods and isopods. Of the grazers, gastropods had twice as strong a negative effect on ephemeral algae as amphipods/isopods, but responded weakly to both predators and fertilization. In accordance with theory, top-down effects became stronger with eutrophication. Across studies, top-down effects on ephemeral algae at all trophic levels are on par with eutrophication effects. However, the few studies manipulating piscivorous fish make estimates of their top-down effects uncertain.Synthesis and applications. Consistently strong top-down effects in coastal ecosystems call for an integrated ecosystem perspective. Management should consider measures to improve stocks of predatory fish and reduce mesopredators for restoration and conservation of essential seagrass and seaweed habitats, thereby increasing the long-term viability of ecosystem services from coastal habitats.Editors Choice

No barrier to emergence of bathyal king crabs on the Antarctic shelf

Significance For tens of millions of years, cold conditions have excluded shell-crushing fish and crustaceans from the continental shelf surrounding Antarctica. Rapid warming is now allowing predatory crustaceans to return. Our study of the continental slope off the western Antarctic Peninsula showed that abundant, predatory king crabs comprise a reproductively viable population at 841- to 2,266-m depth. Depth profiles of temperature, salinity, habitat structure, food availability, and predators indicate that there are no barriers to prevent king crabs from moving upward onto the outer shelf at 400–550 m. A cold-water barrier above 200 m could be breached within the next few decades. Emergence of king crabs on the shelf could have catastrophic consequences for the unique seafloor communities of Antarctica. Cold-water conditions have excluded durophagous (skeleton-breaking) predators from the Antarctic seafloor for millions of years. Rapidly warming seas off the western Antarctic Peninsula could now facilitate their return to the continental shelf, with profound consequences for the endemic fauna. Among the likely first arrivals are king crabs (Lithodidae), which were discovered recently on the adjacent continental slope. During the austral summer of 2010‒2011, we used underwater imagery to survey a slope-dwelling population of the lithodid Paralomis birsteini off Marguerite Bay, western Antarctic Peninsula for environmental or trophic impediments to shoreward expansion. The population density averaged ∼4.5 individuals × 1,000 m−2 within a depth range of 1,100‒1,500 m (overall observed depth range 841–2,266 m). Images of juveniles, discarded molts, and precopulatory behavior, as well as gravid females in a trapping study, suggested a reproductively viable population on the slope. At the time of the survey, there was no thermal barrier to prevent the lithodids from expanding upward and emerging on the outer shelf (400- to 550-m depth); however, near-surface temperatures remained too cold for them to survive in inner-shelf and coastal environments (<200 m). Ambient salinity, composition of the substrate, and the depth distribution of potential predators likewise indicated no barriers to expansion of lithodids onto the outer shelf. Primary food resources for lithodids—echinoderms and mollusks—were abundant on the upper slope (550–800 m) and outer shelf. As sea temperatures continue to rise, lithodids will likely play an increasingly important role in the trophic structure of subtidal communities closer to shore.

Genome architecture enables local adaptation of Atlantic cod despite high connectivity

Adaptation to local conditions is a fundamental process in evolution; however, mechanisms maintaining local adaptation despite high gene flow are still poorly understood. Marine ecosystems provide a wide array of diverse habitats that frequently promote ecological adaptation even in species characterized by strong levels of gene flow. As one example, populations of the marine fish Atlantic cod (Gadus morhua) are highly connected due to immense dispersal capabilities but nevertheless show local adaptation in several key traits. By combining population genomic analyses based on 12K single nucleotide polymorphisms with larval dispersal patterns inferred using a biophysical ocean model, we show that Atlantic cod individuals residing in sheltered estuarine habitats of Scandinavian fjords mainly belong to offshore oceanic populations with considerable connectivity between these diverse ecosystems. Nevertheless, we also find evidence for discrete fjord populations that are genetically differentiated from offshore populations, indicative of local adaptation, the degree of which appears to be influenced by connectivity. Analyses of the genomic architecture reveal a significant overrepresentation of a large ~5 Mb chromosomal rearrangement in fjord cod, previously proposed to comprise genes critical for the survival at low salinities. This suggests that despite considerable connectivity with offshore populations, local adaptation to fjord environments may be enabled by suppression of recombination in the rearranged region. Our study provides new insights into the potential of local adaptation in high gene flow species within fine geographical scales and highlights the importance of genome architecture in analyses of ecological adaptation.

Valuing Multiple Eelgrass Ecosystem Services in Sweden: Fish Production and Uptake of Carbon and Nitrogen

Valuing nature’s benefits in monetary terms is necessary for policy-makers facing trade-offs in how to spend limited financial resources on environmental protection. We provide information to assess trade-offs associated with the management of seagrass beds, which provide a number of ecosystem services, but are presently impacted by many stressors. We develop an interdisciplinary framework for valuing multiple ecosystem services and apply it to the case of eelgrass (Zostera marina), a dominant seagrass species in the northern hemisphere. We identify and quantify links between three eelgrass functions (habitat for fish, carbon and nitrogen uptake) and economic goods in Sweden, quantify these using ecological endpoints, estimate the marginal average value of the impact of losing one hectare of eelgrass along the Swedish northwest coast on welfare in monetary terms, and aggregate these values while considering double-counting. Over a 20 to 50 year period we find that compared to unvegetated habitats, a hectare of eelgrass, including the organic material accumulated in the sediment, produces an additional 626 kg cod fishes and 7,535 wrasse individuals and sequesters 98.6 ton carbon and 466 kg nitrogen. We value the flow of future benefits associated with commercial fishing, avoided climate change damages, and reduced eutrophication at 170,000 SEK in 2014 (20,700 US

Seed Predation by the Shore Crab Carcinus maenas: A Positive Feedback Preventing Eelgrass Recovery?

) or 11,000 SEK (1,300 US