Despite marine traits,the endemic Fucus radicans (Phaeophyceae) is restricted to the brackish Baltic Sea

Many of the marine species that were introduced to the Baltic Sea during the Littorina stage (c. 8500–3000 years BP), e.g. Fucus vesiculosus and F. serratus, have adapted to the present low salinity. These marine species have gone from marine conditions into lower salinity environments. In this paper we ask why the recently discovered endemic brown alga Fucus radicans shows the opposite pattern. Fucus radicans is only present in the northern parts of the Baltic Sea, the low salinity Bothnian Sea (4–6 psu). Potentially, the fitness of F. radicans might be reduced in higher salinities if it is better adapted to brackish conditions. We hypothesize, however, that the southern distribution limit of F. radicans is set by biotic factors, e.g. competition with F. vesiculosus and higher grazing pressure by Idotea balthica and not by salinity. Our results show that the reproductive output of F. radicans is limited by low salinity (4 psu) but increases in higher salinities. However, the southern distribution limit, i.e. the northern Baltic Proper, is regulated by biotic factors, where the additive effects from shading by taller F. vesiculosus thalli and grazing on F. radicans by the isopod I. balthica limit the biomass production of F. radicans. We suggest that F. radicans still maintains marine traits due to its ability to propagate clonally and is restricted to the Bothnian Sea by interactions with F. vesiculosus and I. balthica. We also propose that increased precipitation due to climate change might affect the northern range limit and that the distribution of F. radicans could be expected to shift further south into the Baltic Proper.     

Geographic variation in fitness‐related traits of the bladderwrack Fucus vesiculosus along the Baltic Sea‐North Sea salinity gradient

In the course of the ongoing global intensification and diversification of human pressures, the study of variation patterns of biological traits along environmental gradients can provide relevant information on the performance of species under shifting conditions. The pronounced salinity gradient, co‐occurrence of multiple stressors, and accelerated rates of change make the Baltic Sea and its transition to North Sea a suitable region for this type of study. Focusing on the bladderwrack Fucus vesiculosus, one of the main foundation species on hard‐bottoms of the Baltic Sea, we analyzed the phenotypic variation among populations occurring along 2,000 km of coasts subjected to salinities from 4 to >30 and a variety of other stressors. Morphological and biochemical traits, including palatability for grazers, were recorded at 20 stations along the Baltic Sea and four stations in the North Sea. We evaluated in a common modeling framework the relative contribution of multiple environmental drivers to the observed trait patterns. Salinity was the main and, in some cases, the only environmental driver of the geographic trait variation in F. vesiculosus. The decrease in salinity from North Sea to Baltic Sea stations was accompanied by a decline in thallus size, photosynthetic pigments, and energy storage compounds, and affected the interaction of the alga with herbivores and epibiota. For some traits, drivers that vary locally such as wave exposure, light availability or nutrient enrichment were also important. The strong genetic population structure in this macroalgae might play a role in the generation and maintenance of phenotypic patterns across geographic scales. In light of our results, the desalination process projected for the Baltic Sea could have detrimental impacts on F. vesiculosus in areas close to its tolerance limit, affecting ecosystem functions such as habitat formation, primary production, and food supply.     

Photosynthesis and UV-B Tolerance of the Marine Alga Fucus vesiculosus at Different Sea Water Salinities

The marine algal species in the Baltic Sea are few due to the low sea water salinity. One of the few species that can be found is Fucus vesiculosus. Even this species is affected by the low salinity and becomes smaller in size in the Baltic. In present work the photosynthesis of F. vesiculosus in the northern Baltic (Bothnian Sea) was compared to the photosynthesis of F. vesiculosus in the Atlantic. Oxygen evolution was measured before and after exposure to 2.3 W of UV-B (280–320 nm) radiation for 5 h, as well as after 48 h recovery in low light. The plants were kept in their own sea water salinity as well as in a changed salinity, this to examine possible correlations between salinity and photosynthesis. The results show a significant higher initial maximal photosynthesis (P _max) for Atlantic plants (10.3 nmol O₂ g⁻¹ FW s⁻¹) compared to Baltic plants (4.0 nmol O₂ g⁻¹ FW s⁻¹). The Baltic plants were found more sensitive to UV-B with a 40–50% decrease of P _max as well as a lower degree of recovery (60–70% compared to 75–95% for the Atlantic plants). The higher salinity (35 psu) had a positive effect on the Baltic F. vesiculosus with increased P _max as well as increased tolerance to UV-B. The lower salinity (5 psu) had a negative effect on the Atlantic plants with a decreased P _max as well as a lower tolerance to UV-B. Pigment content was found higher in Atlantic F. vesiculosus. The pigment content decreased then the Atlantic plants were transferred to 5 psu. The concentration of Chl a as well as the total content of violaxanthin, diadinoxanthin and zeaxanthin in Baltic plants increased when transferred to 35 psu. The Atlantic F. vesiculosus can not survive the low salinity in the northern Baltic (died within 8 weeks). It is likely that a long time acclimation or adaptation to low salinity has taken place for F. vesiculosus in northern Baltic. If this is an ecotypic or genotypic development it is too early to say.     

FLUORESCENCE EMISSION SPECTRA OF MARINE AND BRACKISH‐WATER ECOTYPES OF FUCUS VESICULOSUS AND FUCUS RADICANS (PHAEOPHYCEAE) REVEAL DIFFERENCES IN LIGHT‐HARVESTING APPARATUS1

The Bothnian Sea in the northerly part of the Baltic Sea is a geologically recent brackish‐water environment, and rapid speciation is occurring in the algal community of the Bothnian Sea. We measured low‐temperature fluorescence emission spectra from the Bothnian Sea and the Norwegian Sea ecotypes of Fucus vesiculosus L., a marine macroalga widespread in the Bothnian Sea. Powdered, frozen thallus was used to obtain undistorted emission spectra. The spectra were compared with spectra measured from the newly identified species Fucus radicans Bergström et L. Kautsky, which is a close relative of F. vesiculosus and endemic to the Bothnian Sea. The spectrum of variable fluorescence was used to identify fluorescence peaks originating in PSI and PSII in this chl c–containing alga. The spectra revealed much higher PSII emission, compared to PSI emission, in the Bothnian Sea ecotype of F. vesiculosus than in F. radicans or in the Norwegian Sea ecotype of F. vesiculosus. The results suggest that more light‐harvesting chl a/c proteins serve PSII in the Bothnian Sea ecotype of F. vesiculosus than in the two other algal strains. Treatment of the Bothnian Sea ecotype of F. vesiculosus in high salinity (10, 20, and 35 practical salinity units) for 1 week did not lead to spectral changes, indicating that the measured features of the Bothnian Sea F. vesiculosus are stable and not simply a direct result of exposure to low salinity.     

LOCAL ADAPTATION OF CERAMIUM TENUICORNE (CERAMIALES,RHODOPHYTA) WITHIN THE BALTIC SEA1

Ceramium tenuicorne (Kützing) Wærn is a red alga that is widely distributed in the brackish Baltic Sea. We studied the growth response of Ceramium to low salinity and nutrient enrichment, using isolades from two regions of the Baltic Sea where the alga approaches its inner distribution limit. Ecotypic differentiation was observed in that differences in growth response among isolates corresponded to salinity conditions in their regions of origin. Isolates from the Gulf of Bothnia (4 psu) had inherently lower growth rates that were not increased when transferred to higher salinity, but were better adapted to very low salinity levels than isolates from the Baltic Proper (7 psu). Further, the results indicate that Ceramium from different regions of the Baltic Sea vary in their responses to nutrient enrichment. The observed differences may be best described as a quantitative difference in the proportion of isolates with hyposaline adaptation. The results indicate that the wide distribution of Ceramium in the Baltic Sea is better explained by the occurrence of locally adapted genotypes than by a generalist life strategy, and provide example of adaptive differentiation in a marine edge environment.     

Distributional success of the marine seaweedFucus vesiculosus L. in the brackish Baltic Sea correlates with osmotic capabilities of Baltic gametes

To understand the unique success of the marine seaweedFucus vesiculosus L. (PHaeophyceae) in the brackish Baltic Sea, the performance of gametes from Baltic [4.1–6.5S (Salinity)] and marine populations was studied. Sperm from BalticF. vesiculosus swam with a path velocity of c. 30–110 m/s and could fertilize eggs in waters of salinities from 4 to 33S. In their natural water, Baltic sperm were not negatively phototactic, unlike marine sperm in seawater; this should decrease the sperm:egg concentration at the seafloor and reduce the likelihood of polyspermy. Marine (Iceland, Sweden) sperm in seawater had a path velocity of c. 80–100 m/s, but performed poorly and could not fertilize eggs in natural or artificial Baltic water 6S; therefore, Baltic populations have adapted or acclimated to their brackish habitat. Baltic populations appear better adapted to their natural low salinities because, even after culturing Baltic and marine individuals in water from both the Baltic (6.5S) and the marine Skagerrak (21S), Baltic sperm were in both cases still able to swim and fertilize eggs at lower salinities (4S) than marine sperm; fertilization never occurred between marine gametes at 4–6S. However,F. vesiculosus acclimates to some salinities, since sperm from Baltic and marine males that had been cultured at 21S swam better (higher velocity, proportion that were motile and/or linearity) in marine salinities (21–33S) than when they were cultured at 6.5S. The effects of salinity on sperm motility and fertilization were osmolar rather than due to specific ionic requirements, over the tested range. The osmolalities (< c. 100 mmol/kg) at which fertilization success of Baltic gametes decreases nearly to zero correspond to the osmolality of Baltic water at the northernmost limit of distribution ofF. vesiculosus in the Baltic Sea. Therefore, the present range ofF. vesiculosus in the Baltic appears to correspond to the osmotic tolerance of the gametes. Very small natural or anthropogenic increases in ambient osmolality would be likely to cause a substantial expansion of this species into the inner Baltic.     

Letters to the editor

A decrease in salinity and temperature over the past 3000 years has presented the marine algae of the Baltic Sea with very considerable problems in adaptation. The effects of salinity upon a number of Baltic algae have been measured. The results showed cell mortality to be severe in 0, 68 and 102‰, and minimal in 6 and 11‰: there was most variation in tolerance to 34 and 51‰. The salt tolerances of Baltic marine algae have proved more hyposaline than those of British intertidal algae. Water uptake and loss in tissues of Chorda filum and Fucus vesiculosus from Baltic and British populations have been measured in response to salinity changes. The results revealed significant population differences in both live and killed tissues. Receptacle development and oogonial maturation have been observed in Baltic and British F. vesiculosus, and found to differ in seasonality. Some observations were associated with local sea temperatures but differences in the timing of receptacle initiation and in oogonial size were not. Th depauperate thallus, commonly ascribed to the effects of low salinity, was found to be a complicated phenomenon, comprising numerous attributes which are combined differently in different taxa. The morphological differences between Baltic and British marine algae were usually striking.

The marine algae of the Baltic Sea have therefore diverged in a number of ways from their N. Atlantic counterparts. The naturally high variability of these taxa has enabled them to survive the period of increasingly strong selection pressure which followed the Littorina Sea episode. Divergence seems not to have advanced to the point where speciation may be said to have occurred. The Baltic may therefore be contrasted with the much older Mediterranean Sea, which contains a large number of endemic species. Nevertheless, the Baltic is a site of very considerable evolutionary importance.     

GENETIC STRUCTURE IN POPULATIONS OF FUCUS VESICULOSUS (PHAEOPHYCEAE) OVER SPATIAL SCALES FROM 10 M TO 800 KM1

Recent studies showing consequences of species’ genetic diversity on ecosystem performance raise the concern of how key ecosystem species are genetically structured. The bladder wrack Fucus vesiculosus L. is a dominant species of macroalga in the northern Atlantic, and it is particularly important as a habitat‐forming species in the Baltic Sea. We examined the genetic structure of populations of F. vesiculosus with a hierarchical approach from a within‐shore scale (10 m) to a between‐seas scale (Baltic Sea–Skagerrak, 800 km). Analysis of five microsatellite loci showed that population differentiation was generally strong (average F_ST = 12%), being significant at all spatial scales investigated (10¹, 10³, 10^4–5, 10⁶ m). Genetic differentiation between seas (Baltic Sea and Skagerrak) was substantial. Nevertheless, the effects of isolation by distance were stronger within seas than between seas. Notably, Baltic summer‐reproducing populations showed a strong within‐sea, between‐area (70 km) genetic structure, while Baltic autumn‐reproducing populations and Skagerrak summer‐reproducing populations revealed most genetic diversity between samples within areas (<1 km). Despite such differences in overall structure, Baltic populations of summer‐ and autumn‐reproducing morphs did not separate in a cluster analysis, indicating minor, if any, barriers to gene flow between them. Our results have important implications for management and conservation of F. vesiculosus, and we raise a number of concerns about how genetic variability should be preserved within this species.     

Photosynthesis in relation to D1, PsaA and Rubisco in marine and brackish water ecotypes of Fucus vesiculosus and Fucus radicans (Phaeophyceae)

The aim of this study was to investigate photosynthetic differences between the marine, Norwegian Sea ecotype and the brackish, Bothnian Sea ecotype of F. vesiculosus and F. radicans and to see whether photosynthetic differences could be connected with the relative amounts of D1 protein (PSII), PsaA (PSI) protein and/or Rubisco. For this purpose, we tested if a higher photosynthetic maximum (P _max) in the Atlantic Ocean ecotype of F. vesiculosus relative to the Baltic Sea ecotype, and an increase of the P _max in Baltic Sea ecotype of F. vesiculosus at higher salinity, could be due to an increase in the relative amounts of Rubisco. The proteins have been evaluated on a relative basis. Immunoblot signals showed that the amount of Rubisco was higher in both ecotypes of F. vesiculosus than in F. radicans, but no differences could be detected between the two ecotypes of F. vesiculosus. The results suggest an uneven photosystem protein stoichiometry in Fucus, with more of the PSI protein PsaA relative to the PSII protein D1. The difference in P _max between the two ecotypes of F. vesiculosus might be related to the difficulties for the algae to adapt to the environment in Bothnian Sea.     

Effects of filamentous algae and deposited matter on the survival of Fucus vesiculosus L. germlings in the Baltic Sea

As a result of increased nutrient levels in the Baltic Sea during thepast 50 years, mass developments of filamentous algae have become a commonfeature along the Swedish east coast and deposition of organic matter has alsoincreased. To test whether these two factors have any effects on the early lifestages of Fucus vesiculosus a number of laboratory andfield studies were conducted. The amount of epilithic and epiphytic filamentousalgae on F. vesiculosus and the amount of deposited matterin the littoral zone were quantified during the two reproductive periods ofF. vesiculosus, early summer (May–June) and lateautumn (September–October). Both filamentous algae (Cladophoraglomerata) and deposited matter (introduced either before or aftersettlement of fertilized eggs) were shown to significantly decrease the numberof surviving germlings. The survival of germlings seeded on stones withfilamentous algae, or seeded on culture dishes concurrently with the lowestconcentration of deposited matter (0.1 g dm^–2),was 5% or less. In the field, the amount of filamentous algae was significantlyhigher during F. vesiculosus summer reproduction, whereasthe amount of deposited matter collected in traps was significantly higherduring the period of autumn reproduction. The greatest biomass of filamentousalgae was observed at sheltered sites. Based on the negative effects offilamentous algae and deposited matter on Fucusrecruitmentand the observation of local and seasonal differences in abundance offilamentous algae and deposition, we suggest that the prerequisites for thesurvival of either summer or autumn-reproducing populations of F.vesiculosus in the Baltic Sea may differ locally.     

Local adaptation and oceanographic connectivity patterns explain genetic differentiation of a marine diatom across the North Sea–Baltic Sea salinity gradient

Drivers of population genetic structure are still poorly understood in marine micro‐organisms. We exploited the North Sea–Baltic Sea transition for investigating the seascape genetics of a marine diatom, Skeletonema marinoi. Eight polymorphic microsatellite loci were analysed in 354 individuals from ten locations to analyse population structure of the species along a 1500‐km‐long salinity gradient ranging from 3 to 30 psu. To test for salinity adaptation, salinity reaction norms were determined for sets of strains originating from three different salinity regimes of the gradient. Modelled oceanographic connectivity was compared to directional relative migration by correlation analyses to examine oceanographic drivers. Population genetic analyses showed distinct genetic divergence of a low‐salinity Baltic Sea population and a high‐salinity North Sea population, coinciding with the most evident physical dispersal barrier in the area, the Danish Straits. Baltic Sea populations displayed reduced genetic diversity compared to North Sea populations. Growth optima of low salinity isolates were significantly lower than those of strains from higher native salinities, indicating local salinity adaptation. Although the North Sea–Baltic Sea transition was identified as a barrier to gene flow, migration between Baltic Sea and North Sea populations occurred. However, the presence of differentiated neutral markers on each side of the transition zone suggests that migrants are maladapted. It is concluded that local salinity adaptation, supported by oceanographic connectivity patterns creating an asymmetric migration pattern between the Baltic Sea and the North Sea, determines genetic differentiation patterns in the transition zone.     

FREQUENT CLONALITY IN FUCOIDS (FUCUS RADICANS AND FUCUS VESICULOSUS; FUCALES,PHAEOPHYCEAE) IN THE BALTIC SEA1

Asexual reproduction by cloning may affect the genetic structure of populations, their potential to evolve, and, among foundation species, contributions to ecosystem functions. Macroalgae of the genus Fucus are known to produce attached plants only by sexual recruitment. Recently, however, clones of attached plants recruited by asexual reproduction were observed in a few populations of Fucus radicans Bergström et L. Kautsky and F. vesiculosus L. inside the Baltic Sea. Herein we assess the distribution and prevalence of clonality in Baltic fucoids using nine polymorphic microsatellite loci and samples of F. radicans and F. vesiculosus from 13 Baltic sites. Clonality was more common in F. radicans than in F. vesiculosus, and in both species it tended to be most common in northern Baltic sites, although variation among close populations was sometimes extensive. Individual clonal lineages were mostly restricted to single or nearby locations, but one clonal lineage of F. radicans dominated five of 10 populations and was widely distributed over 550 × 100 km of coast. Populations dominated by a few clonal lineages were common in F. radicans, and these were less genetically variable than in other populations. As thalli recruited by cloning produced gametes, a possible explanation for this reduced genetic variation is that dominance of one or a few clonal lineages biases the gamete pool resulting in a decreased effective population size and thereby loss of genetic variation by genetic drift. Baltic fucoids are important habitat‐forming species, and genetic structure and presence of clonality have implications for conservation strategies.     

In situ study of relative electron transport rates in the marine macroalga <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> in the Baltic Sea at different depths and times of year

The brown alga Fucus vesiculous is one of the few marine species in the Baltic Sea. Fucus vesiculosus shows high morphological and physiological variability as a response to its environmental conditions. The salinity in the Baltic Sea is 4–5 psu, compared to 35 psu in the Atlantic. Photosynthesis of algae is usually measured after collection and transportation to constant culture conditions. However, in this study, relative photosynthetic electron transport rates, calculated from chlorophyll a fluorescence parameters were compared in algae collected from 1 and 4 m depths by SCUBA divers. Measurements of light response curves from the same individuals of F. vesiculosus at different depths and times of the year have, to our knowledge, not been made previously. Measurements were performed on four different occasions during the spring of 2005 (25 February, 3 and 29 April, and 26 May) in the Baltic Sea, using rapid light curves generated with a Diving PAM. In addition, samples were collected for photoinhibition studies in the laboratory. The light response curves obtained in situ at 1 and 4 m depths for F. vesiculosus showed lower values of light saturation with depth. When algae from 1 and 4 m depths were exposed to high irradiances of photosynthetically active radiation (1,400 μmol photons m⁻² s⁻¹), algae from 1 m depth showed a higher degree of photoinhibition in comparison to algae from 4 m depth.     

Diversity of Aphanizomenonflos-aquae (Cyanobacterium) Populations along a Baltic Sea Salinity Gradient

Colony-forming cyanobacteria of the genus Aphanizomenon form massive blooms in the brackish water of the Baltic Sea during the warmest summer months. There have been recent suggestions claiming that the Baltic Sea Aphanizomenon species may be different from Aphanizomenon flos-aquae found in lakes. In this study, we examined variability in the morphology and 16S-23S rRNA internal transcribed spacer (ITS) sequences of A. flos-aquae populations along a salinity gradient from a string of lakes to a fjord-like extension of the Baltic Sea to the open Baltic Sea. Morphological differences among the populations were negligible. We found that the Baltic Sea was dominated (25 out of 27 sequences) by one ITS1-S (shorter band of ITS 1 [ITS1]) genotype, which also was found in the lakes. The lake populations of A. flos-aquae tended to be genetically more diverse than the Baltic Sea populations. Since the lake ITS1-S genotypes of A. flos-aquae are continuously introduced to the Baltic Sea via inflowing waters, it seems that only one ITS1 genotype is able to persist in the Baltic Sea populations. The results suggest that one of the ITS1-S genotypes found in the lakes is better adapted to the conditions of the Baltic Sea and that natural selection removes most of the lake genotypes from the Baltic Sea A. flos-aquae populations.     

Density dependent grazing effects of the isopod Idotea baltica Pallas on Fucus vesiculosus L in the Baltic Sea

In the Baltic Sea, abiotic factors are often supposed to explain the distribution of the key species Fucus vesiculosus. Still, in many areas, decline of F. vesiculosus has coincided with mass occurrence of the herbivorous isopod Idotea baltica. The aim of this work was to examine whether, how and at what densities I. baltica can affect the distribution of F. vesiculosus in the central Baltic proper. Both large-scale field surveys and a two-week grazing experiment have been performed.In the field survey there was a correlation between density of I. baltica and reduction in depth penetration of F. vesiculosus. At 80 animals per 100 g F. vesiculosus wet weight, the depth penetration of the F. vesiculosus belt was reduced by 2.5 m within a year. In the grazing experiment there was a correlation between density of I. baltica and loss of F. vesiculosus biomass and meristems. In the controls biomass and number of meristems increased by 50%, while at 20 animals per 100 g of F. vesiculosus there was no net growth of F. vesiculosus. Intensity of grazing did not differ between isopod densities of 20, 40 and 60/100 g. At isopod densities of 80 and 100/100 g though, biomass and meristems decreased by 50%, indicating a threshold for the survival of F. vesiculosus in the experiment. At all densities the isopods preferred younger tissue to older.Our results indicate that grazing by Idotea baltica is an important structuring factor in the Baltic Fucus vesiculosus populations.     

Fitness declines towards range limits and local adaptation to climate affect dispersal evolution during climate‐induced range shifts

Dispersal ability will largely determine whether species track their climatic niches during climate change, a process especially important for populations at contracting (low‐latitude/low‐elevation) range limits that otherwise risk extinction. We investigate whether dispersal evolution at contracting range limits is facilitated by two processes that potentially enable edge populations to experience and adjust to the effects of climate deterioration before they cause extinction: (i) climate‐induced fitness declines towards range limits and (ii) local adaptation to a shifting climate gradient. We simulate a species distributed continuously along a temperature gradient using a spatially explicit, individual‐based model. We compare range‐wide dispersal evolution during climate stability vs. directional climate change, with uniform fitness vs. fitness that declines towards range limits (RLs), and for a single climate genotype vs. multiple genotypes locally adapted to temperature. During climate stability, dispersal decreased towards RLs when fitness was uniform, but increased when fitness declined towards RLs, due to highly dispersive genotypes maintaining sink populations at RLs, increased kin selection in smaller populations, and an emergent fitness asymmetry that favoured dispersal in low‐quality habitat. However, this initial dispersal advantage at low‐fitness RLs did not facilitate climate tracking, as it was outweighed by an increased probability of extinction. Locally adapted genotypes benefited from staying close to their climate optima; this selected against dispersal under stable climates but for increased dispersal throughout shifting ranges, compared to cases without local adaptation. Dispersal increased at expanding RLs in most scenarios, but only increased at the range centre and contracting RLs given local adaptation to climate.     

GENETIC AND MORPHOLOGICAL IDENTIFICATION OF FUCUS RADICANS SP. NOV. (FUCALES,PHAEOPHYCEAE) IN THE BRACKISH BALTIC SEA1

Brown seaweeds of the genus Fucus occupy a wide variety of temperate coastal habitats. The genus is evolutionary dynamic with recent radiations to form morphologically distinct taxa. In the brackish Baltic Sea, fucoids are the only perennial canopy‐forming macroalgae. The most northern populations of Fucus occur permanently submerged in extremely low salinity (3–5 psu). These are currently referred to as Fucus vesiculosus L. but are morphologically distinct with a narrow frond without bladders. We report here that a population of this unique morphotype is reproductively isolated from a truly sympatric population of common F. vesiculosus and conclude that the northern morphotype represents a previously undescribed species. We describe Fucus radicans sp. nov., which is attached and dioecious with broadly elliptic receptacles, characterized by a richly branched narrow flat frond (2–5 mm), short thallus (<26 cm), and a high capacity for vegetative recruitment of attached plants. Analysis of five highly polymorphic microsatellite DNA loci showed genetic differentiation between sympatric populations of F. radicans and F. vesiculosus, whereas allopatric populations of the same species revealed a coherent pattern of genetic variation. Sequences of the RUBISCO region in F. radicans were identical to or differing at only one to two dinucleotide positions from those of F. vesiculosus, indicating a recent common origin of the two species.     

Combined effects of temperature and salinity on growth rates of germlings of threeFucus species from iceland,helgoland and the north adriatic sea

The growth rate of germlings of threeFucus species (F. distichus subsp.edentatus from Iceland,F. vesiculosus from Helgoland, North Sea, andF. virsoides from the North Adriatic Sea) has been investigated under different temperature and salinity conditions. The highest growth rate and the maximum elongation factor were found at 9° C forF. distichus; in the other two species growth increased with increasing temperature. Growth decreases with dilution in the three species under consideration. A rather low tolerance to dilution was observed inF. vesiculosus germlings. In view of the overall distribution of the latter species the opposite was expected. It seems likely, however, that susceptible strains have developed at Helgoland, where salinity rarely drops below 30.     

Interaction between isopod grazing and wave action: a structuring force in macroalgal communities in the southern Baltic Sea

The macroalgal belt in the southern Baltic Sea may be partly structured by the interaction of physical and biological factors. A field study, spanning the 1990s, describes a rapid decline of the Fucus spp. stands along the wave-exposed Swedish southeast coast. During this period, a relative dominance of Fucus vesiculosus L. shifted to a relative dominance of Fucus serratus L. The decline of F. vesiculosus coincided with observations of large numbers of the grazing isopods Idotea baltica (Pallas) and Idotea granulosa Rathke, or with field observations of frequent grazing marks on Fucus fronds. I. baltica, but not I. granulosa, tended to aggregate in the declining Fucus spp. stands, indicating a strong preference for Fucus spp. In a mesocosm experiment I. baltica, when given a choice, grazed both Fucus species at weak water motion. At strong water motion grazing was concentrated on F. vesiculosus. It is hypothesized that one of the reasons I. baltica preferred F. vesiculosus to F. serratus at strong water motion may have been differences in habitat quality, like width of thallus, influencing the ability to cling to the plant. Smaller thallus, as in F. vesiculosus, thus is the preferred habitat for grazing of I. blatica. We postulate that the existence of F. serratus in the area may be favoured by strong wave action and moderate but not strong grazing by I. baltica, relaxing the interspecific competition from F. vesiculosus.