Håkan Wennhage

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.

Ecosystem goods and services from Swedish coastal habitats: identification, valuation, and implications of ecosystem shifts.

Abstract Coastal areas are exposed to a variety of threats due to high population densities and rapid economic development. How will this affect human welfare and our dependence on natures capacity to provide ecosystem goods and services? This paper is original in evaluating this concern for major habitats (macroalgae, seagrasses, blue mussel beds, and unvegetated soft bottoms) in a temperate coastal setting. More than 40 categories of goods and services are classified into provisional, regulating, and cultural services. A wide variety of Swedish examples is described for each category, including accounts of economic values and the relative importance of different habitats. For example, distinguishing characteristics would be the exceptional importance of blue mussels for mitigation of eutrophication, sandy soft bottoms for recreational uses, and seagrasses and macroalgae for fisheries production and control of wave and current energy. Net changes in the provision of goods and services are evaluated for three cases of observed coastal ecosystem shifts: i) seagrass beds into unvegetated substrate; ii) unvegetated shallow soft bottoms into filamentous algal mat dominance; and iii) macroalgae into mussel beds on hard substrate. The results are discussed in a management context including accounts of biodiversity, interconnectedness of ecosystems, and potential of economic valuation.

Regime Shifts and Ecosystem Service Generation in Swedish Coastal Soft Bottom Habitats: When Resilience is Undesirable

Ecosystems can undergo regime shifts where they suddenly change from one state into another. This can have important implications for formulation of management strategies, if system characteristics develop that are undesirable from a human perspective, and that have a high resistance to restoration efforts. This paper identifies some of the ecological and economic consequences of increased abundance of filamentous algae on shallow soft bottoms along the Swedish west coast. It is suggested that a successive increase in the sediment nutrient pool has undermined the resilience of these shallow systems. After the regime shift has occurred, self-generation properties evolve keeping the system locked in a high-density algae state. The structural and functional characteristics of the new system state differ significantly from the original one, resulting in less valuable ecosystem goods and services generated for society. In Sweden, loss of value results from the reduced capacity for mitigating further coastal eutrophication, reduced habitat quality for commercial fishery species, and the loss of aesthetic and recreational values.

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.

Vulnerability of newly settled plaice (Pleuronectes platessa L.) to predation: effects of habitat structure and predator functional response

Abstract Plaice ( Pleuronectes platessa ) nursery grounds on the Swedish west coast have been subject to increasing cover of annual green macroalgae during recent years, with growth of algae starting at the time of plaice settlement in April to May. A laboratory experiment was performed to investigate how the vulnerability to predation of metamorphosing plaice was affected by the presence of filamentous algae. Predation by shrimps ( Crangon crangon ) on settling plaice larvae was higher on sand than among algae, whereas predation by crabs ( Carcinus maenas ) was unaffected by habitat type, suggesting a lower overall mortality of plaice in the vegetated habitat. When predators and prey were presented with a combination of the two habitats, predation by shrimps was as high as that in the sand treatment alone, whereas predation by crabs was lower than that in the two treatments with one habitat. Based on these results, an additional experiment was performed, investigating the functional response of shrimps to six densities of juvenile plaice in a sand habitat with alternative prey present. The proportional mortality of juvenile plaice (12–16 mm total length (TL)) was density-dependent and was best described by a type III (sigmoid) functional response of the predatory shrimps. The results suggested that the combined predation pressure from shrimps and crabs was lower among algae than on sand, but settling plaice and predatory shrimps chose the sand habitat. Plaice densities in the sigmoid part of the obtained functional response curve represented normal to high field densities of plaice on the Swedish west coast, suggesting that shrimp predation could have a stabilising effect on plaice recruitment. The formation of macroalgae mats could therefore lead to a concentration of plaice juveniles in the remaining sand habitat and increased mortality through density-dependent predation by shrimps.

Diet of the invasive piscivorous fish Fistularia commersonii in a recently colonized area of the eastern Mediterranean

The composition of the diet of the Indo-Pacific cornetfish Fistularia commersonii from the SE Aegean Sea is described. The stomach contents of 245 specimens collected between September 2004 and March 2005 were examined. Dietary comparisons were made reflecting the relationship between diet composition, time of year, and fish size. Correlation between predator length and prey length was significant. The diet of the blue cornetfish consists of 96% by number and 99.95% by weight of fish. Size classification and habitat of prey groups (benthic, supra-benthic, and pelagic) showed that with increased body length the blue cornetfish extended its diet to larger prey and more generalist feeding. Spicara smaris, Boops boops, and Mullidae spp. were the most abundant prey by weight whereas a variety of small benthic fish (especially gobiids) and newly hatched fish constituted the largest number of prey items. Length–weight relationships for the cornetfish were investigated.

Substratum selection by juvenile plaice (Pleuronectes platessa L.): Impact of benthic microalgae and filamentous macroalgae

Abstract Preference by juvenile plaice for vegetated or unvegetated substratum was investigated in laboratory experiments to study the impact of recent changes in vegetation structure in shallow nursery areas on the recruitment of plaice. The experiments were done in circular tanks with filtered flow-through sea-water. The tanks were divided into two parts, one with bare sediment and the other with mat-forming benthic microalgae or patches of filamentous macroalgae ( Enteromorpha intestinalis and Rhizoclonium spp.) attached to small cobbles. Substratum preference were tested for different size-classes of juvenile plaice, ranging in size from 11 to 50 mm. Recently metamorphosed (11–15 mm) and post-settlement (16–26 mm) juveniles showed a significant preference for bare sediment compared with areas containing patches of filamentous algae. Post-settlement juveniles (16–26 mm) also exhibited a significant preference for bare sediment compared with sediment covered with mat-forming benthic microalgae. The differences in substratum selection disappeared 6 to 8 weeks after settlement in individuals that were 35 to 50 mm in total length. Field sampling verified that density of plaice (17–34 mm) was significantly lower in vegetation patches than in areas of bare sediment.

Fish assemblage structure in relation to macrophytes and filamentous epiphytes in shallow non-tidal rocky- and soft-bottom habitats

SynopsisThe fish assemblage in nineteen shallow water (0–3 m) areas on the Swedish west coast, including an estuarine zone, was assessed during spring and autumn 1989 and autumn 1990, using semi-quantitative survey nets. Samples of macrovegetation were collected concurrently for estimates of species composition and biomass. Nine stations had rocky-bottom substrata and ten had soft-bottom substrata all characterized by high coverage of macrovegetation and variously overgrown with epiphytic filamentous algae. Fish assemblage structures were compared and related to vegetation biomass, substrata and estuarine influences. At rocky-bottom stations total fish biomass was positively correlated with total vegetation biomass and negatively correlated with the proportion of filamentous algae during autumn samplings. In soft-bottom habitats variation in vegetation was small between stations, and no correlation existed between vegetation biomass and fish biomass. However, the number of fish species in soft-bottom habitats decreased significantly with increasing dominance of filamentous algae. The component species of the fish assemblage varied in their relation to the vegetation biomass and structure suggesting differences in degree of association with vegetation at the species level. Multivariate analysis based on fish species composition and on vegetation assemblages at the individual stations, yielded two major groups in accordance with division of the substrate into rocky- and soft-bottom habitats. Vegetation biomass superimposed on the fish assemblage ordination indicated a relationship between vegetation biomass and fish assemblage structure. Location of stations, in relation to the estuary was reflected in subgroups formed in the fish assemblage based cluster and ordination, suggesting a substantial estuarine influence on the fish assemblages. Thus, substrate type, vegetation biomass and structure, and estuarine influence are all potential structuring factors for the fish assemblages. In our study, vegetation structure seems to be of major importance and changes such as increased dominance of filamentous algae, like that observed in coastal areas in Sweden, might cause significant changes in fish assemblage structure.

Diversity, structure and function of fish assemblages associated with Posidonia oceanica beds in an area of the eastern Mediterranean Sea and the role of non-indigenous species

Temporal and spatial variation in density, biomass and body size of littoral fish species associated with nearshore Posidonia oceanica meadows was studied over an annual cycle in an area of the eastern Mediterranean Sea. A total of 109,350 littoral fishes were collected, belonging to 34 families and 88 species. Density of fishes peaked during the summer due to high numbers of juveniles. Season was a significant factor determining density, although number of species and biomass did not show any obvious seasonal pattern. Throughout the study, schooling planktivorous fish species such as the picarel Spicara smaris, the bogue Boops boops and the damselfish Chromis chromis were dominant, both in terms of density (80%) and biomass (70%). Temporal variation in density and body size of fishes was used to assess the seasonal and ontogenetic habitat use of each species, with their affinity to seagrass assessed by comparing their respective distribution on sand. Four functional guilds were created (juvenile migrants, seagrass residents, seasonal migrants and occasional visitors) to describe the habitat use of P. oceanica meadows by each species. Several species associated with P. oceanica meadows used this habitat mainly as juveniles during summer, although many others were present concurrently as adults and as juveniles. Among the species encountered, 11 were non-indigenous of Indo-Pacific origin, of which three used seagrasses mainly as juveniles and four as residents. The non-indigenous silverstripe blaasop Lagocephalus sceleratus ranked among the 10 most dominant species in terms of biomass (2%) and was classified as a seagrass resident.

Feeding ecology of indigenous and non-indigenous fish species within the family Sphyraenidae.

The feeding ecology of two common indigenous (Sphyraena viridensis and Sphyraena sphyraena) and one abundant non-indigenous sphyraenid species, Sphyraena chrysotaenia, of Indo-Pacific Ocean origin, was investigated in an area of the eastern Mediterranean Sea. The stomach contents of 738 individuals of varying size, collected during the period December 2008 to August 2009, were examined. The dietary analyses revealed that all three species were specialized piscivores with a diet consisting of >90% fish, both by number and mass. Concurrent sampling of the fish assemblage made it possible to calculate selectivity as well as diet breadth and overlap of these strict piscivores. Even though several prey species were found in the stomachs of the three predators examined, selectivity towards Atherina boyeri was highly significant. For all species examined, >70% of the diet by mass was made up by three indigenous species of commercial value: Spicara smaris, Boops boops and A. boyeri. Diet breadth and size of prey increased with increasing body size for all predators. With increased body size, the diet overlap between indigenous and non-indigenous species decreased. This could be attributed to increased diet breadth and the specific life-history characteristics of indigenous species developing into larger individuals. During winter, the condition factor of the non-indigenous species was significantly lower than that of the indigenous, indicating that winter conditions in the Mediterranean Sea may limit its further expansion north and westward. With this study, the gap in knowledge of the feeding preferences of the most abundant piscivorous species found in coastal areas of the study region is filled. Additionally, the results indicate that non-indigenous species familial affiliation to indigenous ones does not facilitate invasion success.