Limpets as prey: Lipophrys pholis choice for size

Patellid limpets are key species on rocky shores, structuring intertidal assemblages through their grazing. Their role as prey for fish is, however, often overlooked in studies of intertidal ecosystem functioning. The shanny Lipophrys pholis is a common predator of limpets on rocky shores in Northern Europe and, in this study, we examined the characteristics of its predation on limpets based on prey size. The limpet size range that adult L. pholis (10–12.5 cm) can eat was examined in the laboratory using Patella depressa in 0.2 cm classes, varying between 0.5 cm and 3.0 cm in maximum shell length. There was a limpet size refuge above 1.8 cm, while all smaller sizes were readily consumed by the shanny. The predator attacking behaviour was also examined and found to vary with prey size. Limpets up to 0.8 cm were crushed by the jaw‐constricting force and eaten whole, whereas larger prey were prised from the substratum and the body subsequently separated from the shell. To examine whether there was a L. pholis preference for P. depressa size, a two‐stage laboratory experiment was done using two size classes defined as small (0.7–0.8 cm maximum shell length) and large (1.5–1.6 cm maximum shell length). In the first stage, the predator was given each limpet size class separately. In the second stage, the fish was given a choice between the two classes. Lipophrys pholis was shown to have a preference for the large size class (1.5–1.6 cm). The average number of limpets consumed by the shanny was examined for the duration of one high‐tide typical of the central region of Portugal (≈ 4 h). On average, approximately five limpets (up to an eight limpet maximum) were consumed. The present study shows that L. pholis has a feeding preference based on limpet size, which suggests that the limpet population structure and intra‐ and inter‐specific interactions may be influenced by the shanny predation.     

The Effect of Human Exclusion on the Population Structure of Key-Hole Limpets Fissurella crassa and F. limbata on the Coast of Central Chile

Abstract. We describe the basic ecology of two key-hole limpets, Fissurella crassa and Fissurella limbata , which are heavily harvested by coastal shellfishermen (mariscadores) at the rocky intertidal of central Chile. The ecological role of human predation on these species was assessed. Anthropic exclusion from a stretch of the rocky intertidal at Las Cruces (non-harvested area) for about two and a half years resulted in changes of the densities and size distributions of these fissurelid species as compared to populations in control zones (harvested area). Differential human predation generates, on key-hole limpets, responses of different intensities related to the accessibility of man to the mesohabitats (sheltered platforms, exposed platforms and vertical walls) and fringes of the intertidal (mid-low intertidal and Lessonia nigrescens fringe). Thus, size distributions and mean densities of both fissurelids differ widely between the harvested and non-harvested areas on both platform habitats but not on vertical walls. In the mid-low intertidal fringe, differences are greater than in the less accessible L. nigrescens intertidal fringe. Moreover, we discuss the way in which other species, directly harvested by mariscadores (i. e., the mollusc Concholepas concholepas) can affect the keyhole limpet populations of central Chile rocky shores. Finally, we highlight the ecological role of these limpets on the dynamics of intertidal ecosystems and how they can modify community landscape.     

Different cardiac response to copper in limpets from metal polluted and clean shores of Hong Kong

The gastropod limpet, Cellana grata, is common on moderately exposed to exposed Hong Kong rocky shores. On the basis of the existing literature on trace metal contamination in Hong Kong, four sites were identified, representative of highly polluted (Aberdeen and Lei Yue Mun) and clean (Cape d'Aguilar and Clear Water Bay) conditions. Limpets from the two polluted sites showed similar, but significantly higher body concentrations of copper than limpets from the two clean sites, which were also similar. Non-invasive measurements of cardiac activity of limpets from the four sites showed between site differences in baseline heart rates under standard seawater, irrespective of pollution level. When acutely exposed to water borne copper (2 h, 0.5+/-0.06 SD ppm), however, limpets from clean sites showed a significantly higher increase in inter-beating time (bradycardia) than those from polluted sites. These results highlight the potential use of cardiac activity to assess the exposure of natural populations of limpets to trace metal pollution.     

Reproduction in the intertidal limpet Cellana ornata in southern New Zealand

The reproductive cycle of the abundant intertidal limpet Cellana ornata (Dillwyn 1817) was examined over 19 months, covering two breeding periods, on a rocky intertidal platform in southern New Zealand. A gonad index was calculated, and histological sections were used to count the numbers of mature and immature eggs present. Fecundity was estimated using counts of eggs before and after spawning. To determine any variation between sites, the gonad index of limpets at a second site dominated by boulders was examined over 6 months during the second breeding period. C. ornata had a single spawning period annually during summer, with greatest gonad sizes in January‐February. There were significant differences in peak gonad indices between two breeding seasons on the platform. Within the second breeding period, both the timing and magnitude of peak gonad development differed between sites, with lower reproductive output at the boulder site. Fecundity increased with increasing size, but individuals are probably capable of reproducing after their first year on the shore. The life history of this species is discussed in comparison to other limpets in New Zealand.     

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。     

Historical demography of southern African patellid limpets: congruence of population expansions,but not phylogeography

Global climatic oscillations have shaped the contemporary genetic structure of marine taxa in different ways. Previous demographic studies have indicated that various intertidal marine species display genetic signatures of demographic expansion that either pre- or postdate the Last Glacial Maximum. Such expansions and the ability of species to colonise new habitats will influence their genetic structure, but the link between scales of larval dispersal and the strength of phylogeographic structure is not always clear. We analysed a fragment of the mitochondrial COI gene of 11 sympatric species of intertidal southern African patellid limpets to investigate how ancient oceanographic dynamics have shaped and maintained their contemporary spatial genetic variation. Our data show that the patellid limpets investigated display congruent evidence of spatial expansion during the Late Pleistocene or Early Holocene, which corresponds with the establishment of the contemporary southern African shoreline. We argue that closely related and co-distributed southern African intertidal invertebrates responded to ancient climatic oscillations as a cohesive group. In contrast, contemporary oceanographic circulation has shaped the phylogeographic patterns of these limpets in different ways. We show close relationships between phylogeography and biogeography for some species, but not for others, despite the similarities in their life histories and exposure to the same climatic changes.     

Comparative salinity tolerances of four siphonariid limpets in relation to habitat restriction of the rare and endangered Siphonaria compressa

The salinity tolerances of four South African species of limpet in the genus Siphonaria were experimentally tested, in the context that one of them, S. compressa, is South Africa's most endangered marine mollusc and is restricted to two lagoonal localities that normally experience little variation in salinity. Its habitat is additionally restricted in that it occurs only on the eelgrass Zostera capensis. The other three species, S. capensis, S. concinna and S. serrata, are abundant and widespread on temperate rocky shores. We hypothesised that S. compressa would have limited tolerance to low salinities, contributing to its absence from otherwise suitable estuaries that experience regular fluctuations of salinity. The percentage survival of the limpets was recorded in salinities ranging from 2 to 35 over that period. Both salinity and species significantly affected the survival of the limpets over the 96 hours. Survival was highest in salinities close to those of normal seawater, and greatly reduced in the hypo saline waters. S. compressa had the lowest salinity tolerance of the four species, its time-to-50% mortality in hyposaline waters being significantly less than that for the other three species, which differed in a manner that could be correlated with their zonation patterns. Restriction to eelgrass, coupled with low salinity tolerance, probably explains why S. compressa is found at only two localities, both of which normally experience salinities close to that of seawater. Even there, mass mortalities of S. compressa have been recorded in association with die-backs of eelgrass after abnormal freshwater flooding or sedimentation—further emphasising the vulnerability of S. compressa to extinction.     

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.     

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.     

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.     

Intertidal microalgal production and molluscan herbivory in relation to season and elevation on two rocky shores on the east coast of southern Africa

Microalgal production (ungrazed rock) and standing stock (grazed) were measured over a period of 15 months in the lower, middle and upper balanoid zones of two rocky snores in the Transkei region of southern Africa. The mean monthly production of chlorophyll a, which ranged from 0,3 to 2,0 μg·cm^?2. was inversely correlated with tidal elevation. Highest rates of production (3,5–6,0 μg·cm^?2) were recorded in winter at all tidal levels. The amplitude of seasonal fluctuations decreased with tidal height. At the middle and upper levels standing stock usually exceeded monthly production, whereas the opposite often occurred in the lower balanoid. No temporal or spatial trends in standing stock were evident. The intertidal distribution of grazing gastropods (excluding littorinids) correlated positively with primary production but showed no relationship with standing stock. No clear seasonal trends in grazer abundance were evident. The fact that standing stock exceeds monthly production suggests that grazing pressure is insufficient to balance primary production in the middle and upper balanoid zones on these shores. Grazers do, however, remove most of the primary production in the lower balanoid zone. Possible mechanisms underlying the observed patterns are discussed.     

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 Dynamics of an Introduced: Balanus glandula Population in the Southwestern Atlantic Rocky Shores. The Consequences on the Intertidal Community*

Abstract. This paper deals with the population dynamics of the introduced barnacle Balanus glandula in a port shelter and a rocky shore exposed of a southwestern Atlantic area. Replicated areas in both zones were denuded monthly and censused in both the high intertidal and the upper mid intertidal zone. The settlement pattern. recruitment, mortality and growth of B. glandula and a comparative successional pattern in both areas were recorded. Recruitment began in winter, i.e., July in the port and August in the rocky shore. In the port, green algae dominate up to a final community of Balanus glandula all along the intertidal, while in rocky shores the red encrusting alga Hildenbrandia lecanellieri establishes a cover that is replaced by the mytilid Brachidontes rodriguezi in the mid intertidal and by B. glandula in the high intertidal and upper mid intertidal. B. glandula densities were greater in the upper mid intertidal 19 600 and 5050 indiv. · m^-2 in the port and rocky shore, respectively than in the high intertidal 13600 and 1100 indiv.· m^-2, respectively. Mortality was greater in the port up to 62%, but heavy settlement allows a final density 4 times greater than in eulittoral rocky shores and 10 times greater than in the high intertidal. The winter reproduction of Balanus glandula, the absence of predators, the neutral or beneficial effect of algae. with spatial and temporal partitioning of substrate allows it to successfully outcompete mussels and other barnacle species from the intertidal community of both sheltered and exposed areas.     

Population genetic structure of the endangered limpet Cymbula nigra in a temperate Northern hemisphere region: influence of palaeoclimatic events?

Cymbula nigra (da Costa L, 1771) is the largest limpet in Europe and it is at present considered an endangered species. Nevertheless, its biology is poorly understood and there is a lack of knowledge about population genetic structure. A total of 61 C. nigra specimens were sampled and studied for a fragment of the mitochondrial gene cytochrome c oxidase subunit I. The low genetic diversity, the absence of geographic structure, the significant values reported by measures to test for a recent population expansion and a star‐shaped phylogeny pattern are here interpreted as forming a hypothesis concerning the influence of palaeoclimatic events in the genetic pool of limpets, considering also the information available on different limpet species.     

山东南部小场沙滩潮间带生态学研究

世界各海沿岸的潮间带大部分区域是沙滩，这主要是由于波浪的强烈冲击所致。与碎石、石砾滩一样，沙滩对底栖生物的生存显然不利，因而其栖息生物种类比较贫乏，数量也少，完全缺乏大型植物，除了某些特定的沙栖种外，主要是一些小型无脊椎动物和植物。河口附近和内湾水下平台则例外，丰富的有机物质使之成为完全不同的生境。 虽然潮间带生态研究已有一百多年的历史，但有关沙滩生境的研究较少，对沙滩生物的生态特点缺乏系统的探讨。有的学者直接采用岩岸潮间带生态研究方法，这不仅与客观实际不符，而且造成术语上的混乱。另外，在群落结构的研究中也存在不少问题，例如往往没有按生态特点区别底内生物和底表生物，造成对区系性质的错误判断。为此，我们想通过对小场沙滩生境的调查研究，对沙滩潮间带研究中的一些问题进行初步探讨。     

Abundance of biofilm on intertidal rocky shores: Can trampling by humans be a negative influence?

Trampling by human visitors to rocky shores is a known stressor on macroorganisms. However, the effects of trampling on rocky intertidal biofilm, a complex association of microorganisms of ecological importance in coastal communities, have not been quantified. We evaluated the impact of trampling frequency and intensity on total biomass of epilithic microalgae on intertidal rocky shores in the southeast of Brazil. There was a trend of increase in the variability of biomass of biofilm in function of intensity of trampling, but no significant effects emerged among trampling treatments. The low influence of trampling on biofilm might be a result of the small dimensions of the organisms coupled with their natural resilience and roughness of the substrate; the former preventing the removal of biofilm layers by shoes and facilitating their quick recovery. Our results provide insights for management and conservation of coastal ecosystems revealing a weaker impact of trampling on biofilm than that reported on macroorganisms.     

Contrasting genetic diversity with phenotypic diversity in coloration and size in Xantho poressa (Brachyura: Xanthidae), with new results on its ecology

The ecology of Xantho poressa (Olivi, 1792) (Brachyura) was studied during field trips to the Mediterranean Sea, the Black Sea and the Spanish Atlantic Ocean. Our results reveal that X. poressa lives from the intertidal to the shallow subtidal zone, and inhabits relatively protected rocky shores, often with pebble underground, from juvenile to adult stages. A mark–recapture experiment revealed a high population density in this habitat. All stages, but predominantly juveniles, show a variability of colour patterns, which allow the crabs to blend in with the rocky substratum, thereby hiding from predators as passive defence. Adulthood can be reached with a carapace length smaller than 6 mm. The morphometric analysis of the species revealed allometric growth in carapace shape. Variability in overall size could be observed at different collecting sites. Neither the colour morphs nor the size differences could be attributed to differences of Cytochrome Oxidase subunit I mitochondrial DNA sequences, suggesting that ecological rather than genetic patterns are responsible for the different phenotypes.     

Intertidal Invertebrate and Algal Communities on the Rocky Shores of the Bay of Morbihan, Kerguelen (South Indian Ocean)

Abstract. Both the emergent surfaces and tide pools of the rocky intertidal shores of the Bay of Morbihan, Kerguelen are depauperate in number of macroinvertebrate and macroalgal species. Diversity (H') was high and similar for macroinvertebrates in tide pools from the high to low intertidal and on the emergent surface exposed at extreme low-tide when calculated in terms of numbers of individuals. H' was lower when calculated in terms of energy and twice as high for the tide pool in the high intertidal than the lower tide-pools and the emergent surface. H' for macroalgae was 0 in the highest tide-pool and highest in the lower tide-pools. The density/species-rank curves for the macroinvertebrates and the macroalgae are logarithmic but vary in linearity. Algal biomass was 37 % of the macroinvertebrate biomass on the emergent surface at the extreme low-tide, but 106% in the highest tide-pool. Twenty-three months after denudation of the tide pools, the ratios of total densities of the macroinvertebrates (ind m^-2) to the original densities were 48 % for the pool in the high intertidal, 25 % for the middle tide-pool, and 4 % for the pool in the low intertidal. The diversity indexes for the macroinvertebrates were similar to the original ones. Recolonization at Kerguelen is slow and shows an intertidal gradient. The composition of the rocky intertidal community at Kerguelen results from the rigorous physical environment and the small number and functional types of species. In the absence of major predators on macroinvertebrates and of herbivores on macroalgae, interaction among the species present seems to be competitive in nature.     

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.