The distribution of molluscs on littoral rocks in Ceylon,with notes on their ecology

Molluscs inhabiting littoral rocks were collected from 43 stations around Ceylon, and 57 species belonging to 20 families were identified. An account is given of the nature and distrbution of littoral rocks around the island, and the stations examined. On each shore, the distribution of the molluscs was studied in relation to the broad zonation of the shores themselves and observations were made on plant and animal associations of each speices. The limits of the broad zones of the shores were defined with reference to biological indicators. Analysis of the data collected led to a classification of the rocky shores into three groups, each characterized by particular combinations of mollusc species and distinct patterns of shore zonation. Some molluscs were found to be distributed over the whole island, others to be common to two groups of shores only, and still others to be confined to a single shore type. An attempt has been made to explain the presence or absence of species on various groups of shores, and to explain discontinuities in distribution on the basis of environmental factors such as the nature of the substratum, degree of wave exposure, and weed cover. A key, based primarily on external features, has been devised for the identification of the species listed.     

Seaweed richness and herbivory increase rate of community recovery from disturbance

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities.     

A mathematical model for the distribution and abundance of benthic algae in a Norwegian fjord

A mathematical model has been developed to study the distribution and abundance of benthic algae species in a Norwegian fjord. The main forcing functions are the physical factors which have well defined gradients in this area, but biotic interactions are also accounted for in the model.The distribution of algae along vertical transects in the intermediate fjord area has been simulated, and the observed distribution is reproduced fairly well. This indicates that the major factors determining the equilibrium elevation between intertidal and subtidal vegetations in this area are adequately represented in the model.Based on observations in a fjord branch with a high level of freshwater influx and polluted by heavy metals, assumptions are made of the combined influence of these contaminants on growth characteristics of the species considered. The resulting effects on community development and community patterns are simulated. For a certain level of contamination, the decrease in abundance, community development rate and species separation have been determined. It is also shown that the fluctuations in biomass during a year are more pronounced than under unaffected conditions.The relative importance of external growth regulating factors and self-shading is demonstrated for different degrees of community development, and we have indicated that this may be an important factor in the assessment of transient and long-term responses to “natural” and man-made influences on the shore-line vegetation.One of the main problems during model development was the formulation of the interaction function for factors influencing growth parameters. Several alternatives were examined.The differences between the most frequently used forms were small with respect to abundance and not very large with respect to possible changes in distribution. It was noted that even by including only the most limiting factor at each instant, (Liebig's form) a high percentage of the potentially limiting factors did actually limit growth during periods of the year.The model is intended as a working tool for the studies of changes in the shore-line vegetation caused by man-made interferences like heavy metal and oil pollution, and we believe that the results presented demonstrate that it can successfully be used for this purpose.     

An assessment of variation in molluscan grazing pressure on Hong Kong rocky shores

Several methods of assessment have been used to document variation in grazing pressure on temperate rocky shores, although often these methods are applied without consideration of local conditions or species. In this study, a comparison was made between abundance counts of inactive molluscan grazers at low tide, direct observations of grazer activity and distribution throughout day and night tidal cycles, and records of grazing marks on wax discs, for the mid-shore of Hong Kong. The abundance of grazers found during low-tide counts varied among dates, sites and species. This method, however, did not record all grazer species that day/night observations showed to migrate from the low shore with the rising tide. Low-tide counts, therefore, underestimate grazing pressure (number of active grazers per unit area) and grazer guild (number of species). Grazing marks on wax discs also recorded a greater number of species than the low-tide counts of inactive grazers, and included grazers that were seen to migrate up shore during day/night observations. Certain limpet species, however, avoided the wax and did not leave grazing marks, showing this method to also underestimate grazing pressure. All methods showed grazing pressure to be variable at spatial scales of tens of metres or less and also temporally variable between sampling dates. The sole use of either low-tide counts or wax discs is likely to underestimate grazing pressure, due to variation in shore topography and grazer foraging behaviour, especially on shores with a narrow tidal range such as in Hong Kong. To gain a more accurate assessment of total grazing pressure, it is suggested that recording of grazing marks on wax discs should be used in conjunction with direct day/night observations.     

Habitat structure and complexity as determinants of biodiversity in blue mussel beds on sublittoral rocky shores

Habitat-forming, ecosystem engineer species are common in most marine systems. Still, much uncertainty exists about how individual and population-level traits of these species contribute to ecosystem processes and how engineering species jointly affect biodiversity. In this manipulative field experiment, we examined how biodiversity in marginal blue mussel beds is affected by blue mussel (1) body size, density and patch context and (2) presence of fucoid and algal structures. In the study area, bladder-wrack (Fucus vesiculosus), filamentous algae and blue mussels (Mytilus edulis) coexist at shallow depths in a variety of patch configurations and offer complex habitats with a high variability of resources. We hypothesized that complexity in terms of mussel bed structure and algal presence determines species composition and abundance. Results from the experiment were compared with macrofaunal communities found in natural populations of both engineering species. Results show that the physical structure and blue mussel patch context are important determinants for species composition and abundance. Results further show that the presence of algal structures positively affects diversity in blue mussel habitats due to increased surface availability and complexity that these algae offer. This study shows that blue mussel beds at the very margin of their distribution have an indisputable function for promoting and maintaining biodiversity and suggest that facilitative effects of habitat-modifying species are important on Baltic Sea rocky shores with fundamental importance to community structure.     

Amphipods and isopods in the rocky intertidal: dispersal and movements during high tide

Animals in the intertidal, both mobile and sessile, generally exhibit some zonation pattern, in which each species shows a preference for, or is confined to, some height levels. The study of zonation patterns is, however, almost exclusively based on surveys made during low tide, when many animals are relatively inactive. We studied zonation patterns of amphipods and isopods on rocky shores in southwestern Iceland, both by traditional sampling at low tide as well as by sampling during high tide. The distributional patterns seen at high tide differed significantly from that at low tide. One amphipod, Anonyx sarsi, was common around baits at all levels at high tide but absent from the intertidal at low tide. Several other species were either relatively more common or tended to be recorded higher, or in one instance, lower on the shore when the tide was in than at low tide. There was also evidence of some species changing habitats within the intertidal with the tidal cycle. Many species, however, moved little away from their respective zones occupied at low tide, and for some species, including some capable of rapid swimming, very limited mobility was indicated. We conclude that low-tide surveys of the intertidal give an incomplete picture of the community structure, and even key species may be missed in such surveys.Communicated by L. Hagermann, Helsingør     

Microhabitat segregation of the amphipod genus Gammarus (Crustacea: Amphipoda) in the Northern Baltic Sea

Closely related species may occupy very similar niches but are often found to diverge by one or more traits when they share the same habitat. Five indigenous and sympatric Gammarus species are characteristic for the Baltic rocky littoral ecosystem. Yet, the species-specific distribution of these sympatric gammarids has not been well studied in the northern Baltic Sea. This study was undertaken to assess the spatial distribution of gammarid amphipods along wave exposure and depth gradients to study whether they show segregation in their microhabitat use. We sampled 12 rocky sublittoral shores along a wave exposure gradient over a period of 5 years. Samples differed with respect to depth and macroalgal type. Three of the five gammarid species occurred mainly in different depth zones and among different macroalgae at the exposed shores. In contrast, on protected shores, where algal zonation is weaker, a link to zonation and macroalgal type was almost absent. Moreover, the microhabitat use was strongest during the reproductive seasons of the species. The observed microhabitats of the three gammarid species fit well to their species-specific mean body sizes.     

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

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

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.     

Effect of canopy removal on a rocky shore community metabolism and structure

The effects of the permanent removal of the canopy-forming alga Fucus serratus was studied in terms of both functioning and diversity on a mid–low rocky shore ecosystem in the south-west English Channel (48°N 43.686′, 3°W 59.282′). Ecosystem functioning was examined as net or gross primary productivity (NPP or GPP) and respiration (Resp) measured through CO₂ fluxes. Diversity was examined as number and composition of species. Measurements were performed in situ, during emersion times, without altering target assemblages. The experiment was designed with two treatments [control (C) or canopy removed (CR)] and five replicates, and was conducted over an 18-month period (from February 2006 to August 2007) to integrate the seasonal variability. The mean GPP and Resp were severely reduced in CR treatment compared to control throughout the survey. The mean NPP was not affected at first, due to the development of opportunistic green algae, but was drastically reduced after 9 months of experiment. The canopy removal affected neither the number of species nor their distribution among trophic groups, and the algal community was only slightly affected. The abundance and biomass of mobile invertebrates, however, were greatly reduced in the absence of canopy. This indicates an important effect of the dominant alga on the higher trophic levels of the community. At this tidal level, the canopy did not seem to affect the community by dampening the environmental stress but by providing food, habitat or both.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Effects of brackish water incursions and diel phasing of tides on vertical excursions of the keystone predator Pisaster ochraceus

The physical factors that constrain the vertical foraging excursions of the keystone predator, the sea star Pisaster ochraceus, hold considerable interest because they indirectly shape the vivid patterns of zonation of rocky shore communities by impeding or enhancing the ability of P. ochraceus to traverse the intertidal zone. In this paper, we describe a study conducted in the Pacific Northwest of North America in which we examined, in the field and laboratory, the abiotic factors that can affect vertical excursions by P. ochraceus. Our field observations revealed that the extreme upward reach and average shore level height reached by P. ochraceus were significantly lower for daylight high tides than nocturnal high tides. Based on diver observations following a severe storm, it would also appear that these diurnal movements can be impeded by freshwater incursions into the intertidal zone; a regularly occurring event in the Pacific Northwest. As part of an experimental investigation into this phenomenon, we observed that sea stars maintained in tall cylindrical aquaria, without tidal flux, remained near the bottom during daylight and moved to the top of the column at night, suggesting that photoperiod alone can influence the cycle of vertical movement. Adding a freshwater layer to the aquaria restricted these vertical excursions. Our results suggest that on rocky coastlines susceptible to fresh water incursions, the suppression of foraging may be an important factor in the spatial and temporal variation in the intensity of predation. Furthermore, given the relative increase in frequency and intensity of freshwater incursions in the Pacific Northwest and the intolerance of P. ochraceus to lowered salinity, there is the long-term potential to significantly alter patterns of species zonation in this essential marine habitat.     

Quantitative analysis of foraging behaviour in a field population of the South American sun-star Heliaster helianthus

Foraging behaviour in the South American sunstar Heliaster helianthus (Lamarck) was investigated quantitatively on a subtropical rocky shore in central Peru. H. helianthus feeds mainly on two mussel species, Semimytilus algosus (Gould) and Perumytilus purpuratus (Lamarck). A sequence of foraging behaviour was described and observations were made of the timing of attacks on mussel beds by sun-stars. Although H. helianthus is capable of foraging out of water, an unusual trait for asteroids, the timing of such foraging appears to be well adjusted to avoid the risk of prolonged heat and desiccation. Foraging activity began 4 h 52 min before high tide and 3 h 19 min after high tide, with a mode between 2 h 30 min and 2 h before high tide. Fifty-two percent of all foraging activity began between 3 h 30 min and 2 h before high tide, while only 12.4% began after high tide. This suggests that H. helianthus mainly relies upon changes in the rate of tidal increase as a cue to begin foraging. Foraging activity ceased between 3 h 40 min before and 4 h 29 min after high tide, with 48% ending between 2 and 0.5 h before high tide. The duration of foraging ranged from 19 to 190 min, with values between 30 and 80 min accounting for 67.4% of all observations. The median duration was 62 min. No significant correlation was detected between the time when foraging activity commenced and its duration. The intensity of foraging activity varied on consecutive days, with a general pattern of decreasing intensity after a day of relatively high activity. Foraging location in relation to a mussel bed was analysed on a marked, 8 m stretch of rocky shore. The numbers of foraging H. helianthus observed on different sections of the shore were related neither to the width of the mussel zone nor to the vertical position of the lower edge of the mussel zone, indicating that sun-stars do not rely upon these factors to assess prey availability and that ideal free distribution with regard to prey abundance does not occur on the spatial scale examined.     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Ecological effects of intertidal uplifting from nuclear testing

Uplifting of a portion of an intertidal rock bench by the Milrow underground nuclear test at Amchitka Island, Alaska affected algal abundance, species composition, and zonation. Parts of the populations of 8 species died off, primarily in the Hedophyllum sessile zone, and mostly within the first 6 months after the disturbance. The center of abundance of these species shifted seaward. Fucus distichus, a species of the upper intertidal, appeared in the area and recolonized a portion of the H. sessile zone. The observations indicate that intertidal algal communities at Amchitka Island are very sensitive to positive elevation changes.Contribution No. 376 of the College of Fisheries, University of Washinton, USA. This publication is based on work performed under U.S. Atomic Energy Commission Contract AT(26-1)-171 for Battelle Memorial Institute, Columbus Laboratories.     

A comparison of macrofaunal and meiofaunal distribution and standing stock across a rocky shore,with an estimate of their productivities

In 1984 and 1985 algal, macrofaunal and meiofaunal standing stocks were estimated on a exposed rocky shore along the west coast of False Bay, South Africa, using comparable, area-based sampling techniques. The shore supported a rich growth of algae, particularly in summer, when a maximum standing crop of 403 g m^-2 was recorded in the low shore. In winter, the largest component of macrofaunal biomass comprised the filter-feeding barnacle Tetraclita serrata, which attained 75 g m^-2 in the middle balanoid; but as a result of late recruitment and high mortality of this species, the summer shore was dominated by herbivorous gastropods, particularly Patella cochlear, which attained a maximum biomass of 66 g m^-2 on the low shore. Meiofaunal numbers and biomass were closely correlated to the distribution of algal turfs and associated trapped sediments. Numerically, the most important components of the meiofauna were nematodes and copepods, while the biomass was more evenly shared among foraminifera, minute gastropods, copepods and insect larvae. Numbers and biomass peaked in the lower balanoid during winter (1.9×10⁶ individuals, or 8.5 g m^-2). Macrofauna:meiofauna numbers and biomass ratios are presented for each zone and the distribution patterns discussed in relation to the conditons in each. Numerically, meiofauna exceed macrofauna by an overall ratio of 1:391, with values ranging from 1:556 in the lower balanoid to 1:18 in the Littorina zone. Macrofaunal biomass exceeds that of meiofauna in all zones by an overall ratio of 10:1, but values range from 2.1:1 in the upper balanoid to 48:1 in the middle balanoid. By incorporating turnover ratios extrapolated from the literature, mean annual productivity ratios have been calculated. These indicate that macrofauna account for 75% of total secondary production and meiofauna for 25%. Failure to incorporate meiofauna in analyses of energy flow on rocky shore ecosystems would thus lead to considerable errors. The possible trophic role of meiofauna in such systems is discussed.     

Short-term effects of an oil spill on the West coast of the Cape Peninsula, South Africa

In 1994, the sinking of the ‘Apollo Sea’ off the West coast of South Africa led to the deposition of ca. 2 500 tons of heavy fuel oil over 150 km of coastline. The impact of the spill on rocky shore invertebrates, rock-pool fish fauna and rock lobsters was assessed by conducting surveys shortly after the spill, and again two months later. Where possible, results of these surveys were compared with existing data from before the sinking of the Apollo Sea. Among the fish fauna of rock-pools, changes in total density of fishes were largely due to changes in the abundance ofClinus superciliosus and were within the range of natural variation for the species and the community as a whole. Community structure of the rock-pool fish fauna also remained unaffected. At three of four impacted rocky shore sites no changes could be detected in overall benthic community structure, although the winkleNodilittorina africana was affected. At the fourth site, a boulder beach, statistical analysis showed distinct differences in community structure between heavily and lightly contaminated areas, as well as between all areas compared with previously existing data. There were also significant changes in the lightly oiled areas between the first and second surveys after the spill. Oil-fouled lobster were found at one of three sites investigated. Ca. 7 % of the seabed in this particular area was polluted. Antennae and forelegs of almost all lobsters in the vicinity of the oil were fouled. Examination of the gut-contents confirmed that oil inhibits the ability of lobster to feed. Although there was no evidence that mortalities of lobster were caused by the spill, growth rates may be reduced by the decreased feeding rate. Overall, the impact of the spill was strikingly less than might have been predicted from the effects of other oil spills. Nomenclature: Names of fish species follow Smith & Heemstra (1986).     

Vertical zonation of endosymbiotic zooxanthellae within a population of the intertidal sea anemone,Anthopleura uchidai

Intertidal organisms commonly form zonation bands along the shore. Environmental stressors often determine the vertical position of each zonation band. These stressors may similarly affect the distribution pattern of endogenous species in their intertidal hosts. To evaluate this possibility, we investigated the distribution pattern of endosymbiotic zooxanthellae in the genus Symbiodinium in a population of the intertidal sea anemone Anthopleura uchidai. We used molecular genetics to identify the Symbiodinium clades and found that A. uchidai has two clades of Symbiodinium, clades A and F. These Symbiodinium clades were disproportionally distributed along the vertical gradient of the intertidal shore. Anemones on the upper shore exclusively possessed clade F Symbiodinium while clade A Symbiodinium became dominant in the sea anemones on the lower shore. Photosynthesis activity assays showed that these Symbiodinium clades had similar net productivities at 23.3 and 31.8 °C at all irradiance levels. At 35 °C, however, clade A Symbiodinium exhibited substantially lower net productivities than clade F Symbiodinium, demonstrating that these Symbiodinium clades have distinct tolerances to thermal stress. These results suggest that the thermal gradient across tidal height is a major factor shaping the zonation pattern of Symbiodinium clades in A. uchidai.     

Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Horizontal,vertical and seasonal distributions of epiphytic hydrozoa on the algaHalimeda tuna in the Northwestern Mediterranean Sea

The thalli ofHalimeda tuna (Ellis & Solander) grow horizontally from rocky surfaces and present an upper and lower surface which can be colonized by epiphytes. Thalli ofH. tuna were collected twice a month from March 1985 to March 1986 at Tossa de Mar (northeast Spain). Thirty five species of hydroid were recorded living onH. tuna thalli; of these, the 15 most abundant species were classified into three groups on the basis of their horizontal zonation on the thalli: those most abundant on the proximal, medial or distal articles. Those hydroids common on the proximal (oldest) articles were relatively large and were present throughout the year, those common on the medial articles were smaller and often occurred in dense monospecific patches, while those common on the distal (youngest) articles were present for only short periods and were the most opportunistic hydroids found onH. tuna. We hypothesize that this zonation reflects interspecific niche-selection, enabling successful competition for space with other hydroids, algae and bryozonas. Three hydroid species were more abundant on the upper surfaces of the algae than on the lower, probably because of gravitationally enhanced settlement of planula larvae coupled with (unusually) limited hydrorhizal growth on the underside. Three opportunistic species were more common on the lower surfaces of the thalli, possibly reflecting the greater availability of substratum in the shade, where epiphytic algae were less abundant. Many hydroids declined in abundance in summer because of the death of old thalli, the growth of new thalli and of apical articles on existing thalli, and possibly because of interspecific competition with epiphytic algae. The death of old thalli and the growth of new thalli reduced the abundance of epiphytic algae; this may have facilitated the increase in abundance of hydroids recorded in the autumn.