Habitat provision of barnacle tests for overcrowded periwinkles

In habitats where competition for space is a shaping force of animal distribution such as in the intertidal rocky ecosystem, new habitats are readily taken by colonising species. We examined the importance of empty Chthamalus spp. tests as a habitat for the intertidal common periwinkle Melaraphe neritoides on Portuguese rocky shores. The role played by the space between neighbouring barnacles as a habitat for other species has been largely studied with regard to how an ecosystem functions, whereas the equivalent role of empty barnacle tests remains largely unknown. The small periwinkle is one of the most abundant snails in European rocky shores and is an important prey for key mobile predators. Biological facilitation is common in the rocky intertidal zone, where biological structures often potentiate the abundance of other species. The role played by barnacles as biological facilitators through habitat provision is not fully understood. In this study, the abundance and morphometric features of empty barnacle tests and their occupants were examined across shore levels and shores with differing exposures, as these are important gradients explaining barnacle distribution. The availability of empty barnacle tests was also experimentally manipulated to examine the percentage, time and length of occupation. Empty barnacles were more abundant on the midshore of sheltered shores and barnacle tests were wider on the upper shore but taller on the midshore. The minimum barnacle test occupation rate by the periwinkle was 70%. Barnacle shell height was an important factor determining snail occupation, where taller barnacles harboured significantly more, but not necessarily larger, snails. Snails outside the barnacle tests were significantly larger than those found within, suggesting that this crustacean group has an increased importance as a habitat for juvenile snails, thus potentially influencing the population dynamics of M. neritoides. We found that occupation of experimentally created empty barnacles was extremely high (70%) on the day after, and remained at 100% after 3 months of monitoring. Our study is the first of its kind to focus on the features of barnacle tests that snails occupy and their occupying snail traits. According to our results, it is clear that barnacles have an important role in providing additional habitat for young gastropods. The small periwinkle is an important grazer and prey for intertidal and subtidal foraging predators; hence, the potential refuge role of barnacle tests for juvenile M. neritoides may be important in the dynamics of intertidal communities.     

The Decline of the North Sea Whelk (Buccinum undatum L.) Between 1970 and 1990: A Natural or a Human-Induced Event?

Abstract. Recent surveys in the southern and central North Sea and the Dutch Wadden Sea revealed that the whelk Buccinun undarum L. can no longer be found in areas where whelks were abundant twenty or more years ago.
The decline of whelk populations tends to coincide with areas of various, intense human activities, which may cause either pollution or physical damage. Potential causes for the decline of whelks from the southern North Sea and coastal areas are discussed and considered in view of North Sea ecology and protective policies for the North Sea.     

Feeding ecology of whelks on an intertidal sand flat in north‐eastern New Zealand

The dispersion patterns and feeding behaviour of intertidal whelks were investigated in north‐eastern New Zealand. Aggregations of whelks feeding on clams occurred regularly, with an average of 15–20 individuals per aggregation. Death of clams was attributable to predation in up to 50% of the aggregations, although scavenging of dead and moribund animals was the main activity of whelks. Laboratory and field studies showed that Lepsiella scobina (generally not considered a soft shore species) was the main whelk predator at Lews Bay, Whangateau Harbour, drilling a small neat hole through the clam shell. The distinction between Cominella spp. as predators or scavengers is not so clear. Reseeding of intertidal clams, Austrovenus stutchburyi, has been proposed as a technique for reestablishing populations in some degraded New Zealand estuaries. The impact of whelk predation has many implications for A. stutchburyi reseeding. Both L. scobina and Cominella adspersa may potentially prey on newly reseeded clams as they preferentially attacked small clams in the laboratory. On‐growing to a larger size before reseeding may be advantageous although large size was not found to protect prey from predation. Additionally, no seasonal trend in whelk activity was found.     

Succession of a wharf‐pile fauna at Lyttelton,New Zealand

A 31/2‐year succession of wharf‐pile fauna is described from communities of different known ages on piles at Lyttelton, New Zealand. Intertidal regions of the piles were initially colonised by the barnacle Elminius modestus; these populations developed to form a subclimax community which was subsequently replaced by a climax community of the mussel Modiolus neozelanicus after 21/2 years. The stability of this climax is discussed. In the infratidal region initial settlement of the tubicolous amphipod Corophium acherusicum and an erect bryozoan Bugula sp. was followed after 5 months by colonisation by solitary tunicates. These tunicates became dominant only after H years and provided numerous niches for superficial foulers, crevice dwellers, and motile organisms. This succession is compared and contrasted with a terrestrial succession of plants. Four vertical zones were recognised: supralittoral zone, midlittoral zone, in‐fralittoral fringe, and infralittoral zone. Species additional to those recorded by Skerman (1958) are appended.     

Report on the Annual Gametogenesis and Tissue Biochemical Composition in the Gray mussel,Crenomytilus grayanus (Dunker 1853) in the Subtidal Rocky Bottom on the East Coast of Korea

Ocean Science Journal - The Gray mussel Crenomytilus grayanus occurs in high density in the shallow rocky subtidal zone on the east coast of Korea, where the mussel dominates the shallow subtidal...     

The whelk (Buccinum undatum L.) in the western Dutch Wadden Sea in the period 1946–1970: Assessment of population characteristics and fishery impact

In the Dutch Wadden Sea a whelk fishery existed from 1900 to 1970. The high post-war catches, following a period without fishery, and the subsequent decrease in catches when fishery was reopened, allowed us to estimate fishery mortality and recruitment. Information on natural mortality and growth, which were required in the estimation procedure, was obtained from previous studies elsewhere. Natural mortality could be calculated from an experiment with marked whelks in the Thames estuary. Age-length relations could be used from French investigations on whelks near the Channel Islands. The size of a potential whelk population could be estimated. Effects of fishery and possible causes of the disappearance of the whelk from the Dutch Wadden Sea are discussed.     

Deja vu? A second mytilid mussel,Semimytilus algosus,invades South Africa's west coast

A second marine mussel is shown to have invaded South Africa's west coast. Molecular techniques, based on intraspecific gene sequence divergences, prove its identity as Semimytilus algosus, a member of the family Mytilidae, native to Chile. The identity of an older introduced population found in Namibia is also confirmed. The present geographic range of S. algosus in South Africa extends some 500 km, from Groenriviersmond in the north to Bloubergstrand in the south. Together with Mytilus galloprovincialis, another previously established invasive mussel, S. algosus has become a dominant intertidal organism on wave-exposed rocky shores across this region. Both species are now much more abundant intertidally than either of the indigenous mussels Aulacomya ater and Choromytilus meridionalis, which have become largely confined to sublittoral and sand-inundated habitats respectively. The two invasive mussels display strong spatial segregation, with M. galloprovincialis dominating the midshore and S. algosus blanketing the lower shore. Through a combination of its small size and high abundance, S. algosus is likely to greatly increase food availability for a range of intertidal predators, many of which cannot consume mussels above a threshold size.     

Mass mortality and recolonization of Pyura stolonifera (Heller) on the south coast of South Africa

The fauna and flora of two sublittoral communities situated in close proximity on the west coast of South Africa are described and shown to be radically different, despite the fact that both experience similar physical conditions. One of these communities, at Marcus Island, has prolific beds of the black mussel Choromytilus meridionalis (35 954 g wet weight·m^?2) accompanied by a large number of other species, notably sea urchins, holothurians, brittle stars, whelks and barnacles. In contrast, the other site, Malgas Island, is dominated by a large population of rock lobsters Jasus lalandii together with a dense seaweed flora (3 866 and 4 402 g wet weight·m^?2 respectively). The numbers of species are similar at both sites (102 at Malgas Island and 107 at Marcus Island) but only 34 per cent are common to both localities. Although preliminary research indicates that the two sites have fairly similar physical conditions and bottom topography, they maintained their contrasting communities for the whole research period (1983–1986) and have probably done so for at least nine years, suggesting that they represent two stable alternative states of the same ecosystem. In the absence of rock lobsters at Marcus Island, wave action and inter/intraspecific competition are the main forces structuring the benthic community. Stability is maintained here by the resilience and rapid recovery of the mussel population after disturbance by storms. Where rock lobsters are abundant, as at Malgas Island, intense predation pressure is the main force controlling the structure of the benthos, and it is continually maintained because the rock lobsters are resistant to the effect of storms. Therefore, the presence or absence of high densities of rock lobsters is fundamental to the maintenance of these two contrasting communities.     

Predators on rocky shores in the northern Atlantic: Can the results of local experiments be generalized on a geographical scale?

Experimental manipulations of invertebrate prey and predators on rocky shores have been done by many authors. In the northern Atlantic the predators involved are usually the green crab Carcinus maenas and/or the dogwhelk Nucella lapillus, and the prey species studied are acorn barnacles (balanid Cirripedia), mussels (Mytilus spp.) and winkles (Littorina spp.). Usually the predators are found to have a regulating “top–down” effect on the prey species. In Iceland the acorn barnacle Semibalanus balanoides, the blue mussel Mytilus edulis and the flat periwinkle Littorina obtusata (including to some extent Littorina mariae) are found on rocky shores all around Iceland in what would seem to be near-optimal physical conditions. The predators Carcinus maenas and Nucella lapillus, on the other hand, are relatively southern species that do not thrive on the colder coasts of Iceland. Thus general surveys of different coasts of Iceland would seem to offer opportunities to see whether the results of local experiments can be discerned on a geographical scale (hundreds of km). The roughly 4900 km of the rocky coastline in Iceland was in this study subdivided into four regions, I–IV, according to the commonness or presence of the two predators. With the hope of reducing compounding factors the surveys were confined to sheltered or semi-sheltered fucoid shores, which were further divided into Ascophyllum (more sheltered) and Fucus vesiculosus (less sheltered) shores. Estuaries or other low-salinity environments were avoided. The study was based on 761 stations distributed around the rocky coastline on these two types of shores. The results for barnacles and mussels, being generally more abundant in regions were predators were scarce or absent, and being less common in Ascophyllum than F. vesiculosus shores in contrast to the predatory dogwelks, were in large measure in accord with predictions from experiments indicating “top–down” regulation. The results for the periwinkles, however, suggested that “bottom–up” regulation might be of greater importance, as their density was closely linked to the biomass of macroalgae, while the abundance of predators appeared to have little effect. The periwinkle, although certainly eaten, may not be a preferred prey of the two predators in question in Iceland. This paper forms a link between experimental and observational study approaches.     

Stramonita haemastoma as a bioindicator for organotin contamination in coastal environments

Stramonita haemastoma was investigated as a suitable bioindicator of TBT and TPhT contamination in the tropical Atlantic Ocean by: 1. Imposex induction in healthy females after inoculation with TBT and TPhT in the laboratory; and 2. Determining incidence of imposex in S. haemastoma collected from areas with various levels of tributyltin (TBT) and triphenyltin (TPhT) and determining the concentrations of these chemical in its tissues and that of its prey, the mussell Perna pernas. Imposex intensities and organotin concentrations in tissues showed good correlation, indicating S. haemastoma as a reliable bioindicator of TBT and TPhT contamination in coastal waters. Body burden threshold of TBT and TPhT for imposex induction was estimated to be 10-20 ng g(-1).     

Differential recruitment of introduced Pacific oysters and native mussels at the North Sea coast: coexistence possible?

Pacific oysters (Crassostrea gigas Thunberg 1793) have been introduced into the Wadden Sea (North Sea), where they settle on native mussel beds (Mytilus edulis L.), which represent the only extensive insular hard substrata in this soft-sediment environment. As abundances of C. gigas rose, some mussel beds became increasingly overgrown with oysters, whereas others did not. Field experiments revealed that recruitment of C. gigas was higher in the lower intertidal than in the upper subtidal zone, that it was higher on conspecifics than on mussels, and that it was not affected by barnacle epigrowth except when settling on mussels. Mussel recruitment is known from inter- and subtidal zones. It occurred equally on oyster and mussel shells but showed a clear preference for barnacle epigrowth over clean shells. Assuming that settlement and recruitment are key processes for species abundances on the North Sea coast, it is predicted that the positive feedback in oyster settlement will lead to rapid reef formation of this invader at the expense of mussel beds. Mussels, however, may escape competitive exclusion by settling between or on the larger oysters especially when barnacles are abundant. Experimental patches with mussels were more often covered by fucoid algae (Fucus vesiculosus forma mytili Nienburg) than patches with oysters, and oyster recruitment was poor underneath such algal canopies. Thus, fucoids may provide the native mussels with a refuge from the invading oysters and the two bivalves may coexist, provided food is not limiting.     

Meiobenthic diversity in space and time: The case of harpacticoid copepods in two Mediterranean microtidal sandy beaches

Meiobenthic data from two microtidal sandy beaches of the eastern Mediterranean (Crete, Greece) were used to investigate patterns of both alpha and beta diversity in space and time. Copepod assemblages and environmental variables related to sediment characteristics, morphodynamics and food were studied over a year at four distinct habitats at each beach; the retention, resurgence and saturation zones of Salvat's intertidal scheme (midlittoral zone), and the surf zone of the sublittoral. Αlpha diversity analysis indicated similar species richness at both beaches when the whole 13-month data set was considered but was higher at the sheltered site when each sampling period was examined separately. Both beaches supported higher diversity in the sublittoral zone. Species richness increased seawards at the midlittoral zone of the sheltered site whereas, no pattern was evident at the exposed site, where the intense hydrodynamic conditions homogenized the sediments. Beta diversity increased markedly towards the sublittoral, indicating greater differences in alpha diversity between the sublittoral and the midlittoral zone. Species turnover was more variable at the exposed beach and at the most landward stations, where environmental conditions change often between extremes. A proportion of the variation in alpha diversity was explained by food availability at both beaches and additionally by grain size at the sheltered site. However, no environmental variable explained beta diversity patterns. Although the results of our study support the hypothesis of Multicausal Environmental Severity proposed for sandy beach macrofauna, we believe the classic Intermediate Disturbance Hypothesis is a more appropriate framework for the meiofauna communities of the studied sites.     

Demersal fishing disturbance increases predation risk for whelks (Buccinum undatum L.)

Field observations by divers indicated that a high rate of predation of whelks (Buccinum undatum) by starfish (Asterias rubens) occurred in an area disturbed by scallop dredging, although these whelks mostly appeared to be alive and externally undamaged. The ability of whelks to escape from starfish was tested in the laboratory after they were dropped or rolled to simulate direct physical contact with bottom fishing gear. Dropping whelks did not significantly affect their escape behaviour, but whelks which had been rolled took significantly longer to right themselves and were significantly less likely to perform an escape response than whelks that had not experienced this treatment. This study suggests that demersal fishing may indirectly increase whelk mortality by increasing their risk of predation.     

Foraging ecology of an endemic shorebird, the African Black Oystercatcher (Haematopus moquini) on the south–east coast of South Africa

We investigated small–medium (1–300 km) scale variation in the foraging ecology of the African Black Oystercatcher during its breeding season, using traditional diet analysis coupled with carbon and nitrogen stable isotope analysis. Fieldwork was conducted between January and March 2006 and 2007, on rocky shores on the south–east coast of South Africa at East London, Kenton and Port Elizabeth. Middens of shelled prey left by adults feeding their chicks were collected from five territories and the abundances of the collected prey on the foraging areas were estimated using quadrats. Blood samples from 45 birds (16 females, 10 males and 19 chicks) and tissues from the predominant prey species on the territory of each breeding pair were collected for isotope analysis. The Manly–Chesson selectivity index revealed that adults feed their chicks preferentially with the limpet Scutellastra cochlear and the Mediterranean mussel Mytilus galloprovincialis, if available. A slight enrichment in the ¹⁵N stable-carbon isotope signature was observed towards the west in both prey and oystercatchers. Differences in isotope signatures between males and females from the same breeding pair indicate sex-related differences in the diet. Both had signatures indicating a mixed diet, but with males exhibiting a signature closer to that of limpets and females closer to that of mussels. In the single case where mussels were rare on the feeding territory, the two members of a pair showed carbon signatures which were identical and very similar to that of limpets. These results indicate dietary partitioning between genders in breeding pairs.     

南极菲尔德斯半岛潮间带南极帽贝的种群生态学研究──夏季种群数量变化和垂直分布

于1994年12月—1995年3月（南极夏季）期间,对南极菲尔德斯半岛卵石滩、岩石滩潮间带南极帽贝的种群数量、垂直分布进行了定点、定时、定量的调查和观察。结果表明,南极帽贝的种群数量有明显下降,分别仅为起始数量的35％和52％。卵石滩潮间带的帽贝分布于中潮区及低潮区,尤其是低潮区,且有随季节（冬季来临）向潮下带移动的趋势。岩石滩潮间带的帽贝具有类似的垂直分布特征,但其在分布带上的发生频次有很大差异,呈波动状态。在垂直岩面上,帽贝数量在低潮区的底部更为集中。讨论了导致数量变化、垂直分布的可能原因。今后有必要深入研究种群数量的时空动态及其调控机理。     

Marine invasive aliens on South African shores: implications for community structure and tropillc functioning

Up to 12 marine mollusc and three crab species are thought to have been deliberately or accidentally introduced to South Africa. Of these, only the Mediterranean mussel Mytilus galloprovincialis and the European shore crab Carcinus meanas have become invasive. M. galloprovincialis, probably introduced in the late 1970s, is already the dominant mussel throughout the Cape west coast. As a result, indigenous intertidal mussels Aulacomya ater have been largely displaced, mussel standing stock has increased and the upshore limit of mussel beds has become considerably elevated. Implications include competitive exclusion of large limpets but enhanced recruitment of juvenile limpets, increased habitat availability for mussel infauna, reduction of habitat for algal infauna and enhanced food availability for predators, particularly terrestrial species. C. maenas was first recorded from Table Bay Docks in 1983 and currently ranges from Camps Bay to Saldanha Bay. Although it is a voracious predator, it appears unable to colonize wave-swept shores, so is unlikely to displace indigenous crabs, or to severely impact prey species on the open coast. Valuable conservation areas and mariculture sites in sheltered lagoonal areas are, however, threatened by it.     

The colonization of artificial substrata by marine sessile organisms in False Bay. 1. Community development

Marine sessile communities developing on stainless steel test panels set 10 and 20 m deep in False Bay over periods ranging from a month to a year were examined. The establishment of communities was characterized by seasonal succession, colonization rates being lowest during winter. Community development differed between depths. At 10 m, barnacles appeared first but numerous mussels settled on them, eventually smothering them. Later, secondary settlement of barnacles on the mussels increased the bulk of the layer. Initial development was slower at the 20 m site, progressing from barnacle dominance to a more complex community including barnacles, serpulids, hydrozoans and compound ascidians. The settlement of solitary ascidians during summer led to their final dominance alongside barnacles. It is suggested that solitary ascidians and barnacles would normally dominate at sublittoral depths up to 20 m in False Bay, except where mussel dominance develops first.     

Small is profitable: No support for the optimal foraging theory in sea stars Asterias rubens foraging on the blue edible mussel Mytilus edulis

Doubt has been shed recently on the most popular optimal foraging theory stating that predators should maximize prey profitability, i.e., select that prey item that contains the highest energy content per handling time. We hypothesized that sea stars do not forage on blue mussels according to the classical optimal foraging theory but are actively avoiding damage that may be caused by e.g. capture of foraging on too-strong mussel shells, hence the sea stars will have a stronger preference for mussels that are smaller than the most profitable ones. Here we present experimental evidence of the sea star Asterias rubens as a predator that indeed chooses much smaller blue mussels Mytilus edulis to forage on than the most profitable ones. Hence this study does not support the optimal foraging theory. There may be other constraints involved in foraging than just optimizing energy intake, for example predators may also be concerned with preventing potential loss or damage of their foraging instruments.     

Coupling bio-logging with nutritional geometry to reveal novel insights into the foraging behaviour of a plunge-diving marine predator

It is widely believed that predators maximise their energy intake while foraging and consume prey that are nutritionally similar. We combined GPS data loggers, miniaturised cameras, dietary sampling and nutritional geometry to examine the nutritional variability in the prey and selected diet, and foraging performance, of the masked booby (Sula dactylatra tasmani), a wild carnivore and marine top predator. Data loggers also revealed no significant differences between sexes in the foraging performance of chick-rearing adults. Females provided more food to their chicks than the males and, regardless of the nutritional variability of prey consumed, both sexes showed similar amounts of protein and lipid in their diets. Miniaturised cameras combined with nutritional analysis of prey provided, for the first time, fine-scale detail of the amounts of macronutrients consumed in each plunge dive and the overall foraging trip. Our methodology could be considered for future studies that aim to contribute to the general understanding of the behavioural and physiological mechanisms and ecological and evolutionary significance of animal foraging (e.g. energy expenditure budgets and prey selection for self- and offspring-feeding that could lead to sex-specific foraging strategies).     

Patterns of distribution and abundance of the stalked barnacle (Pollicipes pollicipes) in the central and southwest coast of continental Portugal

The stalked barnacle Pollicipes pollicipes is a cirriped crustacean that lives on very exposed rocky shores. This barnacle is the most important economical resource on intertidal rocky shores of continental Portugal. It is highly prized as food and heavily exploited (professional and recreational fishery), but fishery data are scarce and do not estimate the real pressure upon this resource.Despite its socio-economic interest, specific regulations on this fishery are recent and different along the Portuguese coast. Four regions with different regulation can be identified: the marine reserve “Reserva Natural das Berlengas” (RNB) and the marine park “Parque Marinho Prof. Luiz Saldanha” (PMLS) (both in central Portugal); the natural park located in SW Portugal (“Parque Natural do Sudoeste Alentejano e Costa Vicentina”, PNSACV); and the rest of the coast.The main objective of the present study was to study the spatial patterns of percentage cover, biomass, density and size structure of P. pollicipes in areas with different exploitation regimes, including harvested areas and no-take areas. Additionally, variability between mid shore and low shore barnacles was also analysed.Seven areas were sampled with a variable number of sites (a total of 24) randomly sampled in each area during 2011. Photographs and image analysis (percentage cover) and destructive sampling (density, biomass and size) were used.In general, percentage cover, biomass and density were higher in mid shore when compared to low shore, namely in harvested areas. Low shore barnacles had a higher proportion of adults with moderate and high commercial value, while juveniles were relatively more abundant at mid shore.There were no consistent differences in the patterns of distribution and abundance of P. pollicipes among areas subject to different exploitation regimes. The most different area was the harvested area by professional fishers in RNB, where the highest biomass within the study was registered (mid shore, 7.7 kg·m^− 2). Barnacles within this area presented a higher proportion of adults with commercial value, while recruits and juveniles were relatively more abundant in other areas.The hypothesis of a highest percentage cover, density and biomass in the low shore of no-take sites was not supported.