LONG‐TERM CHANGES IN THE STRUCTURE OF INTERTIDAL ASSEMBLAGES AFTER INVASION BY SARGASSUM MUTICUM (PHAEOPHYTA)1

The macroalgal assemblages at the low intertidal zone were studied at three localities on the north coast of Spain between 1977 and 2002. Two of these localities were invaded at the end of the 1980s by the brown seaweed Sargassum muticum (Yendo) Fensholt (Phaeophyta, Sargassaceae), whereas the third locality remained free of the invader. In 2002, distinct algal assemblages were noticed in invaded and noninvaded localities. No major changes were detected in the noninvaded locality. Apart from the obvious presence of S. muticum, the changes observed in the invaded localities included a significant reduction in abundance of the previous dominant species (the red alga Gelidium spinosum (S. G. Gmelin) P. C. Silva) as well as an increased number of species and diversity, increased primary productivity, and variations in the seasonal abundance patterns of some species. We speculate that the arrival of S. muticum had a negative effect on the dominant native G. spinosum, probably related to competition for light. This resulted in indirect positive effects on other species of the assemblage (such as Bifurcaria bifurcata R. Ross). Other small epiphytic opportunistic species might also have been benefited from the presence of S. muticum, because the invader has a rich associated epiphytic assemblage.     

Faunal differences between the invasive brown macroalga Sargassum muticum and competing native macroalgae

Interactions between macroalgae and their associated fauna are of great interest for marine invasions, because fauna may increase the biotic resistance of a system and macroalgal invasions may cause shifts in faunal composition. We tested for differences in faunal community structure between a macroalgal invader, Sargassum muticum, and several native macroalgae in intertidal pools on both the west and south coast of Portugal. On each coast, we compared the faunal diversity and composition associated with the invader with that of the competing native macroalga(e). On the west coast, the diversity of the fauna associated with S. muticum was equal to or lower than with the native competitor, Cystoseira humilis. Fauna composition differed between S. muticum and C. humilis at both locations, but within each species, no differences between locations were detected. In contrast, the fauna diversity on S. muticum of the south coast varied among locations. S. muticum fauna differed from the fauna of all native macroalgae at one location, but only from three out of seven native macroalgae at the other location. Discriminating fauna species did not show a consistent pattern towards higher or lower abundances in S. muticum compared to most native macroalgae, and species-specific contributions were small. Differences in fauna community also depended on the identity of the native macroalga. In conclusion, the fauna associated with S. muticum differs from many native brown macroalgae, but these differences were not consistent as they depend both on the native macroalgal species and on location. This invader does not seem to have a severe negative impact on local macroalgae-associated fauna.     

Response of macroalgal assemblages from rockpools to climate change: effects of persistent increase in temperature and CO2

Anthropogenically induced global climate change has important implications for marine ecosystems with unprecedented ecological and economic consequences. Climate change will include the simultaneous increase of temperature and CO₂ concentration in oceans. However, experimental manipulations of these factors at the community scale are rare. In this study, we used an experimental approach in mesocosms to analyse the combined effects of elevated CO₂ and temperature on macroalgal assemblages from intertidal rock pools. Our model systems were synthetic assemblages of varying diversity and understory component and canopy species identity. We used assemblages invaded by the non‐indigenous canopy forming alga Sargassum muticum and assemblages with the native canopy species Cystoseira tamariscifolia. We examined the effects of both climate change factors on several ecosystem functioning variables (i.e. photosynthetic efficiency, productivity, respiration and biomass) and how these effects could be shaped by the diversity and species identity of assemblages. CO₂ alone or in combination with temperature affected the performance of macroalgae at both individual and assemblage level. In particular, high CO₂ and high temperature (20°C) drastically reduced the biomass of macroalgal assemblages and affected their productivity and respiration rates. The identity of canopy species also played an important role in shaping assemblage responses, whereas species richness did not seem to affect such responses. Species belonging to the same functional effect group responded differently to the same environmental conditions. Data suggested that assemblages invaded with S. muticum might be more resistant in a future scenario of climate change. Thus, in a future scenario of increasing temperature and CO₂ concentration, macroalgal assemblages invaded with canopy‐forming species sharing response traits similar to those of S. muticum could be favoured.     

Effects of macroalgal identity on epifaunal assemblages: native species versus the invasive species Sargassum muticum

Seaweeds are a refuge from stressful conditions associated with life on rocky intertidal shores, and there is evidence that different macrophytes support different assemblages of mobile epifauna. Introduction of non-indigenous macroalgae may have a great impact on associated epifaunal assemblages and ecosystem processes in coastal areas. Previous studies have reported conflicting evidences for the ability of epifauna to colonize non-indigenous species. Here, we analyzed epifaunal assemblages associated with three species of macroalgae that are very abundant on intertidal shores along the Galician coast: the two native species Bifurcaria bifurcata and Saccorhiza polyschides and the invasive species Sargassum muticum. We collected samples of each species from three different sites at three different times to test whether variability of epifaunal assemblages was consistent over space and time. Epifaunal assemblages differed between the three macroalgae. Results suggested that stability and morphology of habitat played an important role in shaping the structure of epifaunal assemblages. This study also showed that the invasive S. muticum offered a suitable habitat for many invertebrates.     

Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a synthetic macroalgal assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of S. muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of S. muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming, producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.     

Ecosystem functioning impacts of the invasive seaweed Sargassum muticum (Fucales,Phaeophyceae)

Ongoing changes in natural diversity due to anthropogenic activities can alter ecosystem functioning. Particular attention has been given to research on biodiversity loss and how those changes can affect the functioning of ecosystems, and, by extension, human welfare. Few studies, however, have addressed how increased diversity due to establishment of nonindigenous species (NIS) may affect ecosystem function in the recipient communities. Marine algae have a highly important role in sustaining nearshore marine ecosystems and are considered a significant component of marine bioinvasions. Here, we examined the patterns of respiration and light‐use efficiency across macroalgal assemblages with different levels of species richness and evenness. Additionally, we compared our results between native and invaded macroalgal assemblages, using the invasive brown macroalga Sargassum muticum (Yendo) Fensholt as a model species. Results showed that the presence of the invader increased the rates of respiration and production, most likely as a result of the high biomass of the invader. This effect disappeared when S. muticum lost most of its biomass after senescence. Moreover, predictability–diversity relationships of macroalgal assemblages varied between native and invaded assemblages. Hence, the introduction of high‐impact invasive species may trigger major changes in ecosystem functioning. The impact of S. muticum may be related to its greater biomass in the invaded assemblages, although species interactions and seasonality influenced the magnitude of the impact.     

Spatio-temporal distribution patterns of the invasive macroalga Sargassum muticum within a Danish Sargassum-bed

Sargassum muticum was first observed in Scandinavia in Limfjorden (Denmark) in 1984, where it is now the most abundant and conspicuous macroalga. Despite the ecological importance of Sargassum, few studies have described seasonal patterns within Scandinavian Sargassum beds. We quantified the dynamics of macroalgae among years and seasons along a depth transect through a typical Sargassum bed in Limfjorden. The annual investigations (summer transects 1989–1999) showed a gradual increase in the dominance of Sargassum, especially at the 2–4-m depth interval. Significant seasonal dynamics in macroalgal abundance and assemblage structure were observed in this depth interval; the mean cover of Sargassum varied from ca. 5% (autumn and winter) to 25% (mid-summer). In comparison, encrusting algae had high and relatively stable covers throughout the year (ca. 20%). Other perennial macroalgae had low mean covers (<2%) characterized by a few patches of higher abundances. Except from a spring bloom, filamentous algae had low covers throughout the year. Within this relatively uniform bed, Sargassum abundance was positively related to boulders >10 cm in diameter and species richness was negatively correlated to depth and stones <10 cm in diameter, and non-correlated to other algal form-groups or grazer densities. Thus, in Limfjorden, the distribution of Sargassum is determined by large- (>6 m) and small-scale (<1 m) depth differences where low light limits Sargassum at depth, physical disturbance and sediment stress limits Sargasum in shallow waters, and the presence of stable boulder substratum facilitate Sargassum. Competition for space from other macroalgae and herbivory are probably of minor importance.     

Epibiota communities of the introduced and indigenous macroalgal relatives<Emphasis Type="Italic"> Sargassum muticum</Emphasis> and<Emphasis Type="Italic"> Halidrys siliquosa</Emphasis> in Limfjorden (Denmark)

Sargassum muticum (Phaeophyceae, Fucales) has recently been introduced to Limfjorden (Denmark) where its closest relative is the indigenous Halidrys siliquosa. Previous studies have demonstrated large quantitative (canopy biomass) and qualitative (canopy persistence) differences in the habitat available to epibiota within the canopies of these two macroalgae. We therefore hypothesised that these algae would support different epibiota communities and tested this by sampling the epibiota of S. muticum and H. siliquosa on seven occasions throughout 1997 by enclosing entire thalli in mesh bags. We found 53 epibiota taxa and, with only one exception, they were all recorded on both host species. Species richness and abundance of epibiota exhibited clear seasonal variation on both host species, although epibiota biomass was seasonally constant on H. siliquosa but not on S. muticum. These patterns were consistent with the different life histories of the host species. There was a weakly negative correlation between thallus size and epibiota biomass for both host species. When taking species-specific seasonal variation in thallus size into consideration, S. muticum and H. siliquosa were found to support significantly different epibiota biomasses. Multivariate analyses showed that epibiota community structure was different, although highly overlapping, between the two species, whereas there was an almost parallel temporal development in epibiota community structure. We conclude that it is unlikely that the introduction of S. muticum to Limfjorden has caused major changes in local epibiota community structure. However, the standing stock of epibiota is likely to have increased.Communicated by H.-D. Franke     

Algal communities at Gouqi Island in the Zhoushan archipelago, China

The kelp bed of Gouqi Island lies in the east of the East China Sea, in the Shengsi Archipelago. It is a key component of the island-reef ecosystem. This study evaluated kelp beds on intertidal and subtidal zones from 2004 to 2006. We evaluated seasonal variations in the community structure of macroalgae, and their relationship with water temperature, nitrogen and phytoplankton. The Gouqi Island kelp bed consists mainly of species of Sargassum. Sargassum horneri was the dominant species in subtidal zones, representing 90% of the total biomass. The dominant species in intertidal zones were S. fusiforme, S. horneri, S. thunbergii and Undaria pinnatifida. Fifty-four phytoplankton species were found in the Gouqi Island kelp bed, of which 51 species were found inside the kelp beds, 43 species outside the kelp beds, and 40 species were present both inside and outside the kelp beds. Diatoms were dominant both inside and outside the kelp beds. Except in winter, Skeletonema costatum was dominant, and its abundance in autumn was over 98% of the total abundance. Water temperature was found to be the primary factor influencing the growth of S. horneri in the Gouqi Island kelp bed. S. horneri grew slowly from November to March, and rapidly from March to June. The upper temperature limit for growth of S. horneri was about 18°C. Higher water temperatures can result in a decline in the kelp bed. Nitrogen seemed to be a limiting factor for macroalgal growth. This was particularly true for S. horneri and phytoplankton. Since nitrate is the primary nutrient for S. horneri, its absorption by S. horneri resulted in seasonal changes of nitrate in the ecosystem.     

The importance of shore height and host identity for amphipod assemblages

The spatial distribution of organisms associated with marine intertidal macroalgae may be a direct result of their tolerance to air exposure or an indirect consequence of the distribution of their host. We compared amphipod assemblages from five intertidal macroalgae to investigate their relationship with algal identity. To test the effect of height regardless of algal characteristics, we transplanted coralline algal turfs to three different levels within the intertidal zone and compared amphipod assemblages after 1 and 14 days. Interstitial volume was positively correlated to the abundance of amphipods, suggesting that this attribute may correspond better to the potential space for their occupation when compared to algal biomass, thallus volume or the ratio between thallus and interstitial volume. Algal level determined the structure of the amphipod assemblages. Upper-level (Acanthophora spicifera and Caulerpa racemosa) and intermediate-level (coralline) algae host similar amphipod assemblages dominated by Apohyale media, but different from lower-level algae (Padina gymnospora and Sargassum cymosum), which were dominated by Hyale niger. Ten of the 15 amphipod species reported from natural communities were found in the transplanted plots. Distinct pools of amphipod species colonized coralline transplants at upper and lower levels after 1 day. However, regardless of the position on the shore, transplanted coralline turfs supported similar assemblages after 14 days, indicating that algal identity is also important for species assemblages. Our results suggest that both height on the shore and host identity combine to determine the vertical structure of amphipod assemblages in the rocky intertidal.     

Differential effects of native and invasive algal wrack on macrofaunal assemblages inhabiting exposed sandy beaches

Many sandy beaches worldwide receive large amounts of drift seaweed, known as wrack, from offshore algal beds and closer rocky intertidal shores. Despite the important influence of algal wrack on macrofaunal assemblages from different coastal systems, relatively little attention has been paid to the macrofaunal responses in sandy beaches to macrophyte wrack supplies. Algal wrack is a key resource, i.e. for food and/or refuge, for beach invertebrates while its availability can affect diversity and abundance of intertidal animals including shorebirds, but the role of certain types of wrack and its location on the shore has not been examined experimentally to date. In this paper, we use experimental manipulation of two species of brown seaweeds, i.e. artificial wrack patches made up of the native macroalgae Saccorhiza polyschides and the invasive species Sargassum muticum, to test hypotheses about influences on macrofaunal assemblages inhabiting the drift line and supratidal levels of exposed beaches. Results pointed out that different types of wrack deposits were not used uniformly by invertebrates. Nutritional value differed between the two species of wrack. In most cases, the carbohydrates, lipids and organic carbon content were greater in patches of S. muticum than in patches of S. polyschides. Data also provided evidences that nutritional content and microclimatic conditions of wrack deposits, i.e. temperature and humidity, might affect macrofaunal assemblages.     

Comparative Studies on the Allelopathic Effects of Ulva pertusa Kjellml, Corllina pilulifera Postl et Ruprl, and Sargassum thunbergii mertl O. Kuntze on Skeletonema costatum （Grev.）Cleve

In the present study, we evaluated the allelopathic effects of three macroalgae, namely Ulva pertusa Kjellml, Corallina pilufifera Postl et Ruprl, and Sargassum thunbergii Mertl O. Kuntze, on the growth of the microalga Skeletonema costaturn （Grev.） Creve using culture systems in which the algae coexisted. The effects of the macroalgal culture medium filtrate on S. costatum were also investigated. Moreover, isolated co-culture systems were built to confirm the existence of allelochemicals and preclude growth inhibition by direct contact. The coexistence assay data demonstrated that the growth of S. costaturn was strongly inhibited when fresh tissues, dry powder and aqueous extracts were used; the allelochemicals were lethal to S. costatum at relatively higher concentrations. The effects of the macroalgal culture medium filtrate on the microalga showed both species specificity and complexity. The inhibitory effect of fresh macroalgal tissue and culture medium filtrate on the microalga was due to the alleochemicals released by the macroalgae. The results of the present study show that the allelopathic effects of macroalgae on the microalga are complex. The present study could shed light onto the basis of the interaction between macro- and microalgae.     

HABITAT DIFFERENCES IN THE TIMING OF REPRODUCTION OF THE INVASIVE ALGA SARGASSUM MUTICUM (PHAEOPHYTA,SARGASSACEAE) OVER TIDAL AND LUNAR CYCLES1

Sargassum muticum (Yendo) Fensholt is an invasive species that is firmly established on intertidal and subtidal rocky shores of Europe and the Pacific coast of North America. Local success and spread of S. muticum is thought to rely on its reproductive potential that seems dependent on exogenous factors like tidal and lunar cycles. This study is the first to compare the reproductive patterns (periodicity of egg expulsion and embryo settlement) of this invader in two different habitats: the middle and low intertidal. The combination of monthly, daily, and tidal samples at triplicate sites within each habitat showed a semilunar periodicity of egg expulsion and embryo settlement coincident with increasing tidal amplitude just before full and new moons. In both habitats, duration of each egg expulsion event was ～1 week, and embryo settlement occurred during the first daily low tide and with the incoming high tide during spring tides. However, both expulsion and settlement started 1–2 d earlier, expulsion saturation was faster, and settlement was higher in the mid‐ compared to the low intertidal. Our results suggest that the exact timing of gamete expulsion and embryo release of S. muticum responds to local factors, including tidal cues, which result in differences between mid‐ and low‐intertidal habitats.     

Historical changes and inventory of macroalgae from Königshafen Bay in the northern Wadden Sea

Knowledge on the distribution, abundance and species richness of intertidal macroalgae occurring on sandy and muddy flats of the German Wadden Sea is still incomplete. We summarize published and unpublished information available on the presence of macroalgae on the tidal flats of Königshafen Bay (island of Sylt, North Sea), one of the more extensively studied areas of the Wadden Sea. A total of 46 green algal species, 36 brown algal species and 26 red algal species has been recorded within the last 120 years on soft and hard substrata of Königshafen Bay (disregarding species found unattached or drifting). Several of these species were only temporarily resident on the tidal flats. Today, at least 35 green, 15 brown and 12 red algal species occur within or close to Königshafen Bay. Significant long-term changes in species abundances have occurred in all three major groups of algae: Since the late 1970s, dense green algal mats dominated byEnteromorpha flexuosa, E. radiata andE. prolifera have occurred regularly on the intertidal flats, whereas a general decrease of brown and red algae has been documented. Two red algal species,Gracilaria verrucosa and its epiphyteCallithamnion corymbosum, were conspicuous members of the macroflora until the middle of this century. Although still present in the 1980s, they have now disappeared completely. On the other hand, the brown algaSargassum muticum has begun to colonize mussel beds. The causal background of long-term changes in the macroalgal flora of Königshafen Bay is discussed. Owing to substantial nomenclatural changes during the last 120 years, a revised species list with authors’ names and synonyms is included.     

Direct evaluation of macroalgal removal by herbivorous coral reef fishes

Few studies have examined the relative functional impacts of individual herbivorous fish species on coral reef ecosystem processes in the Indo-Pacific. This study assessed the potential grazing impact of individual species within an inshore herbivorous reef fish assemblage on the central Great Barrier Reef (GBR), by determining which fish species were able to remove particular macroalgal species. Transplanted multiple-choice algal assays and remote stationary underwater digital video cameras were used to quantify the impact of local herbivorous reef fish species on 12 species of macroalgae. Macroalgal removal by the fishes was rapid. Within 3 h of exposure to herbivorous reef fishes there was significant evidence of intense grazing. After 12 h of exposure, 10 of the 12 macroalgal species had decreased to less than 15% of their original mass. Chlorodesmis fastigiata (Chlorophyta) and Galaxaura sp. (Rhodophyta) showed significantly less susceptibility to herbivorous reef fish grazing than all other macroalgae, even after 24 h exposure. Six herbivorous and/or nominally herbivorous reef fish species were identified as the dominant grazers of macroalgae: Siganus doliatus, Siganus canaliculatus, Chlorurus microrhinos, Hipposcarus longiceps, Scarus rivulatus and Pomacanthus sexstriatus. The siganid S. doliatus fed heavily on Hypnea sp., while S. canaliculatus fed intensively on Sargassum sp. Variation in macroalgal susceptibility was not clearly correlated with morphological and/or chemical defenses that have been previously suggested as deterrents against herbivory. Nevertheless, the results stress the potential importance of individual herbivorous reef fish species in removing macroalgae from coral reefs.     

Usefulness of tissue nitrogen content and macroalgal community structure as indicators of water eutrophication

We tested the hypothesis that the community structure and biochemical composition of macroalgae reflect the degree of nutrient concentrations in the water column. Benthic community structure and tissue nitrogen (N) content of macroalgae on intertidal rocky shores at three sites were investigated in relation to sewage effluents on Mireuk Island, Tongyeong city, on the southern coast of Korea. Ulva australis clearly dominated at site 1, which was close to a sewage treatment plant, where higher dissolved inorganic N and dissolved inorganic phosphate concentrations were observed. U. australis-dominated communities also appeared at site 2 (intermediate levels of nutrient enrichment). The macroalgal assemblage at site 3 (unimpacted site) was significantly different from those at sites 1 and 2. Five species (U. australis, Sargassum fusiforme, Grateloupia elliptica, Gelidium amansii, and Sargassum horneri) were dominant at site 3, representing 87 % of the total coverage throughout the study period. Species richness (d), evenness (J'), and diversity index (H') were highest at site 3, intermediate at site 2, and lowest at site 1, showing a negative relationship with nutrient levels. These results indicate that macroalgal community structure can be used as a bioindicator in water quality assessment. The tissue N content of green and red algae was responsive to nutrient availability, while the tissue N content of brown algae was relatively unchanged among the sites. This suggests that tissue N content as a bioindicator for detecting the influence of sewage effluent should be considered to reflect the N storage capacity of macroalgae.     

Ultraviolet radiation and consumer effects on a field-grown intertidal macroalgal assemblage in Antarctica

Ultraviolet radiation (UVR) research on marine macroalgae has hithero focussed on physiological effects at the organism level, while little is known on the impact of UV radiation on macroalgal assemblages and even less on interactive effects with other community drivers, e.g. consumers. Field experiments on macrobenthos are scarce, particularly in the Antarctic region. Therefore, the effects of UVR and consumers (mainly limpets were excluded) on early successional stages of a hard bottom macroalgal community on King George Island, Antarctica, were studied. In a two‐factorial design experimental units [(1) ambient radiation, 280–700 nm; (2) ambient minus UVB, 320–700 nm and (3) ambient minus UVR, 400–700 nm vs. consumer–no consumer] were installed between November 2004 and March 2005 (n= 4 plus controls). Dry mass, species richness, diversity and composition of macroalgal assemblages developing on ceramic tiles were followed. Consumers significantly suppressed green algal recruits and total algal biomass but increased macroalgal richness and diversity. Both UVA and UVB radiation negatively affected macroalgal succession. UVR decreased the density of Monostroma hariotii germlings in the first 10 weeks of the experiment, whereas the density of red algal recruits was significantly depressed by UVR at the end of the study. After 106 days macroalgal diversity was significantly higher in UV depleted than in UV‐exposed assemblages. Furthermore, species richness was significantly lower in the UV treatments and species composition differed significantly between the UV‐depleted and the UV‐exposed treatment. Marine macroalgae are very important primary producers in coastal ecosystems, serving as food for herbivores and as habitat for many organisms. Both, UVR and consumers significantly shape macroalgal succession in the Antarctic intertidal. Consumers, particularly limpets can mediate negative effects of ambient UVR on richness and diversity till a certain level. UVB radiation in general and an increase of this short wavelength due to stratospheric ozone depletion in particular may have the potential to affect the zonation, composition and diversity of Antarctic intertidal seaweeds altering trophic interactions in this system.     

Functional diversity and climate change: effects on the invasibility of macroalgal assemblages

Climate-driven and biodiversity effects on the structure and functioning of ecosystems are increasingly studied as multiple stressors, which subsequently may influence species invasions. We used a mesocosm experiment to test how increases in temperature and CO₂ partial pressure (pCO₂) interact with functional diversity of resident macroalgal assemblages and affect the invasion success of the non-indigenous macroalga Sargassum muticum. Early settlement of S. muticum germlings was assessed in the laboratory under common environmental conditions across three monocultures and a polyculture of functional groups of native macroalgae, which had previously grown for 3 weeks under crossed treatments of temperature and pCO₂. Functional diversity was a key driver shaping early settlement of the invader, with significant identity and richness effects: higher settlement occurred in low-diversity and low-stature assemblages, even after accounting for treatment biomass. Overall, early survivorship of settled germlings responded to an interaction of temperature and pCO₂ treatments, with survivorship enhanced in one treatment (high pCO₂ at ambient Temperature) after 3 days, and reduced in another (ambient pCO₂ at high Temperature) after 10 days, although size was enhanced in this same treatment. After 6 months in the field, legacy effects of laboratory treatments remained, with S. muticum reaching higher cover in most assemblages previously subjected to ambient pCO₂, but ephemeral green algae appearing disproportionately after elevated-pCO₂ treatment. These results caution that invasion outcomes may change at multiple points in the life cycle under higher-CO₂, higher-temperature conditions, in addition to supporting a role for intact, functionally diverse assemblages in limiting invader colonization.     

Grazing effects of the periwinkle <Emphasis Type="Italic">Echinolittorina</Emphasis><Emphasis Type="Italic">peruviana</Emphasis> at a central Peruvian high rocky intertidal

Echinolittorina peruviana is the most common gastropod in the high intertidal zone of Peru, representing more than 80% of the individuals present at that zone. Experimental removal of snails was used to evaluate their effects on (a) abundance of epilithic biofilm, (b) barnacle recruitment, and (c) abundance of macroalgae under “normal” conditions of the El Niño Southern Oscillation (ENSO). Experiments were carried out from October 2005 to April 2007 at two intertidal levels of a semi-protected rocky shore of central Peru. Results demonstrated that E. peruviana is able to control biofilm abundance and barnacle recruitment at both heights investigated, with marked effects in the lower zone. Erect macroalgae (Ulva spp. and Gelidium spp.) were less affected by grazing; but negative effects were observed on macroalgal crusts. Season and physical stress seem to play a more important role in the abundance of macroalgae in the high intertidal. Our results are similar to those reported elsewhere for high shore littorinids and represent baseline data to understand how the role of intertidal consumers will vary under the cold (La Niña) and warm (El Niño) phases of ENSO on these shores.     

Ecological aspects of Sargassum muticum (Fucales,Phaeophyta) in Baja California,Mexico: reproductive phenology and epiphytes

The reproductive phenology and epiphytic macroalgae of Sargassum muticum were studied through an annual cycle (September 1987 to November 1988) at two sites on the northwestern coast of Baja California, Mexico, which were subjected to different degrees of wave exposure. Sargassum muticum is a brown alga of Japanese origin, now considered a permanent member of the marine flora of Baja California. A similar reproductive development was observed at both sites, with a maximum percentage of reproductive plants from May to July (spring–summer) and minimum from December to March (winter). Reproductive plants were found throughout the year. A total of 48 species of epiphytes were identified and seasonal variation in their diversity was observed. The greatest diversity was found at the more protected site.