Introduction,establishment and expansion of the Pacific oyster <Emphasis Type="Italic">Crassostrea gigas</Emphasis> in the Oosterschelde (SW Netherlands)

The Pacific oyster Crassostrea gigas was first introduced as an exotic species by oyster farmers in 1964 in the Oosterschelde estuary (SW Netherlands). The initial phase is not well documented but first natural spatfall was recorded in 1975. Excessive spatfall occurred in 1976 and this is considered the start of the expansion phase of the wild oysters. Oyster beds in intertidal and subtidal areas of the Oosterschelde estuary have been growing since. The development in the intertidal area has been reconstructed by using aerial photography, validated by ground truth in 2000–2002. In the subtidal areas extensive oyster beds have been detected by using side scan sonar; on hard substrates along the dikes coverage with oysters up to 90% locally has been recorded by scuba diving surveys. Expansion has also occurred into adjacent water bodies including the Wadden Sea. By forming resistant reefs the oysters induce structural changes in the ecosystem. It is concluded that bed area is still expanding while decrease of the fraction live animals may indicate adjustment of the stock size to the local conditions.     

Physical constraints on the foraging ecology of a predatory snail

We studied the effects of aerial exposure and high summer temperatures on the southern oyster drill ( Stramonita haemastoma ), feeding on the American oyster, Crassostrea virginica . In the laboratory, oyster drill feeding rates and growth were highest at 25 and 30°C, some mortality occurred at 35°C, all snails died at 40 and 45°C, and the 28-day LC 50 was 35.7°C. In a second experiment where both water temperature (25 vs . 33°C) and aerial exposure were varied, only simulated tidal exposure lowered oyster drill feeding and growth rates. In field cage experiments, oyster drills had reduced feeding rates and growth at intertidal sites, but snail growth rates increased in late summer with warmer water temperature. We therefore conclude that aerial exposure, not high temperature, is the major factor limiting oyster drill feeding and growth in intertidal oyster reefs. Field experiments with partial cages also suggested that ambient predation rates were much higher at a subtidal than at a nearby intertidal site. Because southern oyster drills have depressed feeding, growth, and possibly lower fitness in intertidal oyster reefs during the summer, this reduced predation risk may provide a refuge for intertidal oysters.     

Subtidal populations of the blue mussel <Emphasis Type="Italic">Mytilus edulis</Emphasis> as key determinants of waterfowl flocks in the southeastern Barents Sea

Ornithological surveys conducted over the Pechora Sea (the southeastern part of the Barents Sea) in the 1990 s revealed huge non-nesting flocks of marine ducks, the largest in the European North. Especially dense waterfowl aggregations are constantly observed at the shallows near Dolgij Island during molting period and migration to wintering places. All the marine ducks flocking there are specialized benthos feeders predominantly consuming mussels Mytilus edulis. At the same time, numerous previous benthic studies in the Pechora Sea did not reveal mussels near Dolgij Island where benthic biomass was somewhat lower than in the adjacent areas (Denisenko in Mar Ecol Prog Ser 258:109–123. 2003) which left the food source for these abundant bird flocks enigmatic. In the course of an expedition in summer 2007 we found subtidal populations of M. edulis in shallows to the southwest of Dolgij Island. These populations were confined to a coastal zone and were characterized by a highly disjunct distribution with the biomass reaching up to 4 kg m⁻². We describe these subtidal populations as well as an intertidal mussel population on the western shore of Dolgij Island.     

An Experimental Evaluation of Different Methods of Restoring Phyllospadix torreyi (Surfgrass)

The surfgrass Phyllospadix torreyi is an abundant seagrass found on rocky exposed shores of the Pacific coast of North America. In southern California surfgrass populations are adversely affected by a range of natural events and anthropogenic activities. Few attempts have been made to develop restoration methods for surfgrass, and none have investigated the efficacy of using different life stages. We evaluated several techniques for restoration in intertidal and subtidal habitats using: (1) laboratory‐reared seedlings transplanted to the field (2) sprigs (short lengths of rhizome containing a few shoots) transplanted from undisturbed populations, and (3) plugs (a cohesive clump of shoots and rhizomes) transplanted from undisturbed populations. We calculated the net change in the aerial coverage of surfgrass after 6 months, taking into account the recovery or additional losses from the donor population, and amount of effort involved in transplanting. Transplanted seedlings survived poorly and had minimal rhizome growth at both the intertidal and the subtidal sites, yet the individuals that did survive showed a 275% increase in leaf number. Survivorship of transplanted plugs was high in both habitats; however, physical disturbances to the donor populations exacerbated damage sustained at the time of collecting, yielding a substantial net loss in surfgrass. Sprigs transplanted to the subtidal had higher survivorship (71 versus 48%) and a greater increase in the aerial coverage of rhizome (86 versus 42%) than those transplanted to the intertidal. Of the three techniques, transplanted sprigs had the greatest overall increase in aerial coverage per unit effort, suggesting that this method may be the most effective approach for restoring P. torreyi.     

The potential for seaweed resource development in subarctic Canada; Nunavik,Ungava Bay

Ungava Bay is ice covered 6–7 months of the year and evidence of ice scouring of seaweeds is extensive in the intertidal and shallow subtidal. Maximum tidal amplitudes of 16 m, among the highest in Canadian waters, compound this impact. Despite this level of annual perturbation, very extensive and dense beds of fucoids in the intertidal and laminarians in the subtidal are common on the western shores of Ungava Bay. Ground surveys of 24 intertidal stations combined with satellite images delineated 82,000 tons standing crop of Fucus vesiculosus and Fucus evanescens in Payne Bay, of which 36,000 tons were considered harvestable. Subtidally, kelp cover reached peak biomass at 5–10 m consisting of three primary species, Saccharina longicruris, Laminaria digitata, and Laminaria solidungula. In the area of Payne Bay, kelp beds of 100 ha were common, averaging 9–12 kg m⁻² wet weight. The productivity of brown algae at these latitudes has been assumed to be low relative to southern latitudes. Direct measurement of lineal growth indicates productivity is intermediate between arctic and temperate populations. The potential for medium level industrial harvest exists under conservative management strategies within the constraints of subarctic logistics.     

Tidal activity pattern and feeding behaviour of the ophiuroid Ophiocoma scolopendrina on a Kenyan reef flat

Ophiocoma scolopendrina exhibits a distinctive pattern of feeding activity on intertidal reef platforms off Kenya. With the first wave of the flooding tide, dense aggregations of these ophiuroids (up to 320 m⁻²) engage in a 1–2 min burst of surface-film feeding, vigorously sweeping the air-water interface and associated sea foam with the ventral surface of 2–4 arms. Suspension feeding (with arms extended in the water column) is the primary feeding mode throughout the rest of the tidal cycle (involving 25–65% of the population at a time), while bottom feeding (with arms extended along the substratum) is infrequent (<10%). Field experiments showed that surface-film feeding is regulated by water depth and can be triggered by suspended particles. This feeding mode appears to be an adaptation to the intertidal habitat, enabling the ophiuroids to exploit a nutrient-rich surface film during a temporal refuge (low tide) from fish predation. Dense populations of O. scolopendrina may represent an important trophic link between producers of particulate organic material and higher-level consumers in coral reef environments.Tara Oak and Robert E. Scheibling contributed equally to this paper. The order of authorship is alphabetical     

Intertidal and subtidal soft-bottom macro- and meiofauna of the Kongsfjord (Spitsbergen)

The intertidal and subtidal soft bottom macro- and meiofauna of a glacier fjord on Spitsbergen was studied after complete ice melt in June 2003. The abundances of the benthic fauna were within the range reported from estuaries and similar intertidal areas of boreal regions. The high proportion of juveniles in the eulittoral zone indicated larval recruitment from subtidal areas. The macrobenthic fauna can be divided into an intertidal and a subtidal community, both being numerically dominated by annelids. Deposit feeders were numerically predominant in intertidal sites, whereas suspension feeders were most abundant in the subtidal area. Among the meiofauna, only the benthic copepods were identified to species, revealing ecological adaptations typical for intertidal species elsewhere.     

Control of biofouling on pearl oysters Pinctada imbricata using wax and Chinese herbs

Abstract

Biofouling poses severe challenges to pearl oyster Pinctada imbricata culture in China, and controlling it is both labor- and capital-intensive. The antifouling properties of wax, and wax mixed with Chinese herbs, sprayed onto pearl oyster shell surfaces during peak biofouling seasons were evaluated. Pearl oysters coated with three wax treatments (plain wax, Chinaberry seed extract, Chinese honeylocust fruit extract) and a control (no treatment), were cultured in nets for up to 60?days. Mortality rate, fouling organism and pearl-oyster weights, and shell height are reported for individual oysters on each of six sampling dates. With the exception of oysters submerged for 12?days, all oysters were significantly affected by treatment type and submersion duration. Fouling weight increased more rapidly over time in the control-treatment oysters. Wax-based coatings deterred fouling-organism settlement on oysters for at least 2?months during the intensive fouling season, reducing mortality and not adversely effecting growth.     

Habitat linkages: the effect of intertidal saltmarshes and adjacent subtidal habitats on abundance, movement, and growth of an estuarine fish

In this study we used pinfish (Lagodon rhomboides) in field experiments to examine linkages between intertidal saltmarsh and adjacent subtidal habitats. Pinfish are more than twice as abundant in intertidal marshes adjacent to seagrass beds than in those adjacent to the unvegetated subtidal bottom. Movement of pinfish between the marsh edge and the adjacent subtidal habitat was greater for fish captured in areas with both intertidal and subtidal vegetation than in those with intertidal vegetation and adjacent unvegetated mudflats. This movement provides an important link between habitats, allowing transfer of marsh-derived secondary production to subtidal seagrass beds and vice versa. Pinfish held in enclosures with both intertidal and subtidal vegetation were, on average, approximately 90% heavier than fish held in enclosures with intertidal vegetation and unvegetated subtidal bottom. Because saltmarshes and seagrass beds contribute to the production of living marine resources, active measures are being taken to preserve and restore these habitats. The results from this study have direct application to decisions concerning site selection and optimal spatial proximity of saltmarsh and seagrass habitats in the planning of restoration and mitigation projects. To maximize secondary production and utilization of intertidal marshes, managers may opt to restore and/or preserve marshes adjacent to subtidal seagrass beds. Received: 31 May 1996 / Accepted: 23 October 1996     

Absence of surface-associated microorganisms in adult oysters (Crassostrea gigas).

Healthy, actively feeding intertidal oysters were removed from an estuarine environment (Pipeclay Lagoon, Tasmania). The epithelial surfaces of various organs of the mantle cavity and alimentary tract were explored by scanning and transmission electron microscopy. All epithelial tissues examined were ciliated, and nearly all were partly covered with secreted mucus. However, microorganisms were seen rarely in the adhesive mucus and never attached to the epithelium. Electron microscopy also failed to demonstrate a surface microflora in emersed oysters which had been incubated at 5 to 25 degrees C for 6 or 24 h. The absence of an internal surface microflora did not vary on a seasonal basis. In laboratory experiments, oysters were allowed to filter feed from seawater containing diverse types of marine bacteria at concentrations of 10(3) to 10(7)/mL. However, no surface microflora could be found within actively feeding oysters or in emersed animals incubated at 20 degrees C for 6 or 24 h. In contrast, surface-associated microorganisms were detected readily by scanning electron microscopy on the external shell of healthy oysters and on various internal tissues in spoiled oysters. It is suggested that the major mechanisms restricting microbial growth within oysters are ciliary movement and mucus secretion.     

Modern marine stromatolites in the Exuma Cays,Bahamas: Uncommonly common

Summary Modern stromatolites in open marine environments, unknown until recently, are common throughout the Exuma Cays, Bahamas. They occur in three distinct settings: subtidal tidal passes, subtidal sandy embayments and intertidal beaches. These stromatolites have a relief of up to 2.5 m and occur in water depths ranging from intertidal to 10 m. Surfaces near the sediment-water interface are typically colonized by cyanobacterial mats, whereas high relief surfaces are commonly colonized by algal turf and other macroalgae such asBatophora, Acetabularia, andSargassum. The internal structure of the stromatolites is characterized by millimeter-scale lamination defined by differential lithification of agglutinated sediment. In thin section, the lithified laminae appear as micritic horizons with distinct microstructures: they consist of thin micritic crusts (20–40 μm thick) overlying layers of micritized sediment grains (200–1000 μm thick); the micritized grains are cemented at point-contacts and are trucated along a surface of intense microboring. The Exuma stromatolites are built by cyanobacterial-dominated communities. These laminated prokaryotic structures grade to unlayered thrombolites built by eukaryotic algae. The variety of sites, settings and shapes of stromatolites in the Exuma Cays present excellent opportunities for future studies of stromatolite morphogenesis.     

Selective choice of sucrose solution concentration by the hovering hawk mothMacroglossum stellatarum

To determine the preference of the hovering hawk mothMacroglossum stellatarum for different sugar concentrations, the foraging behavior of adults were analyzed under laboratory conditions. Six sucrose concentrations (range, 10–60%, w/w) were simultaneously offered in six artificial ab libitum feeders. The number of feeding bouts and the duration of each visit were automatically recorded and stored in a computer. Results showed that the frequency of visits to the feeders did not vary among the different solutions offered, but the gathered volume by the group attained a maximum at between 20 and 50% (w/w). Moths invested more time in front of the feeder with the more concentrated sugar solutions. It was assumed that factors different from maximizing energy intake, such as water balance and viscosity of concentrated nectars, have to be considered in order to understand the observed patterns of nectar choice.     

Intertidal and Subtidal Fouling Assemblages in a Patagonian Harbour (Argentina, Southwest Atlantic)

The composition of the early stages of intertidal and subtidal fouling assemblages in Comodoro Rivadavia harbour (Argentina, 45°52′ S, 67°28′ W) and the influence of shore level and season on their structure were analysed. At the beginning of each season, stones were glued to the substratum with epoxy putty and distributed along 4 vertical transects at intervals of 20 m, at 3 levels: upper intertidal, middle intertidal, and subtidal. Substrata remained in the field for 84–100 days. A total of 48 samples (4 seasons × 3 levels × 4 replicates) were analysed. Species richness increased with depth, with 6 taxa in the upper intertidal, 23 in the middle intertidal and 31 in the subtidal. Seasonal differences in richness were less distinct. Green, red and brown algae were the dominant groups. Invertebrates were mainly represented by filter-feeding, sessile organisms, such as cheilostome bryozoans, spirorbid polychaetes and acorn barnacles. The barnacle Balanus glandula and the bryozoan Cryptosula pallasiana were the only non-indigenous species found in this study. Highly significant differences in structure among shore levels and seasons were evidenced by a two-way ANOSIM test. The upper intertidal is characterized by the filamentous green algae Urospora penicilliformis and Ulothrix flacca. The barnacle Balanus glandula is the most abundant species in the middle intertidal. The subtidal is defined mainly by the presence of the spirorbid polychaetes Paralaeospira levinseni and Romanchella perrieri, and the keyhole limpet Fissurella radiosa. Ordination of samples by season was less clear than by shore level.     

Flexibility of Physiological Traits Underlying Inter-Individual Growth Differences in Intertidal and Subtidal Mussels Mytilusgalloprovincialis

Mussel seed (Mytilusgalloprovincialis) gathered from the intertidal and subtidal environments of a Galician embayment (NW, Spain) were maintained in the laboratory during five months to select fast (F) and slow (S) growing mussels. The physiological basis underlying inter-individual growth variations were compared for F and S mussels from both origins. Fast growing seemed to be a consequence of greater energy intake (20% higher clearance and ingestion rate) and higher food absorption rate coupled with low metabolic costs. The enhanced energy absorption (around 65% higher) resulted in 3 times higher Scope for Growth in F mussels (20.5±4.9 J h⁻¹) than S individuals (7.3±1.1 J h⁻¹). The higher clearance rate of F mussels appears to be linked with larger gill filtration surface compared to S mussels. Intertidal mussels showed higher food acquisition and absorption per mg of organic weight (i.e. mass-specific standardization) than subtidal mussels under the optimal feeding conditions of the laboratory. However, the enhanced feeding and digestive rates were not enough to compensate for the initial differences in tissue weight between mussels of similar shell length collected from the intertidal and subtidal environments. At the end of the experiment, subtidal individuals had higher gill efficiency, which probably lead to higher total feeding and absorption rates relative to intertidal individuals.     

Size and Performance of Juvenile Marine Invertebrates: Potential Contrasts Between Intertidal and Subtidal Benthic Habitats

Newly settled or hatched juveniles of marine benthic invertebratesgenerally experience very high mortality. Juvenile mortalitycan profoundly affect adult populations, but little is knownabout how individual variation in juvenile quality affects performance.Several recent studies have demonstrated that differences insize, larval nutrient stores, or larval feeding history canstrongly affect the performance (measured as growth and survivorship)of juveniles. Additional research suggests that the strengthof the effect of juvenile size on performance may be mediatedby variation in environmental stress in the intertidal, a habitatcharacterized by strong fluctuations in abiotic factors. Themajor sources of juvenile snail mortality are likely to differin intertidal and subtidal habitats; abiotic stresses relatedto exposure, such as desiccation, are important in the intertidalbut far less severe in subtidal environments. Previously observedtrends in hatching or settlement size between intertidal andsubtidal species from three gastropod taxa may be due to differingselective regimes acting on initial juvenile size.     

Relationship between algae and environment: an Early Cretaceous case study, Trascău Mountains, Romania

The relationship between algae and depositional environment was studied in a limestone succession of Berriasian–Valanginian age. Several depositional environments were recorded from shallow subtidal to intertidal and supratidal, with salinity ranging from normal marine to fresh and/or supersaline water. The algal assemblages consist mainly of dasycladaleans, rivulariacean-type cyanobacteria and charophytes. Nipponophycus (Bryopsydales) and Lithocodium-Bacinella (microproblematicum), occur at some levels. Dasycladaleans are associated with subtidal, sometimes restricted (“lagoonal”) environments, while rivulariacean-like cyanobacteria and charophytes characterise intertidal-supratidal and fresh and/or supersaline environments, respectively. Among the dasycladaleans, Salpingoporella annulata is often related to restricted environments where it forms monospecific assemblages. Large dasycladaleans, such as Selliporella neocomiensis, Macroporella praturloni and Pseudocymopolia jurassica are found in subtidal high-energy deposits (bioclastic grainstones). The relationship between environment and algae, characteristic for each depositional unit, can be used to interpret the relative sea-level variations.     

Introduced Pacific oysters (Crassostrea gigas) in the northern Wadden Sea: invasion accelerated by warm summers?

Among the increasing number of species introduced to coastal regions by man, only a few are able to establish themselves and spread in their new environments. We will show that the Pacific oyster (Crassostrea gigas) took 17 years before a large population of several million oysters became established on natural mussel beds in the vicinity of an oyster farm near the island of Sylt (northern Wadden Sea, eastern North Sea). The first oyster, which had dispersed as a larva and settled on a mussel bed, was discovered 5 years after oyster farming had commenced. Data on abundance and size-frequency distribution of oysters on intertidal mussel beds around the island indicate that recruitment was patchy and occurred only in 6 out of 18 years. Significant proportions of these cohorts survived for at least 5 years. The population slowly expanded its range from intertidal to subtidal locations as well as from Sylt north- and southwards along the coastline. Abundances of more than 300 oysters m^–2 on mussel beds were observed in 2003, only after two consecutive spatfalls in 2001 and 2002. Analyses of mean monthly water temperatures indicate that recruitment coincided with above-average temperatures in July and August when spawning and planktonic dispersal occurs. We conclude that the further invasion of C. gigas in the northern Wadden Sea will depend on high late-summer water temperatures.Communicated by H.D. Franke     

Attack responses of the southern Australian whelk, Lepsiella vinosa (Lamarck, 1822) (Gastropoda: Muricidae), to novel bivalve prey: an experimental approach

In their native ranges, muricid gastropods feed on similar prey, often bivalves and barnacles, which they usually drill. Throughout its wide southern Australian distribution, the intertidal Lepsiella vinosa feeds on a range of prey from barnacles and littorinid gastropods in the southeast to mussels in the southwest. A number of muricids have been introduced throughout the world, either with oysters or in ship ballast water. In their new environments, they switch to native prey but their feeding responses to them have never been studied in the laboratory. The object of this study was to study the feeding of L. vinosa on a suite of non-native species. Australian Lepsiella vinosa was taken to Hong Kong, offered five different possible prey species and allowed to feed to satiation for many weeks. Replacement of consumed prey items by similar-sized conspecifics was undertaken until trends emerged. Lepsiella vinosa readily attacked thick- and thin-shelled bivalves from Hong Kong’s sandy beaches, Anomalocardia squamosa and Caecella chinensis, respectively, and from rocky shores, Septifer virgatus and Hormomya mutabilis, again respectively. It attacked them all, as it does its major prey item, Xenostrobus inconstans, in its native southwestern Australia, by either drilling or marginal probing with its proboscis. It also preferred intermediate sized prey (10–15 mm shell length), as with its natural prey. It quickly attacked the sandy shore species, and Hormomya mutabilis and Mytilopsis sallei, the latter two being closely similar in shell form and size to its natural prey X. inconstans. Hormomya mutabilis was the most favored prey, and was most similar in shell form and thickness to X. inconstans. This study therefore suggests that if introduced elsewhere, L. vinosa could radically affect intertidal community structure.     

Tolerance of forced air emergence by a fish with a broad vertical distribution, the rockpool blenny, Hypsoblennius gilberti (Blenniidae)

Many intertidal fishes, particularly among the Blenniidae and Cottidae, possess amphibious adaptations, including the ability to breathe in air and to avoid desiccation in terrestrial conditions. These traits are absent in subtidal species of blennies and cottids. Hypsoblennius gilberti, the rockpool blenny, is found in shallow rockpools in the mid to high intertidal areas of Southern California, and deeper to 18 m in the subtidal zone. This broad vertical distribution could indicate that this blenny is adapted for tidal air emergence, although H. gilberti has not been observed out of water in its natural habitat. H. gilberti does not emerge voluntarily from hypoxic sea water in the laboratory, but it easily withstands 3 h out of water. The aerial respiratory exchange ratio (CO2 released compared to O2 consumed) is 0.70, similar to that of amphibious intertidal fishes in air, indicating sufficient release of metabolically produced CO2 while emerged. There is no increase in aquatic respiration following emergence. However, unlike other amphibious fishes that maintain aerial oxygen consumption at a level similar to aquatic oxygen consumption, H. gilberti has an aerial oxygen consumption rate one-third that in water. H. gilberti can recover rapidly from terrestrial water loss, and shows no change in evaporative water loss rates at 93% and 77% relative humidities. The amphibious capabilities in H. gilberti, even if rarely used, permit survival in air during tidal emergence. These findings suggest that H. gilberti may demonstrate an intermediate condition between the amphibious species of intertidal fishes that regularly emerge from water, and the subtidal fishes that do not survive air emergence and are completely restricted to an aquatic habitat.     

A comparative study of territoriality in intertidal and subtidal blennioids (Teleostei, Blennioidei)

This paper presents a comparative analysis of territoriality in three intertidal (Lipophrys pholis, Coryphoblennius galerita, Salaria pavo) and two subtidal (Tripterygion delaisi, Parablennius pilicornis) blennioid fishes. Focal-animal observations of males guarding eggs showed that: (i) intertidal species had smaller territories that were less frequently patrolled; (ii) in subtidal species feeding was limited to the territory, while in intertidal fishes a substantial proportion of the feeding acts occurred outside the defended area; (iii) intertidal species spent less time out of the nest and showed a lower level of locomotory activity; (iv) subtidal species were subjected to a higher number of territorial intrusions with more species intruding their territory, some of them potential egg predators; (v) subtidal species performed several water column displays that played a significant role in courtship, while in intertidal fishes these displays are almost absent. Signalling without the presence of a female was almost absent in intertidal species. It is argued that intertidal fishes minimize the time spent out of the nest and the loss of contact with the substrate, features that may be of high survival value in conditions of strong turbulence. Although intertidal species have a reduced time available for feeding, this may be compensated by lower levels of locomotory activity, territorial defense and risk of egg predation. This revised version was published online in August 2006 with corrections to the Cover Date.