Site fidelity and homing behaviour in intertidal fishes

The theory of ecological cognition poses that the brains and behaviour of animals are shaped by the environmental challenges they face in their everyday lives. Site fidelity and homing ability was tested for five species of intertidal rock pool fish by tagging and displacing them to new rock pools at various distances from their ‘home’ rock pools. Three of the species were rock pool specialists whilst the remaining two spend a small proportion of their life in rock pools during early ontogeny. The three specialists showed strong site fidelity with >50 % of individuals found in the same pool 42 days after tagging. In contrast, the non-specialist species showed low fidelity and poor homing abilities. Homing success in the rock pool specialists remained relatively stable as displacement distance increased. The effect of body size on homing ability was species dependent, with only one species showing a significantly greater tendency to home with increasing size.     

Do artificial structures alter marine invertebrate genetic makeup?

Human-made structures are increasingly built in marine coastal habitats for a variety of purposes. Offshore oil and gas production platforms are among the largest examples. Yet, biological effects of these increasing density artificial substrata are under evaluated. The objective of our study is to investigate the possible role of offshore platforms in modifying the genetic composition of populations of natural rocky shores species. The serpulid Pomatoceros triqueter was used as a model, and genetic variation was assessed using a 419?bp fragment of the mtDNA COI gene in samples collected on eleven offshore gas platforms, on one coastal buoy on the sandy shore and in four sites located on natural rocky shores in the Adriatic Sea. Deep phylogenetic lineages were uncovered over all samples. Nucleotide diversity and mean number of pairwise differences among haplotypes were significantly smaller in offshore platform samples compared to rocky shores samples. No significant genetic structure was observed over all samples. We found direct evidence of lower genetic diversity on platforms confirming that, although artificial structures attract and support species typical of hard bottoms, they are not analogues of natural rocky habitats.     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

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

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

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

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

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

Ecology of the metahaline pool of Di Zahav,Gulf of Elat,with notes on the siphonocladacea and the typology of near-shore marine pools

The hydrobiology of a metahaline near-shore pool, at Di Zahav on the Sinai coast of the Red Sea, has been studied. Salinity of the pool fluctuates between 45 and 60, and winter temperatures are as low as 10°C. The shore consists partly of beach rock, some of it still in statu nascendi. Serpulid reef-boulders also occur, covered with typical marine species. Three siphonocladacean green algae (Siphonocladus rigidus, Dictyosphaeria cavernosa, and Valonia utricularis), form abundant crusts on the boulders. Several peculiar faunal elements are discussed. Traits in common with the Suez Canal biota are discerned. A typology for the nearshore pools of Sinai is proposed.     

Synergistic effects of ultraviolet radiation and conditions at low tide on egg masses of limpets (<Emphasis Type="Italic">Benhamina obliquata</Emphasis> and <Emphasis Type="Italic">Siphonaria australis</Emphasis>) in New Zealand

The effects of ultraviolet radiation (UVR), desiccation and conditions in tidal pools on embryonic survival were examined for two common pulmonate limpets that lay intertidal benthic egg masses on rocky shores in New Zealand: Benhamina obliquata and Siphonaria australis. Field surveys and manipulative experiments were conducted between December 2006 and September 2007 in the Wellington region of New Zealand (41°17′S, 174°47′E). Egg mass deposition sites in the field were species-specific: B. obliquata deposited eggs primarily in shaded crevices, whereas S. australis predominantly deposited egg masses in the sun and in tidal pools. For both species, however, embryonic mortality was greater in egg masses that had been in full sun compared to shade. For S. australis, there was also high mortality in egg masses in tidal pools or desiccated compared to those that remained submerged in flowing seawater at low tide. In outdoor experiments, embryonic mortality was also always greatest for egg masses exposed to full sun, and lowest for those in shaded treatments. Mortality was also higher if egg masses were in simulated tidal pools, and for S. australis, if desiccated, compared to those submerged in flowing seawater. Periods of particularly sunny conditions with high temperatures also resulted in higher overall mortality. Finally, egg masses of both species that were initially deposited in the shade had greater mortality in response to subsequent UV exposure compared to egg masses initially deposited in full sun. Results from this study suggest that the egg masses of these two species are highly vulnerable to UVR, as well as other intertidal stressors. Embryos of both of these species may be at risk of high mortality particularly during summer when extreme conditions of UV intensity and high temperature coincide with low tide cycles. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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

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

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

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

Spatial and seasonal variations in bacterial communities of the Yellow Sea by T-RFLP analysis

Four typical coastal sites (rocky shore, sandy shore, mud flat shore, and artificial harbor) at the Yellow Sea were chosen to investigate the spatial and seasonal variations in bacterial communities. This was accomplished by using terminal restriction fragment length polymorphism (T-RFLP) analysis of PCR amplified 16S rDNA fragments. Two kinds of tetrameric restriction enzymes, HhaI and MspI, were used in the experiment to depict the bacterial community diversity in different marine environments. It was found that the community compositions digested by the two enzymes separately were different. However, the results of bacterial community diversity derived from them were similar. The MDA analysis results of T-RFLP profiles coming from HhaI and MspI both exhibited a significant seasonal community shift for bacteria and a relatively low spatial variation among the four locations. With HhaI as the sample, the pair wise T-tests also revealed that variations were minor between each pair of marine environments, with R ranging from 0.198 to 0.349. However, the bacterial community structure in the mud flat site depicted a larger difference than each of the other three sites (R ranging from 0.282 to 0.349).     

Zonation of the interstitial polychaeteStygocapitella subterranea (Stygocapitellidae) in European sandy beaches

The zonation ofStygocapitella subterranea Knoellner (Polychaeta, Stygocapitellidae) in sandy beaches is compared for several localities along the European coasts. In tidal beaches the species occupies a zone near average high tide level (AHTL), whereas in non-tidal beaches it is found at a fairly long distance from the water line. The regularities observed inS. subterranea indicate that the interstitial fauna may prove well suited for the establishment of a general scheme of zonation in sandy beaches analogous to that for rocky shores proposed byStephenson andStephenson (1949).     

Comparative species composition and relative abundance of decapod crustaceans in marine habitats of Panamá

The community structure of the decapod crustacean fauna of 7 tropical, shallowwater, marine habitats (sandy beaches, mangrove swamps and rocky intertidal habitats on both the Pacific and Caribbean coasts of Panama, and Pocillopora damicornis coral habitat of the Bay of Panama) were examined and analyzed for species composition and relative abundances. Collections from the 7 habitats yielded 4361 individuals, representing 236 species. The number of species per habitat was (Pacific, Caribbean): sandy beach (16, 7); mangrove (20, 17); P. damicornis (53); rocky intertidal (78, 67). There were more species represented by more individuals in the Pacific habitats. An index of faunal similarity was calculated for each pair (Pacific-Caribbean) of habitats. This index is the number of ecologically similar congeneric species which occurred in both habitats expressed as a percentage of the total number of species present in the pair of habitats. For the sandy beach communities there were three Pacific species which were similar to three Caribbean species, a similarity of 6/23 or 26%. The index of similarity for the mangrove communities is 54% and for the rocky intertidal communities it is 37%. The P. damicornis community has affinities with the Pacific rocky intertidal community (18%), with that of the Caribbean rocky intertidal (16%) and with that of Indo-West Pacific pocilloporid corals (20%). A few specialized species dominated each of the communities. The habitats and the number of species accounting for over half of the individuals present are (Pacific, Caribbean): sandy beach (1, 1); mangrove (4, 4); P. damicornis (5); and rocky intertidal (3, 6). Most of the species in each community were represented by one or a few individuals.     

Role of peat organic matter on isotopic composition of most abundant benthic organisms in intertidal habitats of SW Iceland

The transfer of peat organic matter (OM) from tidal pools of upper littoral to downstream rocky shores and its potential incorporation into marine biota were investigated using the stable isotope approach. Samplings were carried out in September 2004 in two SW Icelandic sites (Osar and Hvassahraun), where we selected (1) areas with shores where grass gently declined towards rocks and, on reaching the sea line, formed small tidal peat pools and (2) areas where grass and rocky shores were not contiguous, rather the grass lower limits were several hundreds of meters back from the shore. In both grass and no grass areas, in the intertidal zone, all benthic organisms and all potential OM sources were sampled. Dominant macroalgae were Ascophyllum nodosum ([Linnaeus] Le Jolis) and Fucus vesiculosus Linnaeus. In grass sites, organisms partially reflected the isotopic composition of peat OM, while in no grass sites, peat represented only a negligible amount of available OM, or its signal was totally absent from the environment. Intertidal organisms could rely on peat both directly by suspension feeding, grazing or scavenging and indirectly by predation (crabs) on low level consumers (barnacles).     

Floristic composition of sandy shore vegetation at Uswetakeiya in Sri Lanka

This study was conducted at Uswetakeiyawa on the southwestern part of Sri Lanka. Twenty-two families, 42 genera and 52 species were recorded from the sandy shore vegetation. Approximately 3.8 % (2 species) were found to be endemic and 67.3 % (35 species) were found to be of medicinal importance. Ipomea pes-capre and Spinifex littoreus were the most abundant species at the study site. The average value for all the zone combinations was β = 0.693, and the highest similarity value was ISj = 0.276, corresponding to two non-contiguous zones. The results show that a variation in plant diversity is to be found across the gradient. The typical zoning nature of the sandy shore vegetation has been disturbed due to on-going development in Uswetakeiyawa. New policy decisions should be taken to restrict development activities in Uswetakeiyawa in order to protect the sandy shore vegetation.     

Seasonal variation in utilization of biogenic microhabitats by littorinid snails on tropical rocky shores

Mobile species may actively seek refuge from stressful conditions in biogenic habitats on rocky shores. In Hong Kong, the upper intertidal zone is extremely stressful, especially in summer when organisms are emersed for long periods in hot desiccating conditions. As a result, many species migrate downshore between winter and summer to reduce these stressful conditions. The littorinids Echinolittorina malaccana and E. vidua, for example, are found on open rock surfaces high on the shore in winter but the majority migrate downshore in summer to the same tidal height as a common barnacle, Tetraclita japonica. In the laboratory, where environmental conditions could be controlled to approximate those occurring on the shore, we tested whether the downshore migration allowed littorinids to select barnacles as biogenic habitats to reduce stress and if this behaviour varied between seasons. In summer, littorinids demonstrated a strong active preference for the barnacles, which was not observed in the cool winter conditions, when animals were found on open rock surfaces even when barnacles were present. Littorinids, therefore, only actively select biogenic habitats during the summer in Hong Kong when they migrate downshore, suggesting that such habitats may play an important, temporal, role in mitigating environmental stress on tropical shores.     

Monitoring spatio-temporal variation in molluscan grazing pressure in seasonal, tropical rock pools

In Hong Kong, low-shore rock pools support a patchy mosaic of different successional stages of crustose coralline algae. Echinoid and molluscan grazers are abundant in these low-shore rock pools and spatial and temporal variation in grazing pressure is thought to play an important role in structuring these assemblages. Direct records of graze marks using a modified wax-disc method were used to monitor spatial (different pools) and temporal (periods, 3 or 4 days each; spring vs. neap tides; summer vs. winter) variations in grazing pressure. Radula marks of Chlorostoma argyrostoma, Lunella coronata and Nerita albicilla were common in all pools. Grazing pressure was highly variable with tidal state and season but also within these two factors. Lunella and Nerita exerted highest grazing pressure at neap tides during summer, whilst Chlorostoma exerted highest grazing pressure in winter. Although overall patterns of temporal variation were similar and consistent in all pools, there was between pool variation in grazing pressure and individual species showed different patterns. Within the seasonal and tidal variations, large period-to-period differences were found for all species, illustrating small-scale temporal variation. Fine-scale spatial and temporal patterns of variation in grazing pressure of different species, therefore, contribute to the highly dynamic nature of these low-shore rock pools.     

The importance of sandflat morphology to recruitment of the intertidal snail Nassarius pauperatus during ten consecutive years at three sites in South Australia

This project was desinged to confirm the temporal consistency of inter-site differences in recruitment of an intertidal gastropod, and to test the hypothesis that these differences are accounted for by differences in shore topography. Annual recruitment of the intertidal sanil Nassarius pauperatus (Lamarck) was estimated at two sandflats in Gulf St Vincent, South Australia, over a period of 12 yr (1979 to 1990). At one sandflat the gradient was slight, the zone containing N. pauperatus was wide, and there were extensive pools of water at low tide, whilst at the other sandflat the gradient was steeper, the zone containing N. pauperatus was narrower, and there were fewer or no pools. Although recruitment at both sites varied temporally, the density of recruits was always higher at the first site. These data, together with data covering 10 yr (1981–1990) from a sandflat where the gradient of the substratum was decreasing due to accretion of sediment, suggest that shore topography is an important determinant of relative recruitment for N. pauperatus in South Australia.     

Chaotic genetic patchiness in an intertidal limpet,Siphonaria sp.

Variation of 4 polymorphic enzymes was studied for 2 yr (1978 and 1979) in an undescribed species of Siphonaria, a pulmonate limpet, from a rocky shore at Rottnest Island, Western Australia. Depending on the locus, significant genetic differences were found among sites along 50 m of shore, between high and low portions of the shore within sites, between adults and recruits, and between recruits in the 2 yr. This genetic heterogeneity does not follow a simple, consistent pattern, but forms a shifting, ephemeral genetic patchiness best described as chaotic. This patchiness may result from temporal variation of numbers and genotypes of recruits, which leads to the proposal that planktonic dispersal, although causing uniformity on a large scale, can give rise to fine-scale genetic patchiness.     

The contribution of seagrass beds (<Emphasis Type="Italic">Zostera noltii</Emphasis>) to the function of tidal flats as a juvenile habitat for dominant,mobile epibenthos in the Wadden Sea

Since the substantial loss of subtidal eelgrass (Zostera marina L.) in the 1930s, seagrass beds in the Wadden Sea are limited to the intertidal zone and dominated by Z. noltii Hornem. This study deals with the effect of vegetated tidal flats on quantities of mobile epifauna and proves empirically the function of seagrass canopies as a refuge for marine animals remaining in the intertidal zone at ebb tide. Drop-trap samples were taken in the Sylt-Rømø Bight, a shallow tidal basin in the northern Wadden Sea, on vegetated and unvegetated tidal flats during July and August 2002, and during the entire growth period of Z. noltii from May to September in 2003. The species composition in Z. noltii and bare sand flats showed minor differences since only two isopod species (Idotea baltica and I. chelipes) occurred on Z. noltii flats exclusively. Juvenile shore crabs (Carcinus maenas L.), brown shrimps (Crangon crangon L.) and common gobies (Pomatoschistus microps Krøyer) were also found abundantly on bare sand flats. However, the results showed significantly higher abundances and production of these dominant species on vegetated tidal flats. Additionally, the analyses of faunal size classes indicated higher percentages of small individuals in the seagrass bed during the entire sampling period. Despite drastic diurnal fluctuations of dissolved oxygen at low tide, faunal density in the residual water layer remaining in seagrass canopies at ebb tide was found to be consistently higher than that found in artificially created tide-pool units. Although species composition of mobile epifauna did not basically differ between vegetated and unvegetated tidal flats, Z. noltii beds are considered to contribute quantitatively to the function of tidal flats, as an extended juvenile habitat for some of the most important species of the Wadden Sea food web.Communicated by O. Kinne, Oldendorf/Luhe