Perennial Algae Research Articles

태안반도 꽃지 약용해조의 계절적 군집구조 및 수직분포

Marine algal flora and community structure of medicinal seaweeds were examined at Kkotji of Taean Peninsula, Korea from May 2005 to January 2006. Seventy-nine seaweeds including 42 medicinal algae and one marine plant were identified. Sargassum thunbergii was the representative alga occurred at all seasons and shore levels. The dominant medicinal seaweeds were perennial S. thunbergii, Neorhodomela aculeata, and Corallina pilulifera, and ephemeral Monostroma grevillei, Porphyra yezoensis, and Ulva pertusa. Their vertical distribution were N. aculeata – P. yezoensis, M. grevillei, and U. pertusa – C. pilulifera from high to low intertidal zone. The average biomass of medicinal seaweeds varied from 34.17 g m–2 in spring to 56.41 g m–2 in summer. At Kkotji shore, the opportunistic species (Enteromorpha, Ulva, and Cladophora) and turf-forming algae (Caulacanthus okamurae and Gelidium divaricatum) were easily observed. Such fast growing ESG II (ecological state group) was 87.50% and slow growing perennial algae, ESG I was only 12.15%. Also, diversity index (H’) and dominance index (DI) indicate that the seaweed community of Kkotji is unstable. Therefore, Kkotji rocky shore should be more protected from human activities such as turbulence and eutrophication in order to maintain species diversity and abundance of medicinal seaweeds.

다공질 인공어초 (반톱니형)에서 진행된 해조천이 및 해중림 조성

The succession of marine benthic algae and the restoration of an artificial seaweed forest on a porous type (shaped half saw teeth) artificial reef at Jeju island, Korea was studied. Young thalli of Sargassum horneri and Ecklonia cava were attached to different artificial substrates. In general, the succession on the artificial reefs led from filamentous algae to perennial algae and involved more than 25 species that are useful fishery resources, including E. cava. Coralline algae were dominant on the artificial reefs at the Kangjung site. The maximum algal biomass on the artificial reef in October 2005 was <TEX>$1,990g/m^2$</TEX> at Biyang. In conclusion, a climax community and seaweed forest can be attained one year after the substrate is constructed.

Spatial patterns of macroalgal abundance in relation to eutrophication

Based on a large monitoring data set from Danish coastal waters we tested the hypotheses: (1) The vertical pattern of algal abundance is regulated by exposure in shallow water and by light limitation towards deeper water, resulting in a bell-shaped abundance curve, with peak abundance located deeper at more exposed sites, (2) in deeper water, total algal abundance and abundance of perennial algae decrease along a eutrophication gradient while (3) abundance and relative abundance of opportunists increase. The vertical pattern of algal abundance showed a peak at intermediate water depths which was located deeper in more exposed areas and thus confirmed our first hypothesis. For algae growing from depths of peak abundance and deeper, the study demonstrated that total algal abundance and abundance of perennials and opportunists at given depths decreased significantly along a eutrophication gradient and the relationships had high explanatory power (R 2 = 0.53–0.73). These results confirmed our second hypothesis. By contrast, the relative abundance of opportunists responded solely to salinity and was largest in the most brackish areas, in contradiction to hypothesis three. The lack of coupling between eutrophication and relative abundance of opportunists arises because both opportunists and the entire algal community were light limited and their ratio therefore relatively insensitive to changing water clarity. The analyses indicated that algal abundance initially responded slowly to increasing eutrophication but showed a more marked response at TN concentrations of 35–40 μM. However, the existence of possible threshold nutrient levels demands further analyses.

Macroalgal Communities Face the Challenge of Changing Biotic Interactions: Review with Focus on the Baltic Sea

In diverse littoral communities, biotic interactions play an important role in community regulation. This article reviews how eutrophication modifies biotic interactions in littoral macroalgal communities. Eutrophication causes blooms of opportunistic algae, increases epibiotism, and affects regulation by grazers. Opportunistic algae and epibionts harm colonization and growth of perennial algae. Grazing regulates the density and species composition of macroalgal communities, especially at the early stage of algal colonization. Eutrophication supports higher grazer densities by increasing the availability and quality of algae to grazers. This may, on the one hand, enhance the capability of grazers to regulate and counteract the increase of harmful, bloom-forming macroalgae; on the other hand, it may increase grazing pressure on perennial species, with a poor tolerance of grazing. In highly eutrophic conditions, bloom-forming algae may also escape grazing control and accumulate. Increasing epibiotism and grazing threaten in particular the persistence of habitat-forming perennials such as the bladderwrack. An interesting property of biotic interactions is that they do not remain fixed but are able to evolve, as the traits of the interacting species adapt to each other and to abiotic conditions. The potential of plants and grazers to adapt is crucial to their chances to survive in changing environment.

Macrophytobenthos of the Black Sea coast of the Kerch peninsular (Crimea, Ukraine)

The data on spatial distribution and qualitative and quantitative composition of macrophytobenthos are presented by the results of investigations carried out in 2004 at Cape Ak-Burun (Crimea: priority site Takyl; the Black Sea). Variations in values of indices along the coast and in the depths of the Black Sea are characterized. In all, 68 species of macroalgae were registered: Chlorophyta − 13 (19%), Phaeophyta − 19 (28%), and Rhodophyta − 42 (62%). The benthos vegetation biomass reaches 0.7 − 1 kg·m−2 in pseudolittoral (32 species of algae) and 1.5 − 7 kg·m−2 in sublittoral (64 species of algae). It is prudent to distinguish two subzones within the pseudolittoral zone. Algoflora is of a highly expressed oligosaprobic character. As to the quantity of species, the algae with short-vegetating periods are the dominants, as for biomass − the perennial algae predominate. The water area is recommended to be included into the Nature Reserve Fund.

Nutrient Addition to Experimental Rocky Shore Communities Revisited: Delayed Responses, Rapid Recovery

Coastal eutrophication may alter the dominance patterns of marine macroalgae, with potential consequences for the associated fauna and the entire ecosystem. Benthic macroalgae and animals in control and nutrient-enriched mesocosms were monitored to investigate eutrophication-induced changes in rocky shore communities. During a 3-year project, nutrient addition had only minor effects on the community structure, such as increased cover and biomass of green Ulva spp. and increased abundance of certain animal species at high nutrient levels. This study is a 4-year extension of a previously reported project, with 2 extra years of effect studies (altogether 5 years) and a subsequent 2 years for recovery. During the 4th year of nutrient enrichment, the cover of Fucus vesiculosus and Fucus serratus started to decline. In the 5th year, these canopy species crashed and there was an evident take-over by green algae at high nutrient addition levels. The previously observed abundance stimulation for fauna disappeared later in the time series, probably due to the loss of the macroalgal canopy. After less than 2 years on regular seawater, the algal and animal communities had returned to within the range of normal variability. The results indicate that established rocky shore communities of perennial algae with associated fauna are able to persist for several years, even at very high nutrient levels, but that community shifts may suddenly occur if eutrophication continues. They also indicate that rocky shore communities have the ability to return rapidly to natural undisturbed conditions after the termination of nutrient enhancement.

Effects of eutrophication, grazing, and algal blooms on rocky shores

Eutrophication can profoundly change rocky shore communities. These changes often cause the replacement of perennial, canopy‐forming algae such as Fucus spp. with annual, bloom‐forming algae such as Enteromorpha spp. Grazing, however, can counteract eutrophication by eliminating the annual algae’s susceptible recruits. We examine these generalizations across large scales. We use replicated “bioassay” experiments to compare the effects of eutrophication and grazing across four paired control versus eutrophied sites in the Northwest Atlantic and four eutrophied sites in the Baltic Sea in spring and summer. At each site, annual algal recruitment and grazing pressure were estimated using tiles seeded with Enteromorpha intestinalis propagules. Tiles were exposed for 3 weeks with grazers excluded or allowed access. Productivity of E. intestinalis recruits was strongly related to eutrophication (10‐fold increase) and grazing (80% decrease) and was weakly related to season. While the absolute grazing rate increased in a linear fashion with algal productivity, the relative grazing rate remained surprisingly constant (;80%). Comparative field surveys showed that perennial algae decreased by 30–60%, while annual algae, filter feeders, and grazers increased across a gradient of eutrophication. As eutrophication increased from control to eutrophied to point source sites, rocky shore communities became increasingly dominated by single species of annual algae or filter feeders, and community diversity declined consistently by 24–46%. We conclude that grazers are important controllers of algal blooms but that, ultimately, they cannot override the effects of increasing eutrophication on rocky shore community structure and biodiversity.

Biomass dynamics of exotic Sargassum muticum and native Halidrys siliquosa in Limfjorden, Denmark—Implications of species replacements on turnover rates

The expansion of Sargassum muticum in the Danish estuary Limfjorden between 1984 and 1997 was followed by a decrease in abundance of native perennial macroalgae such as Halidrys siliquosa. Although commonly associated with the expansion of exotic species, it is unknown whether such structural changes affect ecosystem properties such as the production and turnover of organic matter and associated nutrients. We hypothesized that S. muticum possesses ‘ephemeral’ traits relative to the species it has replaced, potentially leading to faster and more variable turnover of organic matter. The biomass dynamics of S. muticum and H. siliquosa was therefore compared in order to assess the potential effects of the expansion of Sargassum. The biomass of Sargassum was highly variable among seasons while that of Halidrys remained almost constant over the year. Sargassum grew faster than Halidrys and other perennial algae and the annual productivity was therefore high (P/B = 12 year −1) and exceeded that of Halidrys (P/B = 5 year −1) and most probably also that of other perennial algae in the system. The major grazer on macroalgae in Limfjorden, the sea urchin Psammechinus miliaris, preferred Sargassum to Halidrys, but estimated losses due to grazing were negligible for both species and most of the production may therefore enter the detritus pool. Detritus from Sargassum decomposed faster and more completely than detritus from Halidrys and other slow-growing perennial macrophytes. High productivity and fast decomposition suggest that the increasing dominance of S. muticum have increased turnover of organic matter and associated nutrients in Limfjorden and we suggest that the ecological effects of the invasion to some extent resemble those imposed by increasing dominance of ephemeral algae following eutrophication.

Growth and gonad production of the sea urchin Hemicentrotus pulcherrimus in the fucoid bed and algal turf in northern Japan

AbstractTo clarify the differences in the growth and gonad size of the sea urchin Hemicentrotus pulcherrimus among algal sere, the study was conducted in June and July, and September 1998 at three fucoid beds in Oga and at three algal turfs in Hachimori, Akita Prefecture in northern Japan. The most rapid growth was observed in a large perennial fucoid bed at the climax stage. Growth in a small perennial Chondrus ocellatus‐dominated bed in algal turfs in the seral stage was then high. The slowest growth was observed in the small perennial Dictyopteris divaricata‐ and Laurencia spp.‐dominated beds which are known to possess chemicals which act as feeding deterrents against the sea urchins. The gonad index (gonad wet weight × 100/body wet weight) at a fucoid bed was high where standing crops exceeded 3 kg·m−2. The gonad index in the Laurencia bed was lower than those at fucoid beds. These results suggest that growth and gonad production are affected by algal sere and differ among species of small perennial algae in the seral stage with or without chemical defense.

Animal diversity in Baltic rocky shore macroalgae: can Cladophora glomerata compensate for lost Fucus vesiculosus?

The substitution of canopy-forming perennial algae by annual filamentous algae is a common phenomenon on eutrophicated rocky shores with potential consequences for marine biodiversity. In the upper littoral of the northern Baltic Sea, this process is exemplified by decreased occurrence of the brown alga Fucus vesiculosus and increased amounts of filamentous algae (e.g. Cladophora glomerata and Pilayella littoralis). In this study, macrofauna communities of F. vesiculosus and C. glomerata during summer have been compared and the effects on littoral macroinvertebrates of a total or partial replacement of F. vesiculosus by C. glomerata have been estimated. The animal abundance and biomass were higher in C. glomerata than in corresponding amounts of F. vesiculosus (g dwt). Similar results were also obtained when animal abundance and biomass were extrapolated to algal cover per m 2 rocky substrate in the study area. Margalef's species richness was, on the other hand, higher in F. vesiculosus, whereas the Shannon–Wiener diversity was the same. The results indicate that C. glomerata maintains a high diversity of macroinvertebrates in the upper littoral zone of the northern Baltic Sea during summer. A possible substitution of F. vesiculosus by C. glomerata does not immediately affect macroinvertebrate diversity negatively in the littoral zone, although, it is difficult to assess the full consequences due to lack of comparable algal data at wider temporal and spatial scales.

Prolonged Lag in Population Outbreak of an Invasive Mussel: A Shifting-Habitat Model

Biological invasions pose a great threat to the integrity of natural communities. Some invasive species demonstrate a population explosion shortly after arrival while in other cases a prolonged lag between arrival and population outbreak is evident. This paper describes a case of a prolonged lag and explores the possible mechanism for this lag. The Red Sea mussel Brachidontes pharaonis, a Lessepsian migrant, was first recorded in the Mediterranean seven years after the opening of the Suez Canal in 1869. Since then it spread along the Israeli coast and as far northwest as Sicily. Studies conducted in the late 1970s, when B. pharaonis was still rare, predicted that it would not establish dense populations along the Israeli coast and would not outcompete the indigenous mussel Mytilaster minimus, although it has strong negative effects on survival and growth of the native species. It was attributed to the invader’s low intrinsic rate of increase relative to that of the native species, and to strong density-independent mortality generated by exposure to high wave action and sedimentation. In contrast to these predictions, we found massive formations of B. pharaonis beds after lag of about 120 years. We looked for distributional patterns that may explain this lag and found no south–north gradient but a strong habitat-dependent colonization. Most apparent are dense B. pharaonis mussel beds (density >300 per 100 cm 2 ) on rocky platforms where mussel beds were absent in the past. These platforms lack the vermetid rim that is typical to this formation. In platforms protected by a biogenic rim, sediment accumulation is high and perennial algae flourish. None of the mussel species form beds in such habitats. We suggest that the delayed formation of B. pharaonis beds along the Israeli coast is a consequence of a recent shift in habitat conditions on some platforms. It is possible that receding of the biogenic rim at the edge of these platforms allowed more effective washing, reduced sediment accumulation, and reduced perennial algae cover making platforms more suitable for the mussels. Lower density-independent mortality allowed B. pharaonis to dominate on such platforms over the indigenous species. On beachrock, a habitat previously dominated by M. minimus, we recorded a rapid shift in numerical domination to B. pharaonis (from 1 : 7 to 1.4 : 1 Brachidontes/Mytilaster individuals) over a period of 4 years (1995–1999). This is probably a result of saturation of the habitat by B. pharaonis recruits originating from the established populations on platforms. Salinity changes and a potential genetic shift may also have contributed to the invasive mussel outbreak.

Biomass response and changes in composition of ephemeral macroalgal assemblages along an experimental gradient of nutrient enrichment

Most conceptual marine eutrophication models predict that ephemeral and epiphytic macroalgae will become substantially more abundant with increasing nutrient richness. This expectation is based on the fact that most of these fast-growing algae possess high requirements for inorganic N and P and, therefore, suffer from nutrient limitation under nutrient poor conditions. We tested the hypothesis that nutrient enrichment will stimulate the abundance of ephemeral macroalgae by studying the abundance and species composition of these algae on several types of substrata along an artificial nutrient gradient. The total biomass of ephemeral macroalgae differed considerably among different types of substrata. The total ephemeral load was significantly higher on consolidated substrata than on large perennial algae, but we were unable to detect any systematic increase in the abundance of these ephemerals with increasing nutrient richness. We found, however, indications that the composition of the ephemeral assemblage changed with nutrient richness. Hence, corticated filamentous algae (mostly red algae) were more abundant at low nutrient richness while thin foliose algae (mainly green species) tended to become more abundant with increasing nutrient richness in most of the surveyed assemblages.

The Response of Experimental Rocky Shore Communities to Nutrient Additions

The aim of this study was to determine whether the experimental nutrient enrichment of littoral rocky shore communities would be followed by a predicted accumulation of fast-growing opportunistic algae and a subsequent loss of perennial benthic vegetation. Inorganic nitrogen (N) and potassium (P) was added to eight concrete mesocosms inhabited by established littoral communities dominated by fucoids. The response to nutrient enrichment was followed for almost 2 1/2 years. Fast-growing opportunistic algae (periphyton and ephemeral green algae) grew significantly faster in response to nutrient enrichment, but the growth of red filamentous algae and large perennial brown algae was unaffected. However, these changes were not followed by comparable changes in the biomass and composition of the macroalgae. The biomass of opportunistic algae was stimulated only marginally by the nutrient enrichment, and perennial brown algae (fucoids) remained dominant in the mesocosm regardless of nutrient treatment level. Established rocky shore communities thus seem able to resist the effects of heavy nutrient loading. We found that the combined effects of the heavy competition for space and light imposed by canopy-forming algae, preferential grazing on opportunistic algae by herbivores, and physical disturbance, succeeded by a marked export of detached opportunistic algae, prevented the fast-growing algae from becoming dominant. However, recruitment studies showed that the opportunistic algae would become dominant when free space was available under conditions of high nutrient loading and low grazing pressure. These results show that established communities of perennial algae and associated fauna in rocky shore environments can prevent or delay the accumulation of bloom-forming opportunistic algae and that the replacement of long-lived macroalgae by opportunistic species at high nutrient loading may be a slow process. Nutrient enrichment may not, in itself, be enough to stimulate structural changes in rocky shore communities.

Temporal variation and succession in an algal-dominated high intertidal assemblage

We determined whether temporal variation and succession were similar among sites with similar species composition by sampling unmanipulated and cleared plots in a high intertidal assemblage dominated by Endocladia muricata and Mastocarpus papillatus. Sampling was done for 6 years at six sites spanning over 4° of latitude in California. Ten 1×2-m permanent plots were chosen in the central portion of the assemblage at each site. Four of these served as unmanipulated controls, three were cleared (scraped and burned) in the spring of 1985, and three were cleared in the fall of 1985. The cover of sessile and density of motile species were determined by subsampling within the plots from 1985 until 1991. Recovery of the clearings was determined by their similarity to the controls. The algae E. muricata, M. papillatus, and Fucus gardneri, and the barnacle Balanus glandula, were the most abundant sessile organisms in the control plots, although the latter never exceeded 12% cover at any site. The grazing gastropods Littorina scutulata/ plena, various limpets, and Tegula funebralis were the most common mobile organisms. The species composition of the common species remained constant in the control plots over the study period and there were few large changes in relative abundance. Significant seasonal variation was detected in 11 species but variation was commonly site-specific. Ephemeral algae were abundant during early succession at only two of the six sites, and barnacle cover was low (<15% cover) at four sites and moderate (15–50% cover) at the remaining two throughout succession. Recovery rate varied considerably among sites and between times of clearing (1–10%/month). Correlations between ephemeral algae and grazer abundance, and between these variables and recovery rate were not significant. The effects of grazers on recovery rate were only evident at one site where they appeared to reduce an initially high cover of ephemeral algae and delay the establishment of perennials. Some of the largest differences in recovery rate were between clearing times, associated with differences in the phenology of the dominant perennial algae. In spite of these differences, most plots recovered by the end of the study period. These results indicate that the assemblages in the control plots at each site were relatively stable and, while successional pathways and processes varied, the assemblage at most sites still recovered. Current models, based largely on biological interactions, that attempt to explain within assemblage structure and succession were not broadly applicable.

LONG‐TERM CHANGES IN THE MACROALGAL VEGETATION OF THE INNER GULLMAR FJORD, SWEDISH SKAGERRAK COAST1

We examined long‐term changes in the macroalgal vegetation at Stora Bornö Island in the inner Gullmar Fjord on the Swedish Skagerrak coast. This was made possible by access to a 1941 diving investigation. The same sites were reinvestigated in 1998. Community composition and depth distributions of species were compared and changes were analyzed with focus on functional groups (size, thallus shape, and life‐history traits). We discovered a significant decrease in the depth extension of macroalgal species and a dramatic decline of species richness in the lower littoral (below 16 m of depth) compared with 57 years earlier. Ordination analysis revealed that there was a significant difference in the community composition between the two study periods. In general, small (&lt;10 cm), thin, filamentous, and aseasonal ephemerals increased in relative abundance, whereas larger (&gt;10 cm), coarsely branched, and perennial algae decreased. Calibrations of individual species to local sediment cover, using canonical correspondence analysis, indicated that part of the change in species composition was related to sediment load. Furthermore, large‐scale climate differences (NAO Winter Index) between the study periods indicated a higher impact of Baltic Sea and Kattegat water in the nutrient dynamics of the fjord in the 1998 study. We concluded that the observed long‐term changes in the macroalgal community at Stora Bornö Island were consistent with an increased nutrient availability.

The effects of grazing on the distribution and composition of low-shore algal communities on the central coast of Portugal and on the southern coast of Britain

A red algal turf is often found just below the barnacle/limpet zone of many European shores, especially on steep shores of moderate exposure. The hypothesis that grazing by limpets determines the upper limit of distribution of this red algal turf was tested on moderately exposed shores in Portugal and Britain. We also aimed to assess whether the grazing effect is modified at various spatial scales. Grazers were excluded by fences, with half-fenced and unfenced controls. Exclusion plots were rapidly colonised by green ephemeral algae in the months immediately after the beginning of the experiment (summer); these algae were later replaced by perennial algae. The percentage cover of turf-forming macroalgae showed a significant increase at both locations. The upper limit of distribution extended more than 50 cm on most of the shores studied. In contrast, control and half-fenced plots remained devoid of algae. After 2 years, ungrazed plots were mainly colonised by a red algal turf (e.g. Caulacanthus ustulatus, Gelidium spp., Laurencia pinnatifida) in Portugal, while canopy cover ( Fucus serratus and Himanthalia elongata) dominated in Britain in marked contrast to the grazed plots. Physical factors acting at both local and geographical scales may explain these differences. However, although physical factors probably have an important influence on the identity, size and abundance of sublittoral fringe macroalgae, grazers play a major role in directly setting their upper limits. The effect of grazing by limpets was not consistent for all of the morphological algal groups and spatial scales considered in the present study. The effect of grazing on the cover of turf algae varied between Portugal and Britain (location scale), while effects on ephemeral and canopy algal cover varied at the shore scale within location.

Constant short-day treatment of outdoor-cultivated Laminaria digitata prevents summer drop in growth rate

Previous laboratory studies in species of the Laminariales have revealed that both the onset of growth in early winter and the summer drop in growth rate are controlled by the annual course of daylength synchronizing endogenous, circannual clocks within the thallus. Moreover, it is known for some laminarian species that cultivation in the laboratory in constant short days (SD) leads to arhythmic, continuous growth activity of the blade throughout the year. Such a prolonged SD treatment has now been performed for the first time in outdoor-cultivated Laminaria digitata. Field-grown sporophytes were collected in May from the sea near Helgoland (North Sea) and cultivated on the island of Sylt (North Sea) in temperature-controlled outdoor tanks (300 l) at 10°C for 1·5 years either in a constant 8 h of light per day, controlled by an automatic blind on top of the tank, or in ambient daylengths. In constant SD, the growth rate remained at a steady-state level of approximately 0·4 cm day−1 from the end of June until mid-October, whereas growth rate in ambient daylengths declined steadily after June to half the rate in SD in October. Growth became light-limited between October and February in both treatments and, in the second year from July onwards, higher growth rates were again observed in constant SD than in ambient daylengths. Further work is required to find out whether SD treatment in summer would also prevent the summer drop in growth rate in other perennial seaweed species, e.g. commercially valuable red algae. This would potentially increase the chance of more constant biomass production throughout the year, and decrease the danger of the cultivated perennial algae being overgrown by annual epiphytes in summer.

Two reproductive strategies in Baltic Fucus vesiculosus (Phaeophyceae)

Marked seasonal reproduction is a widespread phenomenon in seaweeds in temperate waters strongly influenced by seasonal variations. In the Baltic Sea, however, the perennial brown alga Fucus vesiculosus exhibits two periods of reproduction: early summer (May–June) or late autumn (September–November). In this study, we compared the geographical distribution, the reproductive biology and the recruitment success for F. vesiculosus that reproduce during either of these two periods on the east coast of Sweden. Both monocultures and mixed populations of summer- and autumn-reproducing F. vesiculosus occurred along the mainland coast from the southern parts of the Stockholm archipelago to the southernmost part of Sweden. F. vesiculosus reproduced only during summer around the coasts of the large islands of Oland and Gotland. After reproduction, in July and January respectively, plants that reproduced in summer abscised their receptacles and part of their branches, while plants reproducing in autumn abscised only ...

Algal propagule banks modify competition, consumer and resource control on Baltic rocky shores.

We hypothesized that supply from macroalgal propagule banks may influence the relative abundance of annual and perennial algae and that this may alter the effects of grazers and nutrients on species composition. In a factorial field experiment in the Baltic Sea littoral system we tested the effects of manipulating propagule banks, the abundance of crustacean and gastropod grazers, and nutrient supply on recruitment and growth of macroalgae over a year. Moreover, we determined seasonal patterns of macroalgal propagule dispersal at the experimental site and quantified algal abundance and recruitment at 25 locations throughout the Baltic Sea. Experimental manipulations had minor effects on adults of the dominating perennial alga, Fucus vesiculosus. Instead, we found that species composition was determined by processes operating at early life stages. Propagule supply from a propagule bank strongly favored the fast-growing annual alga Enteromorpha spp. which then blocked settlement and recruitment of Fucus. Grazers reduced the abundance of annual algae and indirectly favored Fucus recruitment. There was an apparent trade-off between gains from the propagule bank and losses to herbivory in five of seven colonizing species. Nutrient enrichment overrode grazer control of annual algae and accelerated the decline of Fucus only when annual algae had already achieved high densities through the propagule bank. Corroborating the experimental findings, field surveys across the Baltic showed that Fucus recruit densities can be predicted from the cover of annual algae during the period of Fucus reproduction and settlement. Recruitment inhibition by annual algae, which is driven by the abundance of annuals in the propagule bank, increasing nutrient levels, and declining consumer control, is suggested as a mechanistic explanation of the current decline of perennial algae in the Baltic Sea.

Long-term changes in macroalgal communities in a Danish estuary

Long-term studies of the composition of marine macroalgal communities are rare because of the lack of quantitative investigations in the past. In Denmark, a single quantitative study was performed in 1941–1943 in the nontidal estuary Isefjord–Roskilde Fjord, at several sites of variable salinity (10–22 ‰) and nutrient availability. This study was used as a reference for evaluating changes in the richness and relative abundance of macroalgal species from different taxonomic groups and functional forms at sites experiencing increasing concentrations of nitrate and light attenuation during the 50 years elapsed since then. Qualitative measures showed no, or few, changes over the period. In contrast, quantitative measures showed a significant increase in the relative abundance of small, opportunistic green algae (e.g. species of Chaetomorpha, Cladophora, Enteromorpha and Ulva) and a significant decrease in the relative abundance of large, perennial brown algae (e.g. Fucus serratus and F. vesiculosus). Along with these changes, the diversity of functional form groups declined. The inner parts of Roskilde Fjord, which were already nutrient-rich 50 years ago, did not show any significant changes in the composition of the macroalgal communities. The temporal changes during the 50 years are consistent with the spatial differences observed both then and now along the nutrient gradients. The results imply that quantitative measures of the abundance of macroalgal species having different thallus form, longevity and taxonomy are more sensitive and robust than qualitative measures in reflecting changes in the communities brought about by cultural eutrophication. The response of macroalgal communities gradually diminishes with progressively higher nutrient loadings, suggesting that the impact of eutrophication on coastal macroalgae will follow a geometric series similar to that established for lakes of different trophic levels.