Selective grazing of the isopod Idotea baltica between Fucus evanescens and F. vesiculosus from Kiel Fjord (western Baltic)

Grazing rates of the isopod Idotea baltica on Fucus evanescens and F. vesiculosus (Phaeophyta) were quantified in laboratory feeding preference experiments. Fucus species were offered alone (no-choice) or simultaneously (choice). In three of four no-choice experiments and in all four choice experiments, I. baltica significantly preferred F. vesiculosus to F. evanescens. F. evanescens recently immigrated into Kiel Fjord and has increased in abundance since 1990. One possible reason for the competitive success of this species may be that, compared to F. vesiculosus, it is less preferred by I. baltica, the most abundant mesograzer at the study site.     

Adaptation of light-harvesting pigments to downwelling light and the consequent photosynthetic performance of the eulittoral rockweeds Ascophyllum nodosum and Fucus vesiculosus

We compared the effect of habitat and water depth on the light-harvesting pigment content for Ascophyllum nodosum and Fucus vesiculosus at two near-shore stations in Long Island Sound (USA). Excised pieces of seaweeds were attached at depth intervals to a vertically buoyed line, and left in situ for 7 days. For comparison, fronds were collected from sun and shade habitats in the littoral zone. The three major antenna (light-harvesting) pigments increased in concentration with depth or shade. Chlorophyll c to a ratios remained stable at about 0.2. Fucoxanthin to chlorophyll a ratios decreased by 20 to 30% with depth or shade. Although pigment composition for the two rockweed species was equivalent, the maximum photosynthetic performance of F. vesiculosus exceeded that of A. nodosum by a factor of 2, while the compensation depths for 4 m-adapted A. nodosum and F. vesiculosus under natural limiting light conditions were equivalent. Plants held at 4 m had higher photosynthetic rates compared with plants held at 0 m, no matter the depth of measurement. Indirect evidence indicates that the enhanced photosynthesis of 4 m-adapted plants is due not only to higher concentrations of antenna pigments but to other physiological factors as well. We conclude that the clearly delineated vertical distribution of these two canopy species, the F. vesiculosus zone over the A. nodosum zone, is not determined by light quantity or quality, but by biotic factors as evidenced by the experiments of Menge which are cited herein.     

How an introduced seaweed can affect epibiota diversity in different coastal systems

Invasions by non-indigenous species have the potential to alter the biodiversity of recipient systems. The magnitude of this effect often depends on the nature of the invaded communities and the ecology of the invader. We investigated the impacts of the Japanese seaweed Sargassum muticum (Phaeophyceae, Fucales) on biodiversity in a rocky and sedimentary environment on two islands in the North Sea. In each case, we compared the epibiota of non-indigenous S. muticum with epibiota communities on taxonomically related and structurally similar native seaweed hosts. Total and average species richness on S. muticum were similar on the rocky shore (60 species and 22±6 species, respectively) and the sandy shore (64 species and 20±3 species, respectively). However, community structure and species composition differed significantly between the environments. On the rocky shore, another native fucoid seaweed, Halidrys siliquosa, supported an epibiota community very similar to that of the invader. On the sandy shore, the only other abundant native habitat-providing algal species was Fucus vesiculosus. This species supported a different and less diverse assemblage of associated taxa. We conclude that S. muticum enhances epibiota diversity in the sedimentary environment, probably by increasing the substratum availability and habitat heterogeneity. In contrast, it has negligible impacts on epibiota diversity in the rocky shore environment, where it does not represent a fundamentally new habitat component. We conclude that even within the same region, the consequences of non-indigenous species on biodiversity cannot be generalised but depend on the composition and structural complexity of the species in the recipient community. This paper is dedicated to Udo Schilling, Chief Diving Officer, Alfred Wegener Institute who trained all of the authors in the waters off Helgoland to be scientific divers. The work reported here and in previous publications has been made possible through Udo’s training and support. We remain grateful for his enthusiasm, effort and patience.     

Effect of diurnal variations in natural irradiance on the apical length growth and light saturation of growth in five species of benthic macroalgae

The diurnal variation in apical length growth rate of five benthic macroalgae from the Swedish West Coast (Fucus spiralis L., F. vesiculosus L., F. serratus L., Ascophyllum nodosum (L.) Le Jolis, and Chondrus crispus Stackhouse) was measured during 1.5 to 2.0-h intervals for 3 to 5 d in September 1984. All species showed a maximum in growth rate in the morning, followed by a continuous decline throughout the rest of the day (most pronounced in F. serratus, A. nodosum and C. crispus). By comparing the growth rates with the average irradiance during the same interval, saturation irradiances for length growth in natural daylight regime were estimated; 60 to 70 Wm^-2 for F. spiralis and F. vesiculosus, 90 to 100 Wm^-2 for A. nodosum, 30 to 40 Wm^-2 for F. serratus and 10 to 20 Wm^-2 for C. crispus. Average growth rates during the dark hours ranged from 33% of average day growth rates in A. nodosum to 63% in C. crispus, respectively.     

Feeding preferences of gammarid amphipods among four species ofFucus

Each of fourFucus species occurring on the Atlantic coast of Canada was presented to two species of gammarid amphipods in no-choice and multiple-choice laboratory experiments. We also measured concentrations of anti-herbivore phlorotannins in each of theFucus species. The study site was Cranberry Cove in Nova Scotia, Canada, and experiments were conducted in 1988. In a multiple-choice experiment,F. vesiculosus was very heavily grazed. Other species were much less affected. In a no-choice experiment,F. evanescens, F. spiralis andF. vesiculosus were eaten at similar rates, whileF. distichus was barely grazed. These results correlate with measurements of phorotannin concentrations in each of the species.F. distichus had much higher concentrations than did the other species (which were not significantly different among themselves). There is no general clear-cut relationship between grazer deterrence and competitive capability within the fucoid guild studied.     

Competition and facilitation between germlings of<Emphasis Type="Italic"> Ascophyllum nodosum</Emphasis> and<Emphasis Type="Italic"> Fucus vesiculosus</Emphasis>

Competitive interactions between germlings of Ascophyllum nodosum (L) Le Jolis and Fucus vesiculosus L. were studied both in the laboratory and on a shore of the Isle of Man, in the Irish Sea. Both intra- and interspecific competition were investigated by comparing the performance of algal germlings both in monocultures and mixed populations of the two species. The growth of germlings of both species reduced with increasing density. F. vesiculosus always grew faster than Ascophyllum and did best in mixed cultures, whereas Ascophyllum did least well when mixed with Fucus germlings. Clearly the adverse effects of F. vesiculosus on A. nodosum were greater than those of Ascophyllum cohorts. At the same total density, the survival and growth of Ascophyllum declined with an increasing proportion of Fucus germlings, implying that poor recruitment of A. nodosum results from strong competition with F. vesiculosus. However, under desiccation stress on the shore, F. vesiculosus enhanced the survival of A. nodosum at the early germling stage even though competition may occur again at the late stage. Thus, whether interactions between germlings take the form of competition or facilitation depends on the environmental conditions.Communicated by O. Kinne, Oldendorf/Luhe     

Flexible habitat selection and interactive habitat segregation in the marine congeners <Emphasis Type="Italic">Idotea baltica</Emphasis> and <Emphasis Type="Italic">Idotea emarginata</Emphasis> (Crustacea,Isopoda)

Habitat segregation among competing species is widespread yet very little is know how this is achieved in practice. In a case study, we examined short-term effects of conspecific and congeneric density on habitat selection in two competing marine isopod species, Idotea emarginata and Idotea baltica. Under semi-natural conditions in large outdoor cylindrical tanks (4 m high; volume 5.5 m³), animal groups of different size and composition had the choice between a set of relevant habitat samples (surface-floating seaweed, the water column, seaweed on the bottom). Habitat selection in both I. baltica and I. emarginata proved to be largely independent of conspecific density (level of intraspecific competition). In single-species treatments, both species showed a similar and stable pattern of distribution, with a clear preference for seaweed on the bottom. In mixed-species treatments (MST), however, the species were largely separated by habitat. While the distribution of I. emarginata was completely unaffected by the mere presence of interspecific competitors, habitat selection of I. baltica changed notably when I. emarginata was present. The habitat use patterns observed in MST conformed to those realized in geographical areas where the two species overlap in distribution: I. emarginata is dominant among decaying seaweed on the sea floor, and I. baltica is the dominant species among surface-floating seaweed. Our findings suggest that habitat segregation between the two species is essentially interactive, resulting from rapid decision-making of I. baltica with respect to habitat selection. The underlying mechanism is discussed. I. emarginata is highly superior to I. baltica in interference competition and rapidly eliminates the latter from one-habitat systems which do not allow I. baltica to escape from this interaction. In more natural, heterogeneous environments, however, I. baltica seems to be able to coexist with the superior competitor due to its broader habitat niche, flexibility in habitat selection, and a behavioural disposition to avoid normally preferred habitats when these are occupied by I. emarginata.     

Feeding preferences of periwinkles among four species ofFucus

A widely accepted view of intertidal community organizatiton in the NW Atlantic proposes that fucoid vegetation is maintained by the actions of predators which remove species competitively superior toFucus species. Herbivory is an important component of these predatory interactions, but has been studied largely with reference to the interaction betweenF. vesiculosus andLittorina littorea. There are many species of fucoids and herbivorous invertebrates on the shores of the NW Atlantic and this paper reports field and laboratory experiments performed in 1987 (in or near Halifax, Canada) on the effects of grazing by three species ofLittorina on adults and juveniles of four species ofFucus. In laboratory experiments, portions ofFucus species were presented singly (no choice) or together (multiple choice) to single species ofLittorina. AdultF. distichus was grazed to only a small extent.F. evanescens andF. vesiculosus adult tissues were heavily grazed in most laboratory experiments.F. spiralis adults were heavily grazed in no choice experiments, but were grazed only slightly in multiple choice tests. When adult fucoid tissues were exposed to a mixed suite of grazers in the field, onlyF. vesiculosus was grazed heavily. Tests done on adult tissues showed clearly thatFucus species are highly variable in their vunerability to grazers. Moreover, we found variability among periwinkle species in grazing rates on adultFucus. We also found a great disparity in the feeding rates of different periwinkle species on juveniles ofFucus. Therefore, it is not possible to extrapolate from a single periwinkle species/single fucoid species interaction when attempting to identify vegetation structuring processes.     

Macro- and mesoherbivores prefer native seaweeds over the invasive brown seaweed <Emphasis Type="Italic">Sargassum muticum</Emphasis>: a potential regulating role on invasions

Herbivory has a strong impact on algal distribution, abundance and community structure and may influence the establishment and spread of introduced seaweed species. In this study, we assess the potential regulating role of herbivory on one of the most invasive brown seaweeds: Sargassum muticum. Multiple choice feeding experiments were conducted with 13 native seaweeds, S. muticum and 5 herbivore species from the Northwest, Southwest and South of Portugal. S. muticum was always the least or among the least preferred seaweeds and attained one of the highest growth rates of the tested seaweeds, with and without herbivores. The addition of herbivores increased the number of cases by 40% in which the invader had higher growth rates. Our results suggest that low grazing pressure on S. muticum by the recipient herbivore community may give the invader a competitive advantage over at least part of the native seaweed community, thereby contributing to the invasiveness of S. muticum along the Portuguese coast.     

Grazer avoidance may explain the invasiveness of the red alga Gracilaria vermiculophylla in Scandinavian waters

The invasive red alga Gracilaria vermiculophylla has quickly spread across Europe, but it is unclear whether its success is based on a high tolerance to variations in environmental conditions or to the absence of native grazers that feed on this alga. We tested whether native invertebrate grazers prefer native algae to G. vermiculophylla. Feeding preferences of three common herbivores were quantified when offered G. vermiculophylla and native Fucus vesiculosus, Ceramium virgatum, and Ulva intestinalis in no-, two- and multiple-choice trials. Herbivore growth was measured when fed each of the algae separately. Grazers consumed G. vermiculophylla in no-choice trials, but avoided generally this alga when having a choice. U. intestinalis was always preferred over G. vermiculophylla, and grazers fed with U. intestinalis grew faster than those fed with G. vermiculophylla. We conclude that grazers avoid G. vermiculophylla to most native algae, which may benefit G. vermiculophylla in northern European estuaries.     

Rehydration of desiccated intertidal brown algae: Release of dissolved organic carbon and water uptake

The release of dissolved organic carbon (DOC) by Ascophyllum nodosum (L.) Le Jol. and Fucus vesiculusus L. during rehydration was investigated after desiccation under experimental conditions. During 30 min of rehydration of plants which had lost up to 70% of their water, A. nodosum released about 2 to 10 mg C/100 g dry weight, while F. vesiculosus released 10 to 50 times more. When dried to water losses exceeding 70%, plants of both species yielded similar amounts of DOC, approximating 2 g C/100 g dry weight. Considerable variance in the amounts of DOC released by individual plant of both species was observed over the whole range of water losses tested. Among the DOC released, the portion of carbohydrate-C was O to 5% with A. nodosum but 2 to 47% with F. vesiculosus. This difference was most pronounced when less than 100 mg C/100 g dry weight were released. A. nodosum regained a smaller percentage of its original weight than F. vesiculosus after 30 min of rehydration when more than 30% of fresh weight were lost during desiccation. Curves were obtained which enable the estimation of water losses in naturally desiccated specimens within 30 min.     

Artificial wounding decreases plant biomass and shoot strength of the brown seaweed Ascophyllum nodosum (Fucales, Phaeophyceae)

Brown seaweeds can serve as habitat and food for a number of mesoherbivore species, but the effect of mesoherbivore grazing on the fitness of individual seaweed plants is not well documented. Here we investigate how mechanical wounding, which mimics the grazing damage made by the herbivorous gastropod Littorina obtusata, affects the biomass change, apical growth, and individual shoot strength of the fucoid seaweed species Ascophyllum nodosum both in natural field populations and in the laboratory. We did not detect a significant effect of wounding on apical shoot growth in the laboratory, but wounding decreased the plant biomass in half of the studied A. nodosum populations in the field. Furthermore, we found that the strength of individual A. nodosum shoots was significantly lower when compared with unwounded control shoots initially after wounding. However, after 11 days incubation in the laboratory, the strength of the wounded shoots had increased almost to the level of the control shoots, although there was still a significant difference in the force required to break the shoots. These results indicate that wounding can increase the tissue loss (and thereby decrease the fitness) of A. nodosum plants in natural populations, probably through weakening of individual shoots. However, A. nodosum appears to have a mechanism to strengthen the shoots after mechanical wounding, and therefore, the timing of damage (e.g. prior to a storm) probably determines the amount of plant tissue lost through wave-induced forces.     

Egg release and settlement patterns of dioecious and hermaphroditic fucoid algae during the tidal cycle

The timing of gamete release by fucoid algae, although known to be restricted to calm days is not clearly understood within a circadian time scale. The need for externally fertilizing species to avoid gamete dilution suggests that in wave-exposed areas spawning may occur during particular tidal phases. However, this may differ between mating systems, as selfing species may be less affected by gamete dilution. In this study, two different approaches were used to determine when egg release occurs during the tidal cycle in two sister species with different mating systems. First, egg settlement of Fucus vesiculosus (dioecious) and Fucus spiralis (selfing hermaphrodite) was quantified on removable substrates (egg settlement disks) every day for 2 months and settlement patterns were used to statistically estimate the radius of a circle that would encompass 99% of each patch of settled eggs (the egg dispersal radius). Also, egg release was quantified every 2 h during the tidal cycle. A significantly larger egg dispersal radius (P < 0.02) was found for F. spiralis than F. vesiculosus, and this difference was somewhat site dependent with a greater difference between species in exposed sites. The egg dispersal radius was negatively correlated with significant wave height and positively correlated with sea surface temperature for both the species (P < 0.05), with a greater effect of both the factors for F. spiralis than for F. vesiculosus. Egg release during the tidal cycle was variable between species and experiments, with F. vesiculosus releasing more eggs, later in the day, and at a lower tide, than F. spiralis, which released fewer eggs, throughout the day and at all tides. The dioecious species, F. vesiculosus, may have developed a specific adaptation for timing the egg release to periods when emersed in exposed habitats to avoid rapid dilution of gametes that require outcrossing for fertilization. On the other hand, egg release for F. spiralis, which can self-fertilize, occurred both when emersed and immersed, suggesting this species has developed less synchrony with specific environmental factors. Site dependence also suggests local wave conditions can modulate timing of release. The data are consistent with the relaxation of the selective constraints of water motion on fertilization success in a selfing hermaphrodite, relative to an obligate outcrossing species. Results support the idea that species with different mating systems evolve different sensitivities to environmental cues for gamete release with specific implications for inbreeding and successful external fertilization in the ocean.     

Effects of detrital non-native and native macroalgae on the nitrogen and carbon cycling in intertidal sediments

The decay of non-native and native seaweed mixing may modify sediment biogeochemistry and organic matter transfers within benthic food webs according to their composition and biomass. The non-native species Sargassum muticum was deliberately added to the sediment of an intertidal sandflat at different biomass and mixed to the native species Ulva sp. and Fucus vesiculosus. The sediment porewater was then ¹³C and ¹⁵N enriched to test whether both detrital diversity and biomass influenced the transfer of porewater carbon and nitrogen to the sediment and to the macrofauna consumers. More ¹⁵N-nitrogen was mobilized to sediments and macrofauna when the 3-species detrital mixing was buried, probably because this mixing provided species-specific compounds such as polyphenols due to the presence of S. muticum and F. vesiculosus, as well as large amounts of nitrogen due to the presence of Ulva. Our study revealed the importance of detrital diversity and non-native seaweeds for the nitrogen cycling in the benthic food web.     

Functional diversity of mesograzers in an eelgrass–epiphyte system

Historically, small invertebrate grazers in marine plant communities have been considered to be a relatively homogeneous group in their impact on ecosystem processes. However, recent studies propose that species composition is an important agent in determining grazer effects. We used four mesocosm experiments to test the biomass-specific and density-dependent effects of common mesograzers in temperate regions (Littorina littorea, Rissoa membranacea, Idotea baltica and Gammarus oceanicus) on epiphyte and eelgrass biomass and productivity. Mesograzer species identity strongly influenced epiphyte accumulation and eelgrass growth, where Rissoa was the most efficient mesograzer (per biomass) and Gammarus had the weakest impact. Density-dependent effects varied considerably among species. Both gastropod species reduced epiphyte accumulation in direct proportion to their density, and Littorina had the strongest negative effect on epiphyte biomass. The impact of Idotea seemed to level off to a threshold value and Gammarus had no density-dependent effect on epiphyte accumulation at all. Rissoa and Idotea increased eelgrass productivity in accordance with their effect on epiphyte accumulation, whereas Littorina showed a less positive effect than could be expected by its strong impact on epiphyte biomass. Gammarus had no significant impact on eelgrass growth. Our results show that the different functional traits of superficially similar mesograzers can have important consequences for ecosystem processes in macrophyte systems.     

Functional divergence in heat shock response following rapid speciation of Fucus spp. in the Baltic Sea

In the Baltic Sea, the broadly distributed brown alga Fucus vesiculosus coexists in sympatry over part of its range (south west Gulf of Bothnia) with the Baltic endemic F. radicans sp. nov, while further north in colder and lower-salinity areas of the Baltic F. radicans occurs alone (north west Gulf of Bothnia). F. radicans appears to have arisen via rapid speciation from F. vesiculosus within the recent history of the Baltic (ca. 7500 BP). Possible functional divergence between the two species was investigated by comparing stress-responsive gene expression in a common-garden experiment. The experiment used two allopatric populations of Fucus vesiculosus from the Skagerrak (North Sea) and Central Baltic, as well as F. radicans from the same Central Baltic site. The two species in sympatry displayed divergent heat shock responses, while F. vesiculosus populations from allopatric sites did not. F. radicans was more sensitive to heat shock at 25°C, either alone or together with high irradiance and desiccation, than Baltic or Skagerrak F. vesiculosus. The results indicate that rapid functional divergence in the inducible heat shock response has occurred between sympatric species on a timescale of thousands of years.     

Zonation,vesicle pressure and gas composition in Fucus vesiculosus and Ascophyllum nodosum at Kristineberg (West Coast of Sweden)

Features of fucoid zonation (Fucus spiralis, F. vesiculosus, Ascophyllum and F. serratus) in a vertical range of 60 cm at Kristineberg (Gullmar Fjord, W. Coast of Sweden) are described. In contrast to identical species occupying areas with wider tidal ranges, these plants are smaller in size and inhabit narrow, overlapping zones. Investigation into the magnitude of pressure and gas composition in vesicles of F. vesiculosus and Ascophyllum nodosum revealed an average positive pressure equivalent to 1.7 and 7 cm of water for the 2 plants in succession. These small values compare favourably with the pressure head to which such vesicles are subjected at Kristineberg. O₂-percentage amounts to 28.5% and 32.7% in vesicles of both plants in succession. O₂-values increase in daytime and decrease at night or when plants are maintained in the dark. CO₂-content in the two plants fluctuates between 0 to 2.6% and bears no correlation to the oxygen values. It is suggested that the pressure produced in the vesicles is due mainly to oxygen. Comparison of these results with those by other authors are made whenever possible.     

Behavioural and metabolic adaptations of marine isopods to the rafting life style

Rafting on floating objects is a common dispersal mechanism for many marine invertebrates. In order to identify adaptations to the rafting life style, we compared behavioural and metabolic characteristics of two isopods, the obligate rafter Idotea metallica and the facultative rafter Idotea baltica. In laboratory experiments, I. metallica showed low locomotive activity and a tight association to the substratum. Idotea baltica, in contrast, was more active with more frequent excursions in the surrounding water column. Oxygen consumption rates were similar in both species. Idotea metallica fed on zooplankton making this species widely independent of autochthonous food resources of the raft. Feeding rates and digestive enzyme activities were low in I. metallica. Reduced egestion rates may indicate slow gut passage and, thus, efficient resorption of nutrients. Efficient food utilization and the ability to accumulate high amounts of storage products, i.e. lipids, indicate a possible adaptation of I. metallica to low food availability or starvation. The feeding behaviour of I. baltica, in contrast, was more herbivorous and appeared wasteful and inefficient. Low lipid contents in I. baltica also indicate poor storage reserves. Thus, I. baltica requires a permanent access to food.     

Seasonal fluctuations of benthic macrofauna in the apalachicola estuary,Florida, USA: The role of predation

Long-term monitoring studies revealed a negative correlation between the abundance of bottom-feeding fishes (Leiostomus xanthurus and Micropogonias undulatus) and benthic macrofaunal density, suggesting a causal relationship. To test this hypothesis, large (3x3 m) topless predator-exclusion cages (mesh opening=6 mm) of a design that produced minimal physical artifacts were constructed at two shallow mud-bottom oligohaline stations. Two-sided control cages and uncaged control areas were also sampled Experiments coincided with maximum abundances of bottom-feeding fishes in the spring and motile macroinvertebrates (penaeid shrimp) in the fall. During short-term (two-month) exclusion experiments in the spring and fall of 1980, large predators (those excluded by 6-mm mesh) were not a major factor in regulating the densities of most macrofaunal species. Predation may be important during other seasons, although this possibility was not tested. Densities of Leiostomus xanthurus and Penaeus setiferus during 1980 were lower than in previous years. Despite the reduced abundance of bottom-feeding fishes, macrofaunal densities still declined during late winter/early spring. It is postulated that increased river flow and lowered salinities during the late winter/early spring may contribute to the decline of macrofaunal densities by reducing larval recruitment.     

Robustness in life history of the brown seaweed Ascophyllum nodosum (Fucales, Phaeophyceae) across large scales: effects of spatially and temporally induced variability on population growth

Understanding the demography and function of biotope-forming seaweed species is of great importance for the conservation of the target species itself, as well as its associated organisms. The brown seaweed Ascophyllum nodosum is fundamental for the functioning of coastal marine ecosystems in the North Atlantic. In this study, we use a data-based size-classified matrix model to investigate the temporal and spatial variability in demography, and the environment-specific stochastic sensitivity and elasticity, of two A. nodosum populations, one in western Sweden and one on the Isle of Man in the Irish Sea. A significant difference between the two populations was that the Swedish population had comparably low and more variable stochastic population growth rate (λ _s). This pattern was partly explained by the relatively high and varying mortality rates during extreme ice-years in Sweden, and by the lower survival of small individuals during all years. There were also fewer large individuals in Sweden due to lower transitions to the larger size-classes and higher probability of shrinkage. Sensitivities were analogous in the two populations, and showed a high selection pressure for increased individual growth. Elasticities were also similar, with the exception that survival of the smallest individuals (i.e., transition a _1,1), had a higher elasticity on the Isle of Man. Overall, the stochastic growth rate (λ _s) was most sensitive to proportional changes in loop- (i.e., survival within size-class) and, to some extent, growth-transitions in both study areas. These results show that structurally and demographically diverging A. nodosum populations may be similarly sensitive to changes in vital rates. This, in turn, indicates a plastic life history of A. nodosum that may cope with large environmental variability. The results further suggest that environmental change affecting the survival or growth of the larger, reproductive A. nodosum individuals could have severe and regional effects on the abundance and biomass of this species, with potential negative effects on the biodiversity of the associated communities.