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.     

Repeat photography as a tool for detecting and monitoring historical changes in South African coastal habitats

Repeat photography was used to illustrate long-term changes occurring in coastal habitats in the Western Cape, South Africa. Historic images were sourced from books and theses, the public and subject specialists, and repeat photographs were then taken from the same perspectives. Visible changes could be categorised into four types: (1) changes in species’ ranges; (2) biological invasions; (3) sea level changes; and (4) direct engineering impacts. In terms of range changes, the images depict a progressive easterly spread of the cold-water kelp Ecklonia maxima and parallel easterly contraction of the warmer-water mussel Perna perna, both evidence for declining water temperatures along the South-West Coast. Since c. 1980 most shores have also become conspicuously invaded by the alien Mediterranean mussel Mytilus galloprovincialis, while those on the West Coast have also been visibly invaded by the more-recently introduced Pacific barnacle Balanus glandula. No changes in vertical zonation due to changing sea levels could be detected, despite suitable images being available. Construction along the shore has radically altered the appearance of the shoreline in urban areas. Repeat photography thus proved a useful tool for both detecting and dramatically illustrating historic changes over the past century. These changes have altered substantially both the appearance and ecological attributes of many rocky shores in this region.     

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.     

Biofilm: A crucial factor affecting the settlement of seaweed on intertidal rocky surfaces

We hypothesized that the presence of biofilm accelerated the settlement of dominant seaweed species and maintained high levels of species richness by regulating the blooming of particular species on an intertidal rocky shore. The coverage and species richness of macroalgae on sterile and cleared substrates (225 cm²) were measured to investigate the effects of biofilm on the settlement of macroalgae in intertidal zones at Noryang, Songmoon, and Sangnam in Gawngyang Bay on the southern coast of the Korean peninsula. Green algae coverage on cleared substrates was significantly higher than that on sterile substrates at both Noryang and Songmoon during the study period. This suggests that the presence of biofilm enhances the settlement of green algae by providing various habitat structures and, consequently, may lead to serious ‘green tide’ events. However, the coverage of algae other than green algae and algal species richness on cleared substrates remained high at Sangnam during the experimental period. Biofilm facilitated the settlement of macroalgae and inhibited the blooming of specific algae by inducing inter-specific space competition. Therefore, biofilm plays an important role on seaweed assemblages on intertidal rocky shores by accelerating the settlement of seaweed.     

Breeding,settlement and survival of barnacles at artificially modified shore levels at Leigh,New Zealand

The breeding and settlement seasons of the intertidal barnacles Chamaesipho brunnea, C. columna and Elminius plicatus were determined, and experiments using cages on the shore to examine barnacle survival and growth rate at four levels were carried out at Leigh during 1964–66.

Chamaesipho brunnea bred in spring and summer; C. columna and E. plicatus bred throughout the year, but settled intermittently. When protected from predators and from overgrowth by algae, all three species survived at levels lower than normal. At higher levels C. columna, E. modestus, Balanus trigonus and Tetraclita purpuraseens survived for significant periods (C. brunnea occurs naturally higher on the shore than the highest experimental level).     

An approach to ecological monitoring in the rocky intertidal: A survey of Bantry Bay and Dunmanus Bay,Ireland

The methodology of a monitoring study of the fauna and flora of the rocky intertidal zone in Bantry Bay and Dunmanus Bay in southwest Ireland is described and some representative results are presented. Five shores, four in Bantry and o one in Dunmanus Bay, were visited monthly from May, 1978 to June, 1980, and more limited monitoring is continuing. On each visit to each shore, four research programmes were undertaken—transect studies, recolonisation of cleared areas, barnacle studies and limpet studies. The transect methods are discussed in relation to other methods in current use. Combining community and key species approaches is shown to be most useful in monitoring programmes. The increased effectiveness of continuous shore monitoring when compared with one-off or before and after impact-type studies is stressed.     

Barnacle fouling in the Mediterranean sponges Axinella polypoides and Axinella verrucosa

Secondary metabolites protect many marine sponges (Phylum: Porifera) from settlement by fouling organisms. Previous studies on the subtidal demosponge Axinella verrucosa collected in the Western Mediterranean led to the isolation of compounds that inhibited the settlement of cyprids larvae of the intertidal barnacle Balanus amphitrite, and the enzyme chitinase, which plays a key role in the molting cycle of crustaceans. However, in a field survey conducted at three locations in Israel, Eastern Mediterranean Sea, we observed that A. verrucosa is fouled by the subtidal barnacle Balanus trigonus, a previously unknown association. Settlement inhibition assays using B. amphitrite with chemical extracts from Israeli A. verrucosa and Axinella polypoides, a sympatric, congeneric sponge that seems not to be fouled by B. trigonus, showed that cyprid larvae of B. amphitrite were inhibited by the extracts of both sponges from settlement at concentrations several magnitudes lower than natural volumetric extract concentration in the sponges. These results indicate that, unlike the intertidal barnacle B. amphitrite, the subtidal B. trigonus is unaffected by the compounds from A. verrucosa, stressing and underlining the importance of using suitable target organisms (i.e. from the same habitat) to test for ecologically relevant antifouling activities.     

舟山岩相潮间带动物群落结构的研究

本文应用数理统计方法研究了舟山海区岩岸潮间带中26个测站的春季动物群落结构。根据26个站的位置分为近岸区、近外海区和外海区3个区域。各站均以软体动物和甲壳动物为群落主要组成成分,群落种类组成数从近岸到外海呈增长趋势,且优势种亦从低盐种类向高盐种类演替。近岸区群落主要优势种为短滨螺Littorina brevicula、粗糙滨螺L.scabra、僧帽牡蛎Ostrea cucullata、白脊藤壶Balanus albicostatus等;近外海区为短滨螺、粒结节滨螺Nodilittorinaexigua、僧帽牡蛎、鳞笠藤壶Tetraclita squamosa squamosa、白脊藤壶等;外海区为粒结节滨螺,疣荔枝螺Purpura clavigera、条纹隔贻贝Septifer virgatus、日本笠藤壶T.japonica及藻类等。但群落个体密度和生物量从近岸到外海呈减少趋势。从近岸到外海,群落的Shannon-Wiener多样性指数H和Pielou均匀度J均增加,而McNaughton优势度D减少。群落结构从近岸到外海渐趋复杂。但整体上,舟山海区岩岸春季动物群落结构是比较简单的(绝大多数断面H值小于2.50,J值小于0.5。而D值大于0.6)。近外海区的群落结构比较相似于外海区的群落。根据平均聚类分析,这26个群落可以分为三个生态类型,即隐蔽性岩岸群落,半隐蔽性岩岸群落及开敞性岩岸群落。     

宁波海岛期间带生态学研究──Ⅱ．数量组成与分布

本文研究了宁波海岛潮间带的生物数量组成与分布规律。岩礁潮间带平均生物量（２５１２．９４ｇ／ｍ２）高于沙质潮间带（４７．６４ｇ／ｍ２）高于泥沙质潮间带（２２．３６ｇ／ｍ２），而密度为岩礁潮间带（９８７．２个／ｍ２）高于泥沙质潮间带（２８１．２个／ｍ２）高于沙质潮间带（２０．０个／ｍ２）。主要生物类群有甲壳动物、藻类及软体动物。岩礁潮间带生物量从近岸向外海增加。泥沙质潮间带的生物量和密度从北至南增加。     

舟山潮间带生态学研究：Ⅱ.数量及其分布

舟山潮间带生物种类数量的平面分布与盐度和海岸开敞性有关,垂直分布与潮汐有关;群落数量的水平分布与底质和海岸开敞性有关,垂直分布与潮汐有关。舟山潮间带生物有明显的分带现象,隐蔽的岩岸区分为:滨螺带和藤壶-牡蛎带两个带;开敞性岩岸区分为:滨螺带,藤壶-(虫戚)-藻类带及藻类带3个带。沙滩区分为:沙蟹带和浅蛤-斧蛤带两个带;泥沙滩区分为:盐碱植物-滨螺带和招潮蟹-大眼蟹带;泥滩区分为:蟹守螺-囊螺带和蟹守螺-织纹螺带。     

The effect of life-history variation on the population size structure of a rocky intertidal snail (Littorina sitkana)

On wave-sheltered shores of the northeastern Pacific, the population size structure of Littorina sitkana varies with intertidal height, as larger snails are mostly found only in the upper intertidal. This pattern has been attributed to high predation rates by crabs (and perhaps fish) on large snails inhabiting low-intertidal areas; i.e., large snails are presumed to be rare there simply because predators kill them. In this study we investigate the hypothesis that predation contributes to the shore-level size gradient displayed by L. sitkana by selecting for (or inducing) earlier sexual maturation and reduced somatic growth in low-shore snails relative to high-shore individuals.In the first part of our study, we carried out laboratory dissections, field experiments (mark-release-recapture and caging), and field surveys on a wave-protected shore in Bamfield Inlet, Barkley Sound (British Columbia, Canada). The principal results were: (1) adult survivorship was greater at higher, than at lower, intertidal level, (2) snails displayed a preference for their shore level of origin, (3) immature adults from the high intertidal displayed greater rates of somatic growth relative to immature adults from the low intertidal, and (4) low-shore snails matured at a smaller size than high-shore individuals. In the second part of the study, a large-scale survey showed intra-specific variation in size at sexual maturity (point 4 above) to be relatively consistent over time (winter of 1999 and 2001 for snails from our main study site) and space (13 different sites in winter 2001), although the magnitude of these differences varied greatly from shore to shore.Our results indicate that L. sitkana individuals inhabiting upper and lower parts of their intertidal range allocate resources differently to somatic and gonadal growth, an intra-specific difference that is best interpreted as a response to spatial and size-dependent variation in predation pressure. Taken together, results of this and other recent studies indicate that phenotypic responses to contrasting selection pressures operating in upper- and lower-intertidal areas contribute to the intertidal size gradient of L. sitkana. We believe that greater consideration of evolutionary processes in ecological studies will lead to a more complete understanding of the mechanisms responsible for structuring marine coastal communities.     

Distribution,Population Structure, ­Population Biomass and Morphological Characteristics of the Tunicate Pyura stoloniferain the Bay of Antofagasta,Chile

Abstract. The present study analyzes the distribution, population structure, population biomass and morphological characteristics of the intertidal tunicate Pyura stolonifera (Heller, 1878) in the Bay of Antofagasta, Chile, where the species shows a reduced geographical range extending around 60 – 70 km. Three sites along its distribution were studied. At the central site the tunicate presents the widest intertidal belts, reaching nearly 11 m (the mid‐intertidal‐Pyura‐zone shows the higher percent cover). Population dry biomass here averaged up to 20.45 kg·m^–2, representing one of the highest intertidal specific biomasses reported in the literature. Lower values occur towards the borders of its distribution. Population densities were over 1800 indiv.·m^–2 at mid‐intertidal‐Pyura‐belts. Crowding and competition potentially affect morphological characteristics and the energy/tissue allocation along the intertidal and geographical gradient of P. stolonifera.     

Shore exposure affects mussel population structure and mediates the effect of epibiotic algae on mussel survival in SW Ireland

On rocky shores, the relative importance of abiotic and biotic processes that regulate community structure are thought to vary with levels of shore exposure. This can lead to characteristic features found on sheltered and exposed shores. This study identified differences in the population structure of mussels on exposed and sheltered rocky shores on Atlantic coasts of south-west Ireland. Direct interactions between epibiotic algae and their host mussels were also examined to test if potential effects varied with shore exposure. Mussel beds on sheltered shores were less dense and comprised larger mussels with greater rates of individual survival and growth than those on exposed shores. The results of a field experiment showed that algal epibionts had a negative effect on mussel survival on sheltered shores but not on exposed shores. Surprisingly, the presence of algal epibionts had no effect on mussel growth on either shore type. These findings contrast with those of previous studies. The effects of shore exposure and algal epibionts on mussels may be species-specific and may interact with other factors across different regions. This study shows that predictions of effects of exposure on mussel populations and their epibionts should only be based on specific experimental evidence and cannot be generalised across regions.     

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.     

Habitat use of inter‐tidal chitons – role of colour polymorphism

This study presents a new approach to examining European chiton habitat use according to morphological characteristics. Time‐search transects were conducted on four Portuguese rocky shores at two shore levels. The chiton species, abundance and individual morphometric features were recorded and related to the substratum dominant epiphytes and surface roughness. The chiton Lepidochitona cinerea (Linnaeus 1767) was the dominant species (77%), followed by Leptochiton algesirensis (Capellini 1859) (15%) and Acanthochitona crinita (Pennant 1777) (8%). Species abundance did not vary significantly with shore level or amongst shores. Six colour morphs were considered for Lepidochitona cinerea, four for A. crinita and one for Lepidochiton algesirensis. Substratum characteristics significantly explained the variation in species and colour morph distribution. There was a high correlation between the presence/absence and type of epiphytic algae and the substratum surface roughness level. We hypothesize that three mechanisms operating either independently or in interaction may explain the non‐random distribution of colour polymorphisms according to their habitat characteristics: (i) presence of algae that are part of their diet; (ii) use of rock as shelter – for instance, crevices and cover resulting from algae fronds and; (iii) morph–rock colour matching as a camouflage mechanism to reduce predation pressure.     

Secondary production of the eusirid amphipod <Emphasis Type="Italic">Pontogeneia rostrata</Emphasis> Gurjanova, 1938 (Crustacea: Peracarida) on a sandy shore in Korea

The secondary production of the eusirid amphipod Pontogeneia rostrata Gurjanova, 1938, was examined on a sandy shore in Korea. Secondary production was estimated using a size-frequency method. The biomass structure of P. rostrata included one peak in spring, with maximum biomass occurring in April. The annual secondary production of this species was 0.57 g DW m⁻² yr⁻¹ with an annual P/B ratio of 6.97. Secondary production of P. rostrata fell within the range observed for other amphipods on intertidal sandy shores, although the production/biomass (P/B) ratio was higher than those recorded previously from the sampling site and from other tidal and estuary areas, as the number of generations increases the P/B ratio. The results showing high abundance and P/B ratios suggest that P. rostrata plays an important role in the sandy shore ecosystem as a trophic link between primary producers and higher consumers.     

The invasion of the intertidal canopy-forming alga Fucus serratus L. to southwestern Iceland: Possible community effects

The intertidal serrated wrack, Fucus serratus L. (Fucales: Phaeophyceae), has become an abundant canopy-forming alga along a ca. 100 km stretch of shore in southwestern Iceland in the last century. Its distribution has not changed noticeably since a survey in 1975/1976. Many experimental studies have shown that canopy-forming algae can have profound effects on the community structure. Although such experiments involving F. serratus are few it was decided to test predictions from these experiments on a geographical scale by comparing community compositions within the area where F. serratus is a dominant algae (since at least 1975/1976) with an adjacent area of similar size where F. serratus is absent (with a single exception without consequences). The work is based on measurements on some 372 stations in the F. serratus area, termed region A, and 227 stations from the F. serratus-free area, termed region B. Percentage cover of algae and sessile invertebrates was estimated on 2 m² on each station and animals collected from 800 cm² from each station. The vertical distribution of F. serratus was most similar to that of Fucus distichus and the two species often grew intermingled on the lower part of the shore. F. serratus appeared to have reduced the cover of F. distichus in the lowermost part of the shore, while having little or no effect on other canopy-forming species. The low abundance of Semibalanus balanoides in the F. serratus area (region A) is in line with experiments showing detrimental effect on this barnacle by whiplash of F. serratus. In general algal grazers were more abundant in region A, consistent with the greater attractiveness of F. serratus than other canopy-forming species to grazers, although this may be partly explained by the abundance of understorey algae in region A.     

Trophic relationships on a fucoid shore in south-western Iceland as revealed by stable isotope analyses, laboratory experiments, field observations and gut analyses

Rocky shores in the North Atlantic are known for their zonation patterns of both algae and animals, which can be expected to greatly affect food availability to consumers at different height levels on the shore. We tested the hypothesis that consumers would feed on the most abundant suitable food source in their surroundings. In total 36 species/taxa of common primary producers and consumers were sampled for stable isotope analyses from a sheltered fucoid shore at Hvassahraun in south-western Iceland. A selection of these species was also collected seasonally and from different height levels. Feeding experiments, field observations and gut analyses were also conducted. Our results were in good overall agreement with pre-existing knowledge of trophic relationships in the rocky intertidal. Consumers often appeared to be assimilating carbon and nitrogen from the most common diet in their immediate surroundings. The predator Nucella lapillus was thus feeding on different prey at different height levels in accordance with different densities of prey species. When tested in the laboratory, individuals taken from low on the shore would ignore the gastropod Littorina obtusata, uncommon at that height level, even when starved, while individuals from mid-shore readily ate the gastropod. This indicated that some kind of learned behaviour was involved. There were, however, important exceptions, most noteworthy the relatively small contribution to herbivores, both slow moving (the gastropod L. obtusata) and fast moving (the isopod Idotea granulosa and the amphipod Gammarus obtusatus) of the dominant alga at this site, Ascophyllum nodosum. The recent colonizer Fucus serratus seemed to be favoured. Selective feeding was indicated both by isotope signatures as well as by results of feeding experiments. Seasonal migrations of both slow and fast moving species could partly explain patterns observed.     

Effects of high pressure,hot water shore cleaning after oil spills on shore ecosystems in the Northern Baltic proper

The use of high pressure, hot water hosing techniques in oil spill clean-up operations on rocky and stony-gravelly shores drastically reduces the shore vegetation and macrofauna. The negative effects are more substantial than on oiled shores cleaned by raking and scraping. After one year the hot water cleaned shores were not restored completely.On rocky shores the high pressure, hot water technique is very efficient in terms of freeing the rocks from oil. However, due to its detrimental effects on shore organisms this type of oil spill clean-up operation can only be recommended for bird or wildlife protection areas. The cleanup method is inefficient on stony-gravelly shores due to penetration of oil into the ground and sediment and direct killing of shore organisms. The method should be avoided on these types of shores.