Dynamics of species interaction strength in space, time and with developmental stage

Quantifying species interaction strengths enhances prediction of community dynamics, but variability in the strength of species interactions in space and time complicates accurate prediction. Interaction strengths can vary in response to density, indirect effects, priority effects or a changing environment, but the mechanism(s) causing direction and magnitudes of change are often unclear. We designed an experiment to characterize how environmental factors influence the direction and the strength of priority effects between sessile species. We estimated per capita non-trophic effects of barnacles (Semibalanus balanoides) on newly settled germlings of the fucoid, Ascophyllum nodosum, in the presence and absence of consumers in experiments on rocky shores throughout the Gulf of Maine, USA. Per capita effects on germlings varied among environments and barnacle life stages, and these interaction strengths were largely unaltered by changing consumer abundance. Whereas previous evidence shows adult barnacles facilitate fucoids, here, we show that recent settlers and established juveniles initially compete with germlings. As barnacles mature, they switch to become facilitators of fucoids. Consumers caused variable mortality of germlings through time comparable to that from competition. Temporally variable effects of interactors (e.g. S. balanoides), or spatial variation in their population structure, in different regions differentially affect target populations (e.g. A. nodosum). This may affect abundance of critical stages and the resilience of target species to environmental change in different geographical regions.     

Barnacle larval supply to sheltered rocky shores: a limiting factor?

In northwest Europe, sheltered rocky shores are dominated by fucoid canopy algae and barnacles are rare, although the latter are extremely abundant on exposed shores. The supply of the intertidal barnacle Semibalanus balanoides (L.) to sheltered, fucoid dominated rocky shores was investigated to determine the importance of larval supply in limiting the abundance of adults in shelter. Larval supply was measured at two spatial scales, at the scale of shore (100s of metres), by comparing larval concentrations at exposed and sheltered sites, and at a smaller spatial scale (m), by examining the role of fucoid canopies in limiting supply to the substratum. Replicate plankton trawls were carried out above the intertidal zone at high water at two sheltered sites and nearby exposed headlands. The concentration of S. balanoides cyprid larvae was significantly higher at the sheltered sites on two out of three sampling occasions with up to 14 times greater larvae on one occasion than the nearby exposed site. The effect of the macroalgal canopy on supply to the substratum was assessed in two ways: directly, by pumping water from the substratum in areas with and without Ascophyllum nodosum (L.) Le Jolis, and indirectly by measuring cyprid settlement in a canopy-manipulation experiment. Pumped plankton samples from mid tide level showed that the A. nodosum canopy did not form a barrier to larval supply and may have had a positive effect on larval concentrations at the substratum. Cyprid settlement was assessed in the mid shore A. nodosum and low shore Fucus serratus L. zones to areas with canopy algae (but protected from the sweeping effects of macroalgal fronds) and without canopy. Settlement over three consecutive 24-h periods showed a consistent pattern; settlement was consistently lower beneath the F. serratus canopy than in cleared areas, suggesting that this algal species forms a barrier, limiting supply of cyprid larvae to the substratum.     

Predatory behaviour of juvenile shore crab Carcinus maenas (L.)

Juvenile shore crabs Carcinus maenas (L.) were observed feeding on rock barnacles Semibalanus balanoides (L.) on a Bay of Fundy rocky shore. This previously unreported predatory behaviour was further investigated in the laboratory. When given a choice of three common and abundant gastropods, Nucella lapillus (L.), Littorina littorea (L.), and Littorina obtusata (L.), and the rock barnacle Semibalanus balanoides, juvenile shore crabs of both sexes ate mainly barnacles and consumed proportionately more barnacles than gastropods compared with adults, which ate mainly gastropods. The rock barnacle is an abundant and readily available food source which may be important in sustaining the juvenile crab through periods of moults and rapid growth. As the shore crab attains a certain age (size), it must forage lower on the shore as gastropods become more important in its diet.     

Resource levels and prey state influence antipredator behavior and the strength of nonconsumptive predator effects

The risk of predation can drive trophic cascades by causing prey to engage in antipredator behavior (e.g. reduced feeding), but these behaviors can be energetically costly for prey. The effects of predation risk on prey (nonconsumptive effects, NCEs) and emergent indirect effects on basal resources should therefore depend on the ecological context (e.g. resource abundance, prey state) in which prey manage growth/predation risk tradeoffs. Despite an abundance of behavioral research and theory examining state‐dependent responses to risk, there is a lack of empirical data on state‐dependent NCEs and their impact on community‐level processes. We used a rocky intertidal food chain to test model predictions for how resources levels and prey state (age/size) shape the magnitude of NCEs. Risk cues from predatory crabs Carcinus maenas caused juvenile and sub‐adult snails Nucella lapillus to increase their use of refuge habitats and decrease their growth and per capita foraging rates on barnacles Semibalanus balanoides. Increasing resource levels (high barnacle density) and prey state (sub‐adults) enhanced the strength of NCEs. Our results support predictions that NCEs will be stronger in resource‐rich systems that enhance prey state and suggest that the demographic composition of prey populations will influence the role of NCEs in trophic cascades. Contrary to theory, however, we found that resources and prey state had little to no effect on snails in the presence of predation risk. Rather, increases in NCE strength arose because of the strong positive effects of resources and prey state on prey foraging rates in the absence of risk. Hence, a common approach to estimating NCE strength – integrating measurements of prey traits with and without predation risk into a single metric – may mask the underlying mechanisms driving variation in the strength and relative importance of NCEs in ecological communities.     

Direct and indirect effects of Littorina littorea (L.) on barnacles growing on mussel beds in the Wadden Sea

On the extensive sedimentary tidal flats of the Wadden Sea, beds of the blue mussel Mytilus edulis represent the only major hard substratum and attachment surface for sessile organisms. On this substratum, the barnacle Semibalanus balanoides is the most frequent epibiont. In summer 1998, it occurred on over 90% of the large mussels (>45 mm shell length) and the dry weight of barnacles reached 65% of mussel dry weight. However, the extent of barnacle overgrowth is not constant and differs widely between years. Periwinkles (Littorina littorea) may reach densities >2000 m^–2 on intertidal mussel beds. Field experiments were conducted to test the effect of periwinkle grazing on barnacle densities. An experimental reduction of grazing and bulldozing pressure by periwinkles resulted in increased recruitment of barnacles, while barnacle numbers decreased with increasing snail density. The highest numbers of barnacles survived in the absence of L. littorea. However, a lack of periwinkle grazing activity also facilitated settlement of ephemeral algae which settled later in the year. Field experiments showed that the growth rate of barnacles decreased in the presence of these ephemeral algae. Thus, L. littorea may reduce initial barnacle settlement, but later may indirectly increase barnacle growth rate by reducing ephemeral algae. It is suggested that periwinkle density may be a key factor in the population dynamics of S. balanoides on intertidal mussel beds in the Wadden Sea.     

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.     

Effects of small-scale disturbances of canopy and grazing on intertidal assemblages on the Swedish west coast

The effects of small-scale disturbances (80×30-cm plots) of canopy and grazers on intertidal assemblages were investigated in this 4-year experiment on sheltered rocky shores on the Swedish west coast. Canopy disturbances due to ice scouring were mimicked by removal of adult plants of the seaweed Ascophyllum nodosum (L.) Le Joli. Density of the main epilithic grazing gastropods, Littorina spp., was lowered by exclosure and handpicking. Based on earlier experiments in other areas, the general hypothesis was that canopy removal and grazer exclosure, alone or in combination, should increase the recruitment of A. nodosum or other fucoid juveniles, and change the structure of the understorey assemblage.There was an effect of canopy removal on the development of this assemblage, lasting for more than 31 months. Both increased and decreased abundances of species were found as short-term effects, but there was also a longer-term effect with increased abundance. Grazer exclosure was only effective in combination with canopy removal, causing a short-term increase in ephemeral green algae. Short-term effects of canopy removal were also the increase in recruitment of Semibalanus balanoides (Linnaeus) and the decrease of the red alga Hildenbrandia rubra (Sommerfelt) Meneghini. Fast recruitment and growth of fucoid species (Fucus serratus L. and F. vesiculosus L.) restored the canopy and conditions of the understorey within 18 months. Thus, the canopy removal changed the physical conditions for the understorey, making it possible for other species to coexist in this community. Surprisingly, no effect of canopy removal or grazer exclusion was found on the recruitment of juvenile A. nodosum, neither by canopy removal nor grazer exclosure. The lack of such effects might be due to the early mortality caused by other grazers (small, mobile crustaceans), or to the low density of periwinkles on these shores. However, despite the patchy and generally low recruitment of A. nodosum juveniles, observations suggested that the cover of A. nodosum in manipulated patches would return to initial levels, either by recruitment or regrowth of small holdfasts and from growth of edge plants.     

Thermoregulatory behavior of the pink shrimp Penaeus duorarom Burkenroad

Measurements of Balanus balanoides (L.) distribution at Dipper Harbour, New Brunswick, Canada, indicate as individuals per unit area increase, external morphology changes. This trend is reflected largely by height increase. Directional aspect of substrate effects numbers of barnacles as well as their shapes.     

Divergent succession and implications for alternative states on rocky intertidal shores

It has been hypothesized that rockweed stands and mussel beds in sheltered bays in the Gulf of Maine, USA, are alternative community states. As a test of this hypothesis, experimental clearings of different sizes were established in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis to determine if successional changes in large clearings developed species assemblages distinctly different from the surrounding A. nodosum stands. Clearings ranging from 1 to 8 m in diameter were created at 12 sites in 4 bays on Swan's Island, Maine, in 1996 to mimic the effects of ice scour, and abundances of gastropods, barnacles, mussels and fucoid algae were monitored until 2002. ANOVAs and MDS showed strong effects of clearing size and divergent successional changes in large clearings. Large clearings were quickly filled in and remained dominated by the alga Fucus vesiculosus L. and the barnacle Semibalanus balanoides (L). There was no evidence for site-specific effects, and Mantel tests showed clearing size was a better predictor of species composition than geographic distances among sites. Results suggest that large pulse disturbances using clearings of 8 m in diameter can initiate divergent successional pathways and have a protracted effect on species composition. Results are also consistent with the hypothesis that mussel beds and rockweed stands in sheltered bays may be alternative community states.     

Selective settlement of the barnacle Semibalanus balanoides (L.) facilitates its growth and reproduction on mussel beds in the Wadden Sea

On the unstable sedimentary tidal flats of the Wadden Sea, a suitable attachment substrate for sessile organisms is generally lacking. Epibenthic mussel beds (Mytilus edulis L.) provide the only and strongly limited settlement sites available for the barnacle, Semibalanus balanoides (L.). Field investigations showed that barnacles were non-randomly distributed within a mussel bed. They preferentially occurred near the siphonal apertures of living mussels but rarely grew on dead mussels or shell fragments. Field experiments revealed that this was due to selective settlement of barnacle cyprid larvae. Growth of barnacles was significantly higher upon living mussels than on empty mussel shells. Moreover, a higher reproductive output was obtained by individuals on living mussels which produced twice as many nauplii larvae than barnacles attached to empty shells. This study shows that selective settlement of S. balanoides cyprid larvae on living mussels is adaptive with respect to individual fitness. Received in revised form: 15 January 2001 Electronic Publication     

Sediment on rocky intertidal reefs: Effects on early post-settlement stages of habitat-forming seaweeds

Modification of the coastal environment by human activities often leads to an increase in sedimentation of nearshore waters, with potential impacts on benthic marine assemblages. Here we assess the relationships between the levels of sedimentation, wave exposure and benthic organisms on rocky intertidal platforms around the Kaikoura Peninsula in southern New Zealand. We designed and tested five sediment traps to provide a tool for measuring the relative abundance of sediment across sites. Using field- and laboratory-based experiments, we tested hypotheses concerning whether different levels of sedimentation affected algal germling survival and algal zygote attachment, and whether the interactions of grazers and sediments affected germling survival. Levels of sediment and exposure were inversely related across seven sites. The fucoid alga Hormosira banksii characterized the more sedimented wave-sheltered and intermediately wave-exposed sites, with up to 80% cover in the lower mid-tidal zone, while the bull kelp Durvillaea antarctica characterized the three most wave-exposed sites. Grazing molluscs were found across all sites but species abundances varied by sediment and exposure levels. We did two 11-day trials testing the effects of different levels of sediment and different species of molluscan grazers on the survival of 1-week-old Hormosira germlings. Generally, there was no significant treatment effect of grazers, but mortality varied considerably among sites. In particular, one site had very high levels of sediment, which resulted in 100% mortality of germlings across all grazer treatments. Removing sediment at 1-week, 2-week and 4-week intervals made no difference to the survival of Hormosira germlings. In laboratory-based experiments, a light dusting of sediment reduced the percentage of zygotes of Hormosira by 34% and Durvillaea by 71% that attached to primary substratum, and a complete cover of sediment prevented attachment altogether. Overall, the effects of sediments and its interaction with molluscan grazing were highly variable but often large, particularly on the attachment of zygotes to primary substratum.     

Age-dependent zonation of the periwinkle Littorina littorea (L.) in the Wadden Sea

On sedimentary tidal flats near the island of Sylt (German Bight, North Sea) abundance and size distribution of periwinkles, Littorina littorea L., were studied in low intertidal and in shallow and deep subtidal mussel beds (Mytilus edulis L.). In low intertidal mussel beds, surveys revealed that high densities (1,369±571 m^–2) of juvenile snails (≤13 mm) were positively correlated with strong barnacle epigrowth (Semibalanus balanoides L. and Balanus crenatus Bruguière) on mussels. A subsequent field experiment showed that recruitment of L. littorea was restricted to the intertidal zone. Abundances of periwinkles (213±114 m^–2) and barnacles abruptly decreased in the adjacent shallow subtidal zone, which served as a habitat for older snails (>13 mm). L. littorea was completely absent from disjunct deep (5 m) subtidal mussel beds. Snail abundance varied seasonally with maxima of >4,000 m^–2 in low intertidal mussel beds in October and minima in July, just before the onset of new recruitment. I suggest that the presence of cracks and crevices among the dense barnacle overgrowth in intertidal mussel beds favoured recruitment and survival of juvenile snails. Larger (older) specimens are assumed to actively migrate to the less favourable adjacent subtidal. Therefore, intertidal mussel beds are considered as nurseries for the population of L. littorea in the Wadden Sea. Received in revised form: 25 September 2000 Electronic Publication     

Stylochus tauricus,a predator of the barnacle Balanus improvisus in the Black Sea

The flatworm Stylochus tauricus Jacubova has been found associated with the barnacle Balanus improvisus Darwin, on which it feeds. The predation rate (the number of barnacles eaten by one polyclad in a month) ranges between 5–10. Inside the empty shells of B. improvisus some egg-plates of S. tauricus were observed. Pelagic Götte's larvae aged 2–3 days possess 4 lobes while those aged 7–8 days have 5 lobes. Flatworms can prey on the young of another species Balanus eburneus Gould, whereas predation on the mussels Mytilus galloprovincialis Lam. is rare. There is a direct correlation between predator abundance and prey ingested.     

The habitat and feeding preferences of Littorina obtusata (L.) and L. mariae sacchi et rastelli

The vertical zonation of the flat periwinkles Littorina obtusata (L.) and L. mariae Sacchi et Rastelli is different, but overlaps. At a sheltered location on the Isle of Cumbrae, Scotland, L. obtusata, although most abundant on Fucus vesiculosus L. and Ascophyllum nodosum (L.) Le Jol. on the midshore, ranged over the entire shore from the Pelvetia zone down to the upper limits of Laminaria. Littorina mariae exhibited a much more restricted distribution being associated predominantly with Fucus serratus L. on the lower shore. In laboratory trials with adult plants, flat periwinkles found fucoid algae both more attractive and more palatable than other seaweeds. Littorina obtusata also preferred the reproductive receptacles of fucoid algae to their vegetative tissue. Tiny germlings of a diversity of seaweeds were readily consumed by L. obtusata. In choice experiments germling Ulva lactuca L. was preferred to adult U. lactuca, whereas adult Fucus serratus was preferred to germling F. serratus. Contrasting foraging behaviours between different species of flat periwinkle are discussed in relation to differences in their radular dentition.     

Barnacle growth rate on artificial substrate in the Salton Sea,California

The Salton Sea is one of the few saline, inland lakes in the world with a population of barnacles, Balanus amphitrite. It is also one of California’s most impaired water bodies due to excessive nutrient loading which leads to phytoplankton blooms and low dissolved oxygen. Currently, B. amphitrite growth is limited due to lack of hard substrate in and around the Sea. We have hypothesized that artificial substrate could support the growth of B. amphitrite and their filter-feeding would lead to improved water quality. Periodic harvesting of the barnacles would result in the permanent removal of nitrogen and phosphorus from the Sea. A 44-day in-situ experiment was carried out in the Salton Sea to assess the rate of barnacle growth and phosphorus and nitrogen sequestration on burlap sheets suspended vertically from a floating line. Burlap panels were collected weekly and the barnacles analyzed for Ca, total-P, inorganic-P, total-N, total-C, CaCO₃, and organic matter content. After 44 days of growth, the barnacle mats weighed 7.4 kg m⁻² on a dry weight basis, with 80% of the mass as shell material. The nutrient sequestration was 9.4 g P m⁻² and 100 g N m⁻². Approximately half of the P was inorganic and appears to be coprecipitated with the calcium carbonate shell material. Results indicate that harvesting barnacles grown on artificial substrate in the Salton Sea would not be an effective method for removing N or P from the lake because of the relative proportions of shell material and organic material. Guest editor: S. H. Hurlbert The Salton Sea Centennial Symposium. Proceedings of a Symposium Celebrating a Century of Symbiosis Among Agriculture, Wildlife and People, 1905–2005, held in San Diego, California, USA, March 2005     

Reciprocal Relationships Between Exotic Rusty Crayfish,Macrophytes, and <Emphasis Type="Italic">Lepomis</Emphasis> Species in Northern Wisconsin Lakes

Abstract The non-native rusty crayfish (Orconectes rusticus) has invaded many lakes of northern Wisconsin, profoundly changing littoral zones in the process. There are other lakes that have been invaded, but do not exhibit these changes. We hypothesized that endogenous feedbacks could form involving rusty crayfish, the macrophytes they destroy, and Lepomis species whose abundance is positively related to macrophyte abundance and also consume juvenile crayfish. We assessed this proposal with long-term data from one lake, a regional comparative study, and a case study of Lepomis predation on crayfish. Through time and across lakes, abundances of rusty crayfish, littoral macrophytes and species of the genus Lepomis were related in a fashion that indicated a set of feedbacks that regulate the abundance of all three. Intense predation on juvenile crayfish by abundant Lepomis is capable of maintaining some crayfish populations at low abundance. Thus, some lakes display profound ecological changes where crayfish achieve high abundance, and others sustain crayfish at low abundance. Consequently, lakes invaded by rusty crayfish may take on the appearance of alternative ecological regimes. Direct experimentation is necessary to determine if, and under what conditions, a lake can exist in either regime. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Optimal diet theory: behavior of a starved predatory snail

Summary The tenets of optimal foraging theory are used to contrast the behavior of the predatory snail Acantina spirata when feeding on the barnacles Balanus glandula and Chthamalus fissus under conditions of satiation and starvation. As predicted in optimal diet models, A. spirata is less selective (ratio of attack frequency on a prey species to number of individuals available) when the higher ranking prey has low abundance. When given a choice, starved snails attack both barnacle species equally, whereas satiated individuals preferentially attack B. glandula, the more profitable prey (ash-free dry weight of barnacles ingested per unit handling time). Under starvation conditions, equal attack frequency does not result in equal prey species consumption because Acanthina spirata is more successful at attacking C. fissus than B. glandula.The assumption of constant prey encounter rates in optimal diet models is not met when A. spirata goes from a state of satiation to starvation. The encounter rate on B. glandula is lowered due to a decrease in attack success. A loss of feeding skills in starved A. spirata is responsible for the greater difficulty snails have in gaining access through the opercular plates of B. glandula.Behavioral changes in A. spirata as snails pass from satiation to hunger translate into an energetic disadvantage during feeding for hungry snails for two reasons. First, higher prey handling times result in a decreased rate of biomass intake. Second, alteration in the relative attack frequency between barnacle species, combined with a decrease in attack success on the more profitable prey leads to more frequent ingestion of the less profitable prey.     

Chemical settlement inhibition versus post-settlement mortality as an explanation for differential fouling of two congeneric seaweeds

It has been proposed that seaweed secondary metabolites, e.g. brown algal phlorotannins, may have an ecologically important function as a chemical defence against epiphytes, by acting against colonisation of epiphytic organisms. We tested whether the low epiphytic abundance on the invasive brown seaweed Fucus evanescens, compared to the congeneric F. vesiculosus, is due to a more effective chemical defence against epiphyte colonisation. A field survey of the distribution of the common fouling organism Balanus improvisus (Cirripedia) showed that the abundance was consistently lower on F. evanescens than on F. vesiculosus. However, contrary to expectations, results from experimental studies indicated that F. vesiculosus has a more effective anti-settlement defence than F. evanescens. In settlement experiments with intact fronds of the two Fucus species, both species deterred settlement by barnacle larvae, but settlement was lower on F. vesiculosus both in choice and no-choice experiments. Phlorotannins from F. vesiculosus also had a stronger negative effect on larval settlement and were active at a lower concentration than those from F. evanescens. The results show that Fucus phlorotannins have the potential to inhibit settlement of invertebrate larvae, but that settlement inhibition cannot explain the lower abundance of the barnacle Balanus improvisus on F. evanescens compared to F. vesiculosus. Assessment of barnacle survival in the laboratory and in the field showed that this pattern could instead be attributed to a higher mortality of newly settled barnacles. Observation suggests that the increased mortality was due to detachment of young barnacles from the seaweed surface. This shows that the antifouling mechanism of F. evanescens acts on post-settlement stages of B. improvisus.     

The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

Direct and indirect effects of predation,herbivory and surface rugosity on mussel recruitment

Summary The predatory gastropod Nucella lapillus, commonly preys upon the mussel, Mytilus edulis, and is thought to control the distribution and abundance of mussels on the rocky shores of New England, USA. In this study, done in Maine, USA, not only the presence of Nucella lapillus but also the roughness of the experimental surface and the presence of the herbivorous gastropod, Littorina littorea, were manipulated. Four types of surfaces were used as recruitment substrata for mussels: smooth bare granite, aggregations of the barnacle, Semibalanus balanoides, fiberglass resin castings of smooth bare granite and resin castings of aggregations of S. balanoides. To ensure that caged N. lapillus were not starving, barnacles were provided as alternative prey. Experiments showed no detectable effect of N. lapillus on the recruitment of M. edulis. Mussel recruitment was enhanced by surface rugosity and depressed by the activities of L. littorea. Analysis of covariance, using the number of algal species as the covariate, suggested that L. littorea reduced the number of newlyrecruited mussels by removing algae that provided recruitment sites, but no manipulations were done to test this conjecture. It is likely that previous reports of N. lapillus controlling mussel abundance are attributable to N. lapillus preying upon barnacles, which increase surface rugosity and enhance mussel recruitment. Review of literature on feeding preferences of N. lapillus supports this view. When handling times and prey availability are taken into account, Nucella shows a clear preference for barnacles over mussels.