DIFFERENTIAL LIFE HISTORY PHASE EXPRESSION IN TWO COEXISTING SPECIES OF SCYTOSIPHON (PHAEOPHYCEAE) IN NORTHERN CHILE1

The identity of two phaeophycean taxa that monopolized the middle‐lower rocky intertidal zone of a coastal area chronically exposed to copper mine wastes in northern Chile was unraveled. One of them was preliminarily identified as the gametophytic stage of Scytosiphon lomentaria (Lyngbye) Link. The other, a dark crust, resembled the alternate stage of some Scytosiphon species. Comparative analysis of morphology, life history, and DNA sequences strongly suggests that crusts corresponded to sporophytic S. tenellus Kogame and confirm that erect thalli belonged to S. lomentaria. A clear segregation of erect and crustose thalli was found using internal transcribed spacer region 1 and RUBISCO spacer sequences. Furthermore, whereas crusts always grouped with S. tenellus, erect thalli always grouped with S. lomentaria. Life history studies failed to connect the two entities. First, field‐collected S. tenellus produced progeny that either recycled the crust, which reproduced by unilocular zoidangia, or developed into erect thalli. The latter, unlike typical gametophytic S. lomentaria, developed patchy sori of plurilocular zoidangia. Second, S. lomentaria displayed a direct‐type life cycle, in which progeny from erect individuals only developed into erect thalli and produced only plurilocular zoidangia. This constitutes the first experimental study on Scytosiphon from the Pacific coast of South America and the first report of S. tenellus on this coast. It is also the first report of the crustose stage of Scytosiphon appearing as a perennial and dominant algal species in a temperate rocky intertidal system.     

Spatial and temporal variation of photosynthesis in intertidal Mazzaella laminarioides (Bory)Fredericq (Rhodophyta, Gigartinales)

The red alga Mazzaella laminarioides is an economically important species with an extended latitudinal distribution along the Chilean coast. Its populations form mid-intertidal stands, several meters wide, and therefore are differentially exposed to environmental variables that result in temporal and spatial variability in productivity. We evaluated the effect of latitude and intertidal height on productivity by in situ measurement of photosynthetic performance. Daily and seasonal variations of O₂-evolution rate and maximal quantum yield (F _v/F _m) were determined in plants from the upper and lower intertidal zone at two localities 1500,km apart. Results suggest that plant responses were mainly affected by irradiation, temperature and desiccation. At local level, upper intertidal plants showed a reduced photosynthetic rate and quantum efficiency as compared to those displayed by plants from the lower intertidal, indicating their higher level of excitation energy acclimation. Stronger acclimation differences between upper and lower intertidal plants were observed in spring and summer. Differences in photosynthetic parameters between reproductive phases were recorded in autumn and winter, regardless of the position of the individuals in the intertidal zone. The effects of tidal elevation on seasonal patterns of photosynthesis were also influenced by latitude. Seasonal variation in photosynthetic efficiency was observed in plants from the northern population at both intertidal elevations, but only at the upper intertidal level in the southern population. This study shows that production variability in M. laminarioides results from differences in the intensity of environmental factors observed seasonally at local (intertidal) and latitudinal scales.     

INFECTIOUS DISEASES OF MAZZAELLA LAMINARIOIDES (RHODOPHYTA): CHANGES IN INFECTION PREVALENCE AND DISEASE EXPRESSION ASSOCIATED WITH SEASON,LOCALITY, AND WITHIN-SITE LOCATION1

This study addresses the issues of infection prevalence and disease expression in two wild populations of the red algal host Mazzaella laminarioides and their variability associated with locality, season, and spatial location of the host in the intertidal zone. Our results demonstrated that Endophyton ramosum is the most frequent infective pathogen affecting M. laminarioides in Matanzas and Pucatrihue. This situation prevailed through the year and across the high-to-low intertidal gradient. Although there was a general trend for lower levels of infection in late winter and early spring, only in a few, cases was well-defined seasonality detected. Furthermore, clear seasonal patterns, as displayed by deformative disease in the high intertidal zone of Pucatrihue, were attenuated in the middle and lower intertidal zones. Differences in levels of infection in M. laminarioides between the high intertidal zones of Matanzas and Pucatrihue diminished toward the low intertidal zone. Thus, effects of seasonality and locality on infection prevalence may be influenced, at least in part, by the position of the hosts an the intertidal zone. Spatial distribution of the diseased individuals also varied along the beach. This pattern was consistent between the two sites and seemed related to wave exposure and the specific pathogen. Comparisons of the size distribution of noninfected fronds with their infected counterparts showed that infections by Endophyton ramosum and Pleurocapsa sp. more frequently affected medium-and large-sized fronds. This pattern was consistent temporally and similar in the two localities. Finally, a clear association between maturity and prevalence of infection was detected. This association resulted in most fronds of the noninfected segment of the host population being immature, whereas most mature fronds were infected. In conclusion, infectious diseases affecting the red alga Mazzaella laminarioides are a persistent phenomenon in wild populations of the host, although only a small segment of the infected populations displays the full expression of the disease. In spite of the suggested role of factors such as season, latitude, and spatial location of the host on disease prevalence and expression, additional studies are needed to understand fully the dynamics of infectious diseases in wild populations of algal hosts.     

Changes in Epiphytic Bacterial Communities of Intertidal Seaweeds Modulated by Host, Temporality, and Copper Enrichment

This study reports on the factors involved in regulating the composition and structure of bacterial communities epiphytic on intertidal macroalgae, exploring their temporal variability and the role of copper pollution. Culture-independent, molecular approaches were chosen for this purpose and three host species were used as models: the ephemeral Ulva spp. (Chlorophyceae) and Scytosiphon lomentaria (Phaeophyceae) and the long-living Lessonia nigrescens (Phaeophyceae). The algae were collected from two coastal areas in Northern Chile, where the main contrast was the concentration of copper in the seawater column resulting from copper-mine waste disposals. We found a clear and strong effect in the structure of the bacterial communities associated with the algal species serving as host. The structure of the bacterial communities also varied through time. The effect of copper on the structure of the epiphytic bacterial communities was significant in Ulva spp., but not on L. nigrescens. The use of 16S rRNA gene library analysis to compare bacterial communities in Ulva revealed that they were composed of five phyla and six classes, with approximately 35 bacterial species, dominated by members of Bacteroidetes (Cytophaga-Flavobacteria-Bacteroides) and α-Proteobacteria, in both non-polluted and polluted sites. Less common groups, such as the Verrucomicrobiae, were exclusively found in polluted sites. This work shows that the structure of bacterial communities epiphytic on macroalgae is hierarchically determined by algal species > temporal changes > copper levels.     

Differential responses of tetrasporophytes and gametophytes of Mazzaella laminarioides (Gigartinales,Rhodophyta) under solar UV radiation

The effects of solar UV radiation on mycosporine‐like amino acids (MAAs), growth, photosynthetic pigments (Chl a, phycobiliproteins), soluble proteins (SP), and C and N content of Mazzaella laminarioides tetrasporophytes and gametophytes were investigated. Apical segments of tetrasporophytes and gametophytes were exposed to solar radiation under three treatments (PAR [P], PAR+UVA [PA], and PAR+UVA+UVB [PAB]) during 18 d in spring 2009, Punta Arenas, Chile. Samples were taken after 2, 6, 12, and 18 d of solar radiation exposure. Most of the parameters assessed on M. laminarioides were significantly influenced by the radiation treatment, and both gametophytes and tetrasporophytes seemed to respond differently when exposed to high UV radiation. The two main effects promoted by UV radiation were: (i) higher synthesis of MAAs in gametophytes than tetrasporophytes at 2 d, and (ii) a decrease in phycoerythrin, phycocyanin, and SPs, but an increase in MAA content in tetrasporophytes at 6 and 12 d of culture. Despite some changes that were observed in biochemical parameters in both tetrasporophytes and gametophytes of M. laminarioides when exposed to UVB radiation, these changes did not promote deleterious effects that might interfere with the growth in the long term (18 d). The tolerance and resistance of M. laminarioides to higher UV irradiance were expected, as this intertidal species is exposed to variation in solar radiation, especially during low tide.     

Patterns of organizations of intertidal and shallow subtidal vegetation in wave exposed habitats of central Chile

Wave-exposed rocky intertidal habitats of central Chile exhibit zonation of algal morphologies rather than strict patterns of species zonation. In low shore areas, there is a vertical sequence of perennial belts of calcareous crusts, kelp-like forms and expanded cushions or non-calcareous crusts. The calcareous crusts are represented by species of Mesophyllum, the kelp-like forms include Lessonia nigrescens and Durvillaea antarctica, while the cushions are represented by Gelidium chilense and G. lingulatum and the noncalcareous, expanded crusts by Codium dimorphum. Thin and thick blades, represented by Iridaea laminarioides, Ulva rigida and Porphyra columbia and filamentous forms including Ceramium rubrum, Centroceras clavulatum and Polysiphonia spp. are more patchy than the lower, perennial belts. They may, however, form distinct temporal monocultures at upper intertidal levels. Upper and lower limits of the various zones are set by interactions of several factors, the relative importance of which can change seasonally. When some of the factors restricting species distribution are experimentally removed, other interactions among factors become limiting.Within each zone, species are morphologically similar, with the abundance of species being regulated by symmetric competitive interactions. Competition is often asymmetric at the boundaries of zones except when adults of small-sized forms interact with morphologically similar juveniles of larger forms. Irrespective of their extremely different morphologies, the permanent, zone-forming algal species generally combine escape from grazers or defensive adaptations with clear competitive abilities. Nevertheless, there is a clear competitive hierarchy which is expressed in vertical displacements and zonation. The lowershore habitats could potentially be occupied by any of the different types of algae. Fast growth and large size allow the kelps to occupy this zone pushing the calcareous crust dominated-zone down into shallow subtidal areas and displacing the cushions and fleshy crusts into the low and middle intertidal regions. In turn, these last forms can displace thick and thin foliose forms and filaments to upper levels on the shore. Displaced forms may exist as patches at various levels of the shore.     

Effects of Chiton granosus (Frembly, 1827) and other molluscan grazers on algal succession in wave exposed mid-intertidal rocky shores of central Chile

Molluscan grazers can have important effects on the abundance, colonization rates, and successional pathways of algal assemblages and the entire intertidal community. In general, early successional algae are more readily consumed than corticated algae and kelps, which usually get established later in the community succession. To generalize, however, the effect of different grazers on algal assemblages must be examined on different coasts and under different scenarios. This information could help us understand the mechanisms of ecosystem processes and situations in which general models do not apply. Along the coast of Chile, humans harvest large keyhole limpets, which seem to be the only invertebrate grazers capable of controlling the dominant corticated alga Mazzaella laminarioides, a canopy-forming species that can cover extensive areas of the mid intertidal zone. In this scenario, where large limpets are harvested, the overall effects of the diverse molluscan assemblage of limpets, chitons and snails on algal succession and on corticated algae in particular are not clear. We conducted a 26-month-long experiment to evaluate the effects of molluscan grazers on mid-intertidal algal succession and to isolate the effects of Chiton granosus, the most conspicuous member of the assemblage at these tidal elevations. At sites heavily impacted by humans the molluscan grazer assemblage had strong negative effects on colonization and abundance of green algae such as ulvoids and Blidingia minima. In doing so, the grazer assemblage had a strong negative indirect effect on the establishments of chironomid fly larvae, which were only observed on green algal mats and rarely on bare rock. No significant effects were detected on epilithic microalgae, and effects on sessile invertebrates were highly variable over space and time. C. granosus also had significant negative effects on green algae but did not account for the total grazing pressure exerted by the guild. Limited foraging excursions (ca. 35 cm) from refuges and moderate site (crevice) fidelity in this species may contribute to the patchiness in green algal distribution observed in the field. Nearly 13 months after rock surface were experimentally cleared, M. laminarioides appeared in all experimental plots, but increased over three times faster in enclosures containing C. granosus than in exclosures plots or controls, suggesting that moderate levels of herbivory could actually facilitate the establishment of this alga in the succession and that the green algal cover found in the absence of grazers may delay its establishment.     

COALESCENCE VERSUS COMPETITION: FIELD AND LABORATORY STUDIES OF INTRA‐ AND INTERSPECIFIC ENCOUNTERS AMONG COALESCING SEAWEEDS

Classical ecological theory predicts that whenever growing individuals share a common and limiting resource, such as substratum in mid‐intertidal and shallow subtidal habitats, preemptive competition will occur determining species abundance and distribution patterns. However, conspecificity of several ecologically dominant Rhodophyta may coalesce when grown in laboratory cultures. The extent at which intraspecific coalescence occurs in the field and whether the process may also happens during interspecific encounters remain to be determined. If intra‐ and interspecific coalescence effectively occurs, then coexistence through coalescence rises as an alternative to competition among red‐algal dominated intertidal and shallow subtidal communities. Populations of Mazzaella laminarioides and Nothogenia fastigiata living in mid‐intertidal, semi‐exposed rocky habitats in Central Chile are being used to test the above ideas. Intra‐ and interspecific encounters occur in the field throughout the year. Coalescence does occur among conspecific partners but it has not been detected in interspecific encounters. Rather, a thick interface of compressed cells, necrotic tissues and cyanobacterial nodules is formed between the two contacting partners. In addition, observations of laboratory cultures indicate that spore germination, germling survival and differentiation of erect axes in bispecific cultures may be reduced when compared to single‐species controls. Interspecific differences in growth and differentiation rates appear as the mechanisms explaining a lack of coalescence and negative effects during interspecific contacts. On the other hand, the existence of conspecific coalescence in the field suggests this process should be considered as a real alternative to intraspecific competition among coalescing Rhodophyta.     

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.     

Taxonomic study of Scytosiphon (Phaeophyceae) from temperate coasts of Argentina

Scytosiphon is a common intertidal genus widely distributed on temperate coasts worldwide. Recently, eight species have been delimited with molecular tools. Although S. lomentaria is the only species that predominates in the macroalgal literature of the Southwestern Atlantic Ocean (SwAO), unpublished molecular data obtained for a population study of S. lomentaria revealed hidden species diversity of Scytosiphon among the individuals collected from four localities at the SwAO. The aim of this study was to revise the identity and phylogenetic relationships of Scytosiphon from temperate coasts of the SwAO using DNA data. Thalli were collected from the Argentinean coast between 39° S and 43° S, from which cox1 and rbcL gene sequences were obtained. Phylogenies and haplotype networks were inferred and morphology of gametophytes was studied. Four species were recognized, S. lomentaria, S. promiscuus, S. shibazakiorum, and one species that belongs to a complex of species known as “Scytosiphon Atlantic complex.” This complex was known to occur only in the North Atlantic, however, the results found in this study revealed that it has an extended distribution range that includes the southern hemisphere, where its populations have high genetic diversity and unique haplotypes. The morphological differences among the four species were subtle; denoting that previous Scytosiphon records from the SwAO attributed to the renowned S. lomentaria could represent different species. In addition, sex ratio and genome-wide single nucleotide polymorphisms (SNPs) analyses were done for populations of S. promiscuus presumably introduced to the SwAO, and the results indicated that they included female-dominant parthenogenetic populations, which were probably introduced from Japan.     

Effect of Detachment on the Palatability of two Kelp Species

Many species of macroalgae survive after becoming dislodged from their primary substratum, but little is known about their capacity to express anti-herbivore defences after detachment. We examined the effect of detachment on the relative palatability of the two kelp species Lessonia nigrescens and Macrocystis integrifolia to mesograzers. Laboratory and field experiments were conducted on the northern-central coast of Chile to investigate whether (i) time after detachment and (ii) grazing on detached and attached algae could trigger internal defence mechanisms in the algae, which may have acted as deterrents to grazing. In order to examine palatability, feeding assays were run after each experiment using fresh algal pieces and artificial food. Time after detachment had a significant influence on palatability of L. nigrescens but not of M. integrifolia. During the first 12 days of detachment, detached L. nigrescens held in grazer-free laboratory tanks were not significantly more palatable than attached conspecifics from the field but thereafter detached individuals became more palatable. Floating individuals of M. integrifolia showed no effect of detachment, indicating that this alga maintains its defence after detachment. An experiment conducted in the field confirmed these results for M. integrifolia. An additional laboratory experiment confirmed that attachment status plays an important role on algal defence reaction for L. nigrescens when exposed to grazers. Detached and previously grazed individuals of this species were less palatable than grazer-free control algae, but grazing had no effect on palatability of attached algae. Our results indicate that kelps have varying capacities for development of anti-grazing responses once they become detached, possibly depending on their capacity to float and survive after detachment.     

LAMINARIOCOLAX SP. (PHAEOPHYCEAE) ASSOCIATED WITH GALL DEVELOPMENTS IN LESSONIA NIGRESCENS (PHAEOPHYCEAE)1

As part of a long‐term study on the biology and ecology of the intertidal kelp Lessonia nigrescens Bory, we report on the occurrence of gall development on this alga, identified the possible causal agent, and assessed the extent of the phenomenon in two wild stands of the host. Our results showed that galls affecting natural populations of L. nigrescens were associated with the infection by a filamentous brown algal endophyte of the genus Laminariocolax. Assignment to Laminariocolax of the endophytes isolated from cultured gall tissue was based on the (i) high internal transcribed spacer 1 (ITS1) sequence similarity and phylogenetic relationship between the Chilean isolates and several species of the genus Laminariocolax endophytic in other kelps, (ii) reproductive and vegetative features of the endophyte in culture, and (iii) anatomical agreement of fully developed galls of Lessonia with those described for other kelp galls caused by endophytic members of Laminariocolax. Unequivocal identification at the species level of the endophytes infecting Lessonia, however, awaits further studies.     

Coexistence,size overlap and population regulation in tidal vs. non-tidal Palaemon prawns

Summary The effect of increased habitat heterogeneity in tidal areas on coexistence between Palaemon prawns was studied at eight sites along the European Atlantic coast. Two species which are sympatric in non-tidal areas, Palaemon adspersus Rathke and P. squilla (L.) are largely allopatric in tidal areas, and the sympatric size difference decreases in allopatry. In tidal areas the smaller species, P. squilla, is restricted to brown algal belts and rockpools. A third species, P. serratus (Pennant), larger than the others, occurs under oceanic salinities in subtidal brown algal belts and there has forced P. squilla to restrict its habitat distribution to adjacent intertidal rockpools. At estuarine salinities, however, P. squilla also inhabits the brown algal belts. A larger diel variation in stomach fullness index in P. squilla than in P. adspersus persists in non-tidal areas.Abiotic factors probably restrict the two largest species, viz. P. adspersus and P. serratus, to subtidal environments; these species are sensitive to the extremes in salinity, temperature or O₂ levels characteristic of the intertidal zone. Competition and/or predation probably relegates P. squilla to the intertidal zone. In non-tidal areas, where this zone is reduced, P. squilla increases its niche width and coexists with P. adspersus, and the size differentiation associated with sympatry may reduce interspecific competition.     

Dynamics of total surface bacteria and bacterial species counts during desiccation in the Malaysian sea lettuce, Ulva reticulata (Ulvales, Chlorophyta)

The present study illustrates the dynamics of surface bacteria during post‐harvest desiccation of Ulva reticulata Forsskal. Algal fronds were subjected to desiccation for 31 days. The total surface bacteria and bacterial species counts were monitored for moisture content and water activity index (a_w). There was an 86% decrease in total algal moisture content. However, a_w showed a more gradual decrease. The total bacterial count increased in the first week, reaching a maximum on day 7. After this, there was a drastic drop in the total bacterial count until day 14, and then a more gradual decline towards the end of the process. Six species of bacteria were isolated throughout this process: Azomonas sp., Aeromonas hydrophila, Vibrio alginolyti‐cus, Escherichia coli, Proteus vulgaris and Vibrio para‐haemolyticus. The dynamics of each of these bacterial species exhibited trends similar to the total bacterial count. Based on these findings, the drastic decrease in the total bacterial count after seventh day of desiccation could not be attributed to the a_w or salinity. Therefore, the possible exposure of these bacteria to the algal internal fluid upon the rupture of the thallus cells was seen as the most likely reason for the drop in the bacterial population. Scanning electron microscope micrographs taken after the tenth day of desiccation showed the presence of cracks and areas where the bacteria were exposed to the algal internal fluid. In vitro antibacterial tests of three different solvent extracts of Ulva reticulata were also carried out against these surface bacteria to verify the antibacterial potential of its internal fluid. It was apparent that all these surface bacteria were inhibited by at least one of the three extracts, and there were indications of the possible presence of multicompound antibacterial potential, since extracts of different polarity showed bacteria‐specific activity. Hence, it is possible that Ulva reticulata has the potential to protect itself against the opportunistic bacteria present on its surface and in its environment.     

Changes in the macrozoobenthos of the intertidal zone at Helgoland (German Bight, North Sea): a survey of 1984 repeated in 2002

Changes in the presence and absence of invertebrates as well as in species conspicuousness were documented in a rocky intertidal community based on surveys in 1984 and 2002. In 2002 six vertically and/or morphologically different stations of an intertidal platform were sampled. Five of these six habitats had already been surveyed in 1984. Replicating precisely the method of the first assessment, presence/absence changes as well as changes in species conspicuousness of 83 invertebrate species were documented, indicating that this intertidal community changed considerably during the 18-year interval. Compared with the study in 1984, 27 species newly appeared, whereas 32 species disappeared. Furthermore, 16 species increased in conspicuousness, whereas eight invertebrates decreased. The total number of species in 2002 was 154 versus 158 in 1984. Although algal species were not recorded as thoroughly as invertebrates, a massive decline in cover of Halidrys siliquosa was noted. Conversely, two invasive algal species became established after 1984, Sargassum muticum (since 1988), a cosmopolitan fucoid alga that prefers shallow subtidal areas for colonization, and Mastocarpus stellatus (introduction in the 1980s) that particularly colonized areas in the mid intertidal. In 1984 the mid intertidal zone was dominated by the brown alga Fucus serratus, whereas in 2002 the blue mussel Mytilus edulis and the periwinkle Littorina littorea were the most conspicuous organisms. Annual mean sea surface temperature (BAH measurements) warmed by 1.1°C over the past four decades. Range-related community shifts, introductions of non-indigenous species and the input of pollutants, are considered to explain long-term ecological changes in the invertebrate community at Helgoland.     

The non-native turf-forming alga Caulacanthus ustulatus displaces space-occupants but increases diversity

Non-indigenous seaweeds can be found in coastal habitats worldwide yet the ecological effects of only ~6 % of macroalgal introductions have been studied. The turf-forming red alga Caulacanthus ustulatus, a putative introduction from Asia, was discovered in southern California in 1999, yet has received very little attention despite being common in rocky intertidal habitats in the region. The purpose of this study was to evaluate the potential effects of Caulacanthus on native invertebrate and seaweed community composition. Macrofaunal, meiofaunal, and macroalgal community structure and diversity were compared between patches with (non-native) and without Caulacanthus (native) in the upper intertidal zone at 5 locations in southern California. Caulacanthus appears to displace macro invertebrates, such as barnacles, limpets, and periwinkles, while facilitating a more diverse array of meiofauna and macroalgae. This is likely due to the formation of a novel turf habitat in the upper zone where turfs are uncommon in this region naturally; algal turfs can increase habitat complexity, trap sediment, and maintain moisture during low tide which likely benefits meiofauna and seaweeds by providing food, habitat, or refuge from desiccation stress. Subsequent comparisons of invertebrate and seaweed assemblages were conducted in native and non-native patches at one site in the upper intertidal zone as well as in the middle intertidal zone where a native turf zone exists. Despite differences in community composition in the upper intertidal zone, no differences were observed in the middle zone, providing support that the novel turf created by Caulacanthus in the upper zone drives community differences.     

Experimental studies of herbivory and algal competition in a low intertidal habitat

Summary Ecological factors regulating the species composition and abundance of the plant assemblage in the low intertidal zone were studied in the semiprotected San Juan Islands in Washington state. In particular, the roles of an abundant herbivore (the chiton Katharina tunicata) and competition among plants were evaluated experimentally. Densities of Katharina were manipulated in large areas to establish 3 treatments: Katharina removals (N=2), controls (N=2), and Katharina addition (N=1). These sites were monitored for 4 years. Over this time, algal abundance and diversity increased in the Katharina-removal areas; algae in most functional groups proliferated, and a multi-storied intertidal kelp bed eventually developed. In the Katharina-addition, the abundance of all plants except crusts, diatoms, and surfgrass decreased, and overall diversity declined. Control sites underwent year-to-year fluctuation in the abundance of the most conspicuous alga, Hedophyllum sessile, but remained otherwise unchanged.At the end of the third year, the dominant space occupiers (the large brown algae) were removed from half of each of the Katharina removal and control areas. These kelps positively affect abundances of Katharina but are negatively correlated with limpet numbers. In the absence of competition from the kelps, the abundance and diversity of most other algal groups increased. This change was especially dramatic in the areas lacking both chitons and kelps, indicating that herbivory and competition have negative, additive effects on smaller algae. However, even in the absence of Katharina, abiotic disturbance (desiccation and storms) apparently mediates the competitive dominance of the kelps.     

Facilitative Effect of a Generalist Herbivore on the Recovery of a Perennial Alga: Consequences for Persistence at the Edge of Their Geographic Range

Understanding the impacts of consumers on the abundance, growth rate, recovery and persistence of their resources across their distributional range can shed light on the role of trophic interactions in determining species range shifts. Here, we examined if consumptive effects of the intertidal grazer Scurria viridula positively influences the abundance and recovery from disturbances of the alga Mazzaella laminarioides at the edge of its geographic distributions in northern-central Chilean rocky shores. Through field experiments conducted at a site in the region where M. laminarioides overlaps with the polar range edge of S. viridula, we estimated the effects of grazing on different life stages of M. laminarioides. We also used long-term abundance surveys conducted across ~700 km of the shore to evaluate co-occurrence patterns of the study species across their range overlap. We found that S. viridula had positive net effects on M. laminarioides by increasing its cover and re-growth from perennial basal crusts. Probability of occurrence of M. laminarioides increased significantly with increasing density of S. viridula across the range overlap. The negative effect of S. viridula on the percentage cover of opportunistic green algae—shown to compete for space with corticated algae—suggests that competitive release may be part of the mechanism driving the positive effect of the limpet on the abundance and recovery from disturbance of M. laminarioides. We suggest that grazer populations contribute to enhance the abundance of M. laminarioides, facilitating its recolonization and persistence at its distributional range edge. Our study highlights that indirect facilitation can determine the recovery and persistence of a resource at the limit of its distribution, and may well contribute to the ecological mechanisms governing species distributions and range shifts.     

Thermal tolerance,evaporative water loss,air-water oxygen consumption and zonation of intertidal prosobranchs: a new synthesis

Duration of emergence increases with tidal height on rocky shores therefore, emergence adaptations in intertidal species such as littorine and other prosobranch gastropods have been considered correlated with zonation patterns; temperature tolerance, desiccation resistance and aerial respiration rate all commonly assumed to increase progressively with increasing zonation level. Such direct correlations are rarely observed in nature. Maximal aerial gas exchange occurs in mid-shore, not high shore species. Temperature tolerance and desiccation resistance do not increase directly with shore height. Thus, hypotheses regarding physiological correlates of zonation require revaluation. A new hypothesis is presented that the high tide mark presents a single major physiological barrier on rocky shores. Above it, snails experience prolonged emergence and extensive desiccation; below it, predictable submergence and rehydration with each tidal cycle. Thus, desiccation stress is minimal below the high tide mark and maximal above it. Therefore, species restricted below high tide (the eulittoral zone) should display markedly different adaptive strategies to emergence than those above it (the eulittoral fringe). A review of the literature indicated that adaptations in eulittoral species are dominated by those allowing maintenance of activity and foraging in air including: evaporative cooling; low thermal tolerance; elevated aerial O₂ uptake rates; and high capacity for radiant heat absorption. Such adaptations exacerbate evaporative water loss. In contrast, species restricted to the eulittoral fringe display adaptive strategies that minimize desiccation and prolong survival of emergence including: foot withdrawal, preventing heat conduction from the substratum; aestivation in air; elevated thermal tolerance reducing necessity for evaporative cooling; position maintenance by cementation to the substratum and increased capacity for heat dissipation. In order to test of this hypothesis the upper thermal limits, tissue and substratum temperatures on emergence in direct sunlight and evaporative water loss and tissue temperatures on emergence in 40 °C were evaluated for specimens of six species of eulittoral and eulittoral fringe gastropods from a granite shore on Princess Royal Harbour near Albany, Western Australia. The results were consistant with adaptation to the proposed desiccation barrier at high tide. The eulittoral species, Austrocochlea constricta, Austrocochlea concamerata, Nerita atramentosa and Lepsiella vinosa, displayed adaptations dominated by maintenance of activity and foraging during emergence while the eulittoral fringe littorine species, Bembicium vittatum and Nodilittorina unifasciata displayed adaptations dominated by minization of activity and evaporative water loss during emergence. The evolution of adaptations allowing tolerance of prolonged desiccation have allowed littorine species to dominate high intertidal rocky shore gastropod faunas throughout the world's oceans.