Ecology and dynamics of two intertidal algal communities on the littoral of the island of São Miguel (Azores)

The intertidal benthic algal communities of two sites located on opposite coasts of São Miguel Island (Azores), were studied over a 2-year period (September 1993–September 1995). At both sites (São Roque on the south coast and São Vicente on the north), the littoral region was surveyed from the upper intertidal down to the sublittoral fringe. The survey revealed five distinct zones, with a variable degree of overlapping. The two upper zones were characterized by animals (littorinids and barnacles, respectively). Lower down, algal communities formed three distinct zones: an upper Fucus spiralis/Gelidium microdon association, a more extensive turf zone, and a belt featuring erect or frondose algae. Upper in the eulittoral, the turf was mainly monospecific, and dominated by Caulacanthus ustulatus. The lower eulittoral turf was dominated by articulated coralline algae, the associated species differing between the two sites studied. Mainly erect algae (Pterocladiella, Asparagopsis, etc.) occurred furthest down the shore and extended into the nearby shallow sublittoral. Two intertidal communities were studied at each locality: the upper eulittoral (Caulacanthus turf in São Roque and the Fucus spiralis/Gelidium microdon association at São Vicente), and the lower eulittoral (the coralline turf). The lower littoral communities had a higher algal diversity. A general pattern was observed in the seasonal variation of biomass: the lower levels exhibited the higher values in late summer/early autumn, the period in which the upper levels had the lowest standing crops. Physical factors are proposed to account for this. No significant inter-annual variations could be detected, indicating relatively stable communities, at least on a short-term basis.     

A field investigation into the effects of interspecific competition on the distribution and cover of Ulothrix flacca

Ulothrix flacca is a filamentous green alga found in its greatest abundance in the littoral fringe of the rocky intertidal during the winter months. It is much less abundant below this level. This experiment was undertaken tot determine if the lower abundance below the littoral fringe is due to the effects of interspecific competition or physiological limitations. Clearing experiments were performed on the Coast of Maine, USA to eliminate competition. Belts 10 cm wide that ran from the littoral fringe downward into the eulittoral zone were cleared of all attached organisms down to bare rock. A second condition was set up in which only the brown algal canopy was removed from a 60 cm wide belt. Unaltered belts were designated as controls. Squares 0.25 m² in area were cleared in the littoral fringe and eulittoral zone. U. flacca was able to colonize the cleared areas of the eulittoral zone and reach abundances equal to those found in the unaltered control areas of the littoral fringe. This indicates that physiological limitations do not restrict the abundance of U. flacca in the eulittoral zone. However, evidence of grazing in cleared areas adjacent to high concentrations of the barnacle Semibalanus balanoides suggest that competition alone is not responsible for the limited abundance of U. flacca in the eulittoral zone. Herbivores associated with the presence of barnacles may also have an impact on the abundance of U. flacca, even during the winter months.     

Peculiarities of the Structure of and Trends in the Macrobenthos Community of the Ermolinskaya Bay Lagoon Ecosystem,Kandalaksha Bay,White Sea

The species composition and spatial and trophic structures of the macrobenthos communities of the Ermolinskaya Bay were studied, and the main changes that had occurred since previous explorations (the 1960s) were analyzed. The most significant changes in the spatial structure of the macrobenthos community (species diversity, population density and biomass, and similarities of the species’ distribution) were observed with regard to the sea level zone: the most pronounced at the upper littoral, followed by the middle littoral, lower littoral, and sublittoral. The longitudinal gradient of the community structure (from the innermost area seaward) was pronounced less strongly. Altogether, four macrobenthos communities can be identified in Ermolinskaya Bay; they differ in species composition, diversity, population density and biomass, and trophic structure: (1) community of the upper littoral with the dominance of collecting deposit feeders, Hydrobia ulvae, and scrapers, Littorina saxatilis; (2) community of the middle littoral with the prevalence of deposit feeders, Hydrobia ulvae and Macoma balthica, seston feeders, Mya arenaria, and, to a lesser extent, Mytilus edulis; (3) community of the lower littoral with the domination of the filtering organisms, Mytilus edulis, to a lesser extent collecting deposit feeders, Hydrobia ulvae and Macoma balthica, and scrapers, Littorina littorea; and (4) community of the sublittoral, where the leading positions belong to the group of collecting deposit feeders, Macoma balthica, to a lesser extent also Hydrobia ulvae (in the innermost area) and Capitella capitata, Arenicola marina (in the marine part of the bay). The gradual siltation of the bay and the reduction of its connection to the sea have led to the development of a littoral complex of species in the sublittoral, whereas the species typical at the sublittoral in the 1960s are now mainly found at the outlet of the bay.     

Effect of the Razdol’naya River on structure of the free-living nematode community of the Amursky Bay,Sea of Japan

The effect of the Razdol’naya River on the littoral and sublittoral nematode community was studied in the innermost part of the Amursky Bay. Thanks to the seasonal abundance of juveniles of macrobenthic animals, pseudomacrobenthic species prevailed in the meiobenthos littoral community. Juvenile amphipods dominated (39.5%), the percentage of nematodes was 18.4% (45 species). The nematodes dominated in the meiobenthos community (65.7%, 71 species) at the sublittoral zone. The similarity of the species structure of littoral and sublittoral nematodes reached 36.5%. Low population density (2.6±1.7 thousand ind/m²) and poor species diversity of nematodes (7 species), the dominating form being Oncholaimium paraolium, were recorded at the littoral zone, which was subjected to direct riverine influence (1‰salinity). The correlation analysis revealed no relationship between salinity and population density of nematodes at the littoral zone. At the sublittoral zone (at stations of salinity less than 2.85‰) the parameters of population density (73.9±32.3 thousand ind/m²) and species diversity (12 species) of nematodes were the lowest, Parodontophora timmica dominated. A poor expressed dependence between the density of nematode populations and salinity was revealed. The correlation between the average size of ground particles and population density of nematodes was recorded. The greatest similarity of the species structure of littoral and sublittoral nematodes (26.67%) was recorded on gravel-sand bottoms.     

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.     

三亚岩相潮间带底栖海藻群落结构及其季节变化

为了解三亚潮间带大型海藻生态现状,布设6条岩相断面,于2008年1—11月进行了四季野外的调查,并使用物种多样性指数与聚类分析研究了潮间带大型海藻四季的群落结构和多样性变化。调查共鉴定出大型底栖藻类130种,隶属于4门75属,其中红藻门41属67种、褐藻门16属31种、绿藻门15属29种、蓝藻门3属3种。潮间带海藻种类数季节变化不大,并呈低潮带>中潮带>高潮带的垂直分布特征,且中低潮带有共同分布种,夏季尤为普遍。潮间带海藻夏冬两季平均生物量高于春秋两季,但并无显著差异。优势种季节变化明显:春季有波利团扇藻Padina boryana、日本仙菜Ceramium japonicum、半叶马尾藻Sargassum hemiphyllum、海柏Polyopes polyideoides,夏季有波状软凹藻Chondrophycus undulates、冠叶马尾藻Sargassum cristaefolium、宽扁叉节藻Amphiroa dilatata、石花菜Gelidium amansii,秋季有冠叶马尾藻、波状软凹藻、叶状铁钉菜Ishige foliacea,冬季有瓦氏马尾藻Sargassum vachellianum、苔状鸭毛藻Symphyocladia marchantioides、珊瑚藻Corallina officinalis、波利团扇藻、日本仙菜。群落多样性冬高夏低:Shannon多样性和Margalef丰富度指数最大值出现在冬季,Shannon多样性和Pielou均匀度指数最低值出现在夏季。区域底栖藻类以暖水性热带-亚热带种为主,其次为温水性的暖温带种类,基本符合南海南区暖水区系特点。研究表明,尽管三亚潮间带底栖海藻种类数与生物量上并无显著的季节差异,但群落结构与多样性均显示出一定的季节变化。     

Seaweed of the littoral zone at Cove Island in Long Island Sound: annual variation and impact of environmental factors

A site in the western part of Long Island Sound was monitored from January 2000 to May 2002. The littoral was divided into five different zones from the supra-littoral fringe (A) to the infra-littoral fringe (E). The midshore was dominated by Fucus vesiculosus L. and the sublittoral fringe by Chondrus crispus Stackh. There was a significant change in community structure over the years and the predominant change occurred between 2001 and 2002. The alternation in community structure was caused by an increase in abundance of species like Porphyra suborbiculata Kjellm., Porphyra leucosticta Type A and C (Neefus et al. 2000), Ceramium virgatum Roth, and Codium fragile subsp. tomentosoides (van Goor)Silva and a decrease in abundance in Fucus vesiculosus, Blidingia minima (Nägeli ex Kütz.) Kylin and Ulva lactuca L. The changes in community structure coincided with the change in environmental conditions. Air temperature as well as surface seawater temperature (depth <2 m) were the most important factors of those analyzed. Temperature seems to be the bottom-up force regulating the community structure.     

A comparison of the ecology and life cycles of two species of littoral isopod

A comparison is made of the vertical distribution, life cycles, and ecology of two littoral species of the sexually dimorphic isopod, Dynamene, on rocky shores of northwest France. Adult D. bidentata (Adams) prefer middle and lower eulittoral crevices and empty Balanus perforatus Brug. tests, and juveniles a wide variety of algae in the same zones. Males are capable of killing B. perforatus by wedging themselves tightly in the operculum so preventing feeding and respiration. Adult Dynamene magnitorata Holdich prefer empty Balanus crenatus Brug. tests, channels within sponges such as Halichondria and Hymeniacidon, and spaces within and between ascidian colonies, e.g., Dendrodoa and Didemnum, in the lower eulittoral and upper sublittoral. Juveniles are found in the same zones on a restricted number of algal species, in particular Chondrus crispus Stack. There is little overlap in the habitat preferences of the two species. Availability of food and tolerance of desiccation do not appear to be the primary factors responsible for the maintenance of zonation patterns; temperature would appear to be the main factor involved.The stages of the life cycle of the two species are the same but the sex-ratio in Dynamene magnitorata is much lower, and this is possibly due to the fact that the male may only spend one breeding season in the reproductive habitat as opposed to the two spent by male D. bidentata.Populations of D. bidentata from different latitudes show a tendency to increase the length of their reproductive and growth phases at more southerly locations so that ovigerous females and juveniles occur throughout the year. Winter air temperatures are a possible cause of these differences in timing.Stereoscan electron microscope observations are included to clarify the identification of the three species of Dynamene found on the Atlantic coasts of Europe and northwest Africa.     

Photosynthetic characteristics and UV stress tolerance of Antarctic seaweeds along the depth gradient

The photosynthetic characteristics through P-E curves and the effect of UV radiation on photosynthesis (measured as rapid adjustment of photochemistry, F _v/F _m) and DNA damage (as formation of CPDs) were studied in field specimens of green, red and brown algae collected from the eulittoral and sublittoral zone of Fildes Peninsula (King George Island, Antarctic). The content of phenolic compounds (phlorotannins) and the antioxidant activity were also studied in seven brown algae from 0 to 40 m depth. The results indicated that photosynthetic efficiency (α) was high and did not vary between different species and depths, while irradiances for saturation (E _k) averaged 55 μmol m^?2 s^?1 in subtidal and 120 μmol m^?2 s^?1 in eulittoral species. The studied species exhibited notable short-term UV tolerance along the vertical zonation. In intertidal and shallow water species, decreases in F _v/F _m by UV radiation were between 0 and 18 %, while in sublittoral algae, decreases in F _v/F _m varied between 3 and 35 % relative to PAR treatment. In all species, recovery was high averaging 84–100 %. The formation of CPDs increased (15–150 %) under UV exposure, with the highest DNA damage found in some subtidal species. Phlorotannin content varied between 29 mg g^?1 DW in Ascoseira mirabilis from 8 m depth and 156 mg g^?1 DW in Desmarestia menziesii from 17 m depth. In general, phlorotannin concentrations were constitutively high in deeper sublittoral brown algae, which were correlated with higher antioxidant activities of algal extracts and low decreases in photosynthesis. UV radiation caused a strong decrease in phlorotannin content in the deep-water Himantothallus grandifolius, whereas in D. menziesii and Desmarestia anceps, induction of the synthesis of phlorotannins by UV radiation was observed. The antioxidant activity was in general less affected by UV radiation.     

ICE NUCLEATION IN SEAWEEDS IN RELATION TO VERTICAL ZONATION1

The ice nucleation temperatures of thallus fragments of different macroalgal species were investigated. The samples were collected from the littoral zone on a rocky shore in Trondheimsfjord, central Norway. Thallus fragments of species growing in the upper parts of the eulittoral zone had lower ice nucleation temperatures than those of species living in the lower eulittoral zone and in the upper sublittoral zone. Samples collected in the winter had lower nucleation temperatures than samples from the same species collected in the summer, which indicates that the seaweeds are removing or inactivating ice nucleators as a part of their cold hardiness strategy.     

Algal invasions,blooms and biodiversity in lakes: Accounting for habitat-specific responses

Range expansions of species comprise a pervasive environmental problem worldwide and can cause substantial ecological and economic impact. However, the magnitude of impact may vary across habitats, highlighting the need to account for spatial heterogeneity in assessment studies. Here we compare invertebrate community structure in three habitats (littoral, sublittoral, and profundal) of boreal lakes that suffer recurring blooms of a regionally expanding, nuisance flagellate, Gonyostomum semen (Raphidophyta), with the assemblage structure in lakes were no blooms occur. We contrast community structure over a 6-year period using univariate metrics (total abundance, community evenness, species richness, and Simpson diversity) and multivariate community similarity to infer habitat-specific associations of local (alpha) diversity. We also calculated indices of multivariate dispersion to infer associations with beta diversity; i.e., whether or not habitats in bloom lakes show faunal homogenisation. Results show that the magnitude of assemblage alteration in bloom relative to bloom-free lakes varied with habitat and increased from the littoral to the profundal habitats. Littoral assemblages in bloom and bloom-free lakes shared similar alpha (taxon richness, evenness and Simpson diversity) and beta diversity characteristics, despite differing in multivariate community similarity. By contrast, alteration of assemblage structure was most severe in the profundal and manifested in reduced diversity and faunal homogenisation (i.e. decreased beta diversity) in bloom relative to bloom-free lakes. This was due to numerical dominance of the predatory phantom midge, Chaoborus flavicans, in the profundal of bloom lakes. Not only do the results highlight that spatial heterogeneity should be accounted for to assess the potential broader impact of nuisance species on biodiversity within lakes; more generally, the dominance of a single species suggests a reduced overall resilience of bloom lakes, making them more susceptible to environmental perturbation.     

Intertidal zone of Svalbard 3. Littoral of a subarctic, oceanic island: Bjornoya

Twenty-two stations in the intertidal and shallow sublittoral of Bjornoya (74 °N, 19 °E) were studied in August 1994 revealing a large and diverse standing crop of macro-algae (16 species) and littoral macrofauna (at least 17 species). In most places the biomass of littoral macroorganisms exceeded 100 g ww/m². In the shallow sublittoral, between 2 and 20 m, 45 animal taxa and 23 algae species were collected. Littoral coarse sand meiofauna was dominated by Turbellaria, while, on algae, Halacaridae and Harpacticoida predominated. Meiofauna densities ranged from 0 to 169 ind./10 cm² and biomass from 0 to 0.4 g dw/m². The abundance of littoral species and their zoogeographic origin resemble that of Spitsbergen more than that of the northern Scandinavian coast, although both are of equal distance from Bjornoya. The first record of the boreal bivalve Mytilus edulis from the island is presented. Another striking feature was the presence of the arctic amphipod Gammarus setosus and the absence of its boreal sibling species G.oceanicus. Received: 19 March 1996 / Accepted: 1 December 1996     

Resistance to fresh and salt water in intertidal mites (Acari: Oribatida): implications for ecology and hydrochorous dispersal

The resistance to fresh water and seawater in three intertidal oribatid mite species from Bermuda, Alismobates inexpectatus, Fortuynia atlantica and Carinozetes bermudensis, was tested in laboratory experiments. Larvae are more sensitive to fresh and salt water, nymphs and adults showed equal tolerances. Fortuynia atlantica and A. inexpectatus were more resistant to salt water whereas C. bermudensis survived longer in fresh water. Differences in the resistance to fresh and salt water among the three species may be related to their different vertical occurrences in the eulittoral zone but also to the ability of single species to dwell in periodically brackish waters. In all three species half of the specimens survived at least 10 days in fresh water and more than 18 days in salt water. Maximal submersion time in fresh and salt water ranged from 40 to 143 days. Based on median lethal times it could be estimated that each species would be able to survive transport in seawater along the Gulf Stream over a distance of 3,000 km, from Central America to Bermuda. Thus hydrochorous dispersal should be assumed as the most likely mode of dispersal in intertidal fortuyniid and selenoribatid mites.     

The distribution of Zostera and its exploitation by wildfowl in the solent,Southern England

In the Solent estuarine system on the central south coast of England, Zostera marina L. occurs both in mainly sublittoral beds on firm substrate around the low-water mark of spring tides, and in large eulittoral beds on mudflats between the low-water mark of neap tides and the uppermost level of the shore. Zostera noltii Hornem. occurs with eulittoral Z. marina on uppershore mudflats and also on exposed sandy shores near the low-water mark of spring tides. Since 1960 the eulittoral Zostera beds have spread widely on mudflats in Portsmouth, Langstone and Chichester Harbours, where they now occupy 430–450 ha. The Zostera beds at the bottom of the shore remain winter-green, but the mudflat Zostera beds shed most of their leaves between November and January. Comparisons are made with localities further east, which show earlier and greater leaf loss.The uppershore Zostera beds in the three harbours comprise an important and favoured autumn and early winter food for Dark-bellied Brent Geese, (Branta bernicla bernicla (L.)) and Wigeon (Anas penelope L.), which consume the leaves and rhizomes, and Teal (Anas crecca L.) which consume the seed. The pattern of occurrence of these three species of birds is, to some extent, related to the availability of Zostera.     

Reproduction and lipids in the sub-Antarctic limpet Nacella (Patinigera) macquariensis Finlay, 1927

The reproductive cycle of the sub-Antarctic limpet, Nacella (Patinigera) macquariensis Finlay, 1927 is described from monthly gonad indices and slide smears of gonadal tissue. Two populations were sampled, from the eulittoral zone and at ≈4 m depth into the sublittoral zone. Gametogenesis started 2 months earlier in the eulittoral zone population; the breeding period, however, was sufficiently long to overlap in the two populations. This phase difference in reproductive cycles is discussed with respect to possible genetic divergence and correlation between timing within the reproductive cycle and environmental factors. Lipid was a prominent component of both the digestive gland and the gonad, but not of the foot. Changes in lipid levels during the reproductive cycle indicated that the digestive gland stored lipids which were transferred to the gonad during gametogenesis.     

Population dynamics of intertidal oribatid mites (Acari: Cryptostigmata) from the subtropical archipelago of Bermuda

The population dynamics of the three intertidal oribatid species, Alismobates inexpectatus, Fortuynia atlantica and Carinozetes bermudensis, have been studied on the archipelago of Bermuda over the course of a year. All three species are univoltine, showing a clear seasonal demographic pattern, with reproduction from spring to late autumn and a complete standstill of egg production in winter. A seasonal shift in sex ratio could also be observed in all three species and is supposed to be based on sex-dependent mortality. The subtropical climate of Bermuda allows longer reproductive periods than shown in other intertidal or edaphic temperate species and temperature is supposed to be the main factor influencing the demography of these intertidal dwelling mites. Although all three Bermudian species exhibit the same basic seasonal demographic pattern, there are slight temporal shifts in population dynamics, presumably caused by local microclimatic differences among the populations. Larviparity, shown in other littoral oribatid mites, is clearly absent in the present species.     

Lack of metabolic temperature compensation in the intertidal gastropods,Littorina saxatilis (Olivi) and L. obtusata (L.)

Two intertidal snails, Littorina saxatilis (Olivi, 1972) (upper eulittoral fringe/maritime zone) and Littorina obtusata (Linnaeus, 1758) (lower eulittoral) were collected from a boulder shore on Nobska Point, Cape Cod, Massachusetts, in July and acclimated for 15–20 days at 4 ° or 21 °C. Oxygen consumption rate (Vo₂) was determined for 11–15 subsamples of individuals at 4 °, 11 ° and 21 °C with silver/platinum oxygen electrodes. Multiple factor analysis of variance (MFANOVA) of lo₁₀ transformed values of whole animal Vo₂ with log₁₀ dry tissue weight (DTW) as a covariant revealed that increased test temperature induced a significant increase in Vo₂ in both species (P<0.00001). In contrast, MFANOVA revealed that temperature acclimation did not affect Vo₂ in either L. saxatilis (P= 0.35) or L. obtusata (P= 0.095). Thus, neither species displayed a capacity for the typical metabolic temperature compensation marked by an increase in Vo₂ at any one test temperature in individuals acclimated to a lower temperature that is characteristic of most ectothermic animals. Lack of capacity for metabolic temperature acclimation has also been reported in other littorinid snail species, and may be characteristic of the group as a whole. Lack of capacity for respiratory temperature acclimation in these two species and other littorinids may reflect the extensive semi-diurnal temperature variation that they are exposed to in their eulittoral and eulittoral fringe/maritime zone habitats. In these habitats, any metabolic benefits derived from longer-term temperature compensation of metabolic rates are negated by extreme daily temperature fluctuations. Instead, littorinid species appear to have evolved mechanisms for immediate metabolic regulation which, in L. saxatilis and L. obtusata and other littorinids, appear to centre on a unique ability for near instantaneous suppression of metabolic rate and entrance into short-term metabolic diapause at temperatures above 20–35 °C, making typical seasonal respiratory compensation mechanisms characteristic of most ectotherms of little adaptive value to littorinid species.     

Description of Harpagifer permitini sp. nova (Harpagiferidae) from the sublittoral zone of South Georgia and redescription of the littoral H. georgianus Nybelin

Comparative morphology of Antarctic spiny plunderfishes of the genus Harpagifer from the littoral and sublittoral zones off South Georgia (Scotia Sea) is investigated. A new species H. permitini from the depth 65–80 m is described and the littoral H. georgianus Nybelin, 1947 is redescribed. The significant differences in their morphology are shown. Two morphological and bathopathic forms (“soft” and “spiny”) of spiny plunder-fishes are characterized.     

Die Makrofauna und ihre Verteilung im Nordost-Felswatt von Helgoland

The macrofauna and its distribution in the sheltered, rocky intertidal zone of Helgoland (North Sea) was studied at 9 vertically and/or morphologically different stations from March to September in 1984. Seasonal variations in the communities were described based on each species' conspicuousness. A total of 172 species was found. The macrofauna shows a zoned pattern, but also the different substrata, for example, affect its distribution. The number of species increases from the upper intertidal to the upper sublittoral zone from 23 to 133 species. The upper intertidal is characterized byLittorina saxatilis, Chaetogrammarus marinus andHyale nilssonii. Typical and abundant species of the middle and lower intertidal areFlustrellidra hispida, Littorina mariae/obtusata, Littorina littorea, Mytilus edulis andSpirorbis spirorbis. The upper sublittoral zone is characterized byGibbula cineraria and increasing species numbers of Bryozoa, Nemertini and Opisthobranchia. Only few species (e.g.Dynamena pumila, Laomedea flexuosa, Polydora ciliata, Fabricia sabella, Jaera albifrons, Carcinus maenas) occur in the entire intertidal zone. In comparison to other very sheltered shores in Great Britain, which are also dominated by Fucaceae, the macrofauna in the Helgoland intertidal zone lacks several littoral species, such asPatella spp.,Monodonta lineata, Gibbula umbilicalis, Littorina neritoides, Chthamalus spp., whereasLittorina littorea andGibbula cineraria are highly abundant.