Age-dependent zonation of the periwinkle Littorina littorea (L.) in the Wadden Sea

On sedimentary tidal flats near the island of Sylt (German Bight, North Sea) abundance and size distribution of periwinkles, Littorina littorea L., were studied in low intertidal and in shallow and deep subtidal mussel beds (Mytilus edulis L.). In low intertidal mussel beds, surveys revealed that high densities (1,369±571 m^–2) of juvenile snails (≤13 mm) were positively correlated with strong barnacle epigrowth (Semibalanus balanoides L. and Balanus crenatus Bruguière) on mussels. A subsequent field experiment showed that recruitment of L. littorea was restricted to the intertidal zone. Abundances of periwinkles (213±114 m^–2) and barnacles abruptly decreased in the adjacent shallow subtidal zone, which served as a habitat for older snails (>13 mm). L. littorea was completely absent from disjunct deep (5 m) subtidal mussel beds. Snail abundance varied seasonally with maxima of >4,000 m^–2 in low intertidal mussel beds in October and minima in July, just before the onset of new recruitment. I suggest that the presence of cracks and crevices among the dense barnacle overgrowth in intertidal mussel beds favoured recruitment and survival of juvenile snails. Larger (older) specimens are assumed to actively migrate to the less favourable adjacent subtidal. Therefore, intertidal mussel beds are considered as nurseries for the population of L. littorea in the Wadden Sea. Received in revised form: 25 September 2000 Electronic Publication     

Subtidal and intertidal mussel beds (Mytilus edulis L.) in the Wadden Sea: diversity differences of associated epifauna

In 1997 and 1998, surveys were performed to compare species composition, abundance and diversity of non-attached epifauna (>1 mm) in low intertidal and adjacent shallow subtidal zones of three mussel beds (Mytilus edulis L.) near the island of Sylt in the North Sea. The community structure was similar when compared within tidal zones: no significant differences in species numbers and abundances were recorded between locations and between years. A comparison between tidal zones, however, revealed higher diversity, species densities and total species numbers in the subtidal (per 1,000 cm²: H ′=2.0±0.16; 12 ±1 species density; 22 species) than the intertidal zone (per 1,000 cm²: H ′=0.7±0.27; 6±2 species density; 19 species). Abundances significantly dropped with increasing submergence from 2,052 (±468) m^–2 to 1,184 (±475) m^–2. This was mainly due to significantly higher densities of both juvenile periwinkles, Littorina littorea, and crabs, Carcinus maenas, in intertidal mussel beds. However, many less dominant species were significantly more abundant in subtidal mussel beds. This study revealed that in the non-attached epifaunal community of mussel beds the tidal level effect within metres was strong, whilst the spatial variability in a much wider (kilometre) range but the same tidal level was negligible. The high epifaunal diversity in the subtidal zone suggests that the protective measures for mussel beds against the effects of mussel fishery should be extended from the intertidal to the subtidal zone, if the integrity of the mussel bed community in the Wadden Sea National Park is to be maintained. Electronic Publication     

The introduced clam Ensis americanus in the Wadden Sea: field experiment on impact of bird predation and tidal level on survival and growth

In the Danish Wadden Sea the intertidal distribution of the introduced bivalve Ensis americanus (syn. E. directus) is restricted to a narrow zone around the mean low water level. To test the possible impact of birds and submersion time on dynamics and distribution of the clams, adult specimens of E. americanus collected near the low water line were transplanted to two intertidal sites and established in open and net-covered experimental plots for 9 weeks (autumn 2001). The lowest survival of clams was registered at the low-shore-site (LSS) in plots open to bird predators, suggesting that birds such as Common Eider (Somateria mollissima) or Oystercatcher (Haematopus ostralegus) may control the abundance of E. americanus at the lower tidal levels. For clams showing increment in shell length during the study period, the shell growth rates were highest at the LSS and lowest in the open plots at the high-shore-site (HSS). Differences in immersion time and thus food supply may explain this pattern. Body mass index (BMI) of the clams showed basically the same pattern as the survivorship: lowest BMI in open plots at the LSS and highest in the covered plots at this site. Clams from the HSS were intermediate in their BMI. Disturbance by birds in the open plots at the LSS may explain the low BMI. In conclusion birds may be an important factor controlling abundance of E. americanus in the lower intertidal zone.     

High survival and growth rates of introduced Pacific oysters may cause restrictions on habitat use by native mussels in the Wadden Sea

Pacific oysters Crassostrea gigas (Thunberg, 1793) were introduced to the northern Wadden Sea (North Sea, Germany) by aquaculture in 1986 and finally became established. Even though at first recruitment success was rare, three consecutive warm summers led to a massive increase in oyster abundances and to the overgrowth of native mussel beds (Mytilus edulis L.). These mussels constitute biogenic reefs on the sand and mud flats in this area. Survival and growth of the invading C. gigas were investigated and compared with the native mussels in order to predict the further development of the oyster population and the scope for coexistence of both species. Field experiments revealed high survival of juvenile C. gigas (approximately 70%) during the first three months after settlement. Survival during the first winter varied between > 90% during a mild and 25% during a cold winter and was independent of substrate (i.e., mussels or oysters) and tide level. Within their first year C. gigas reached a mean length of 35-53 mm, and within two years they grew to 68-82 mm, which is about twice the size native mussels would attain during that time. Growth of juvenile oysters was not affected by substrate (i.e., sand, mussels, and other oysters), barnacle epibionts and tide level, but was facilitated by fucoid algae. By contrast, growth of juvenile mussels was significantly higher on sand flats than on mussel or oyster beds and higher in the subtidal compared to intertidal locations. Cover with fucoid algae increased mussel growth but decreased their condition expressed as dry flesh weight versus shell weight. High survival and growth rates may compensate for years with low recruitment, and may therefore allow a fast population increase. This may lead to restrictions on habitat use by native mussels in the Wadden Sea.     

Predation on barnacles of intertidal and subtidal mussel beds in the Wadden Sea

Balanids are the numerically dominant epibionts on mussel beds in the Wadden Sea. Near the island of Sylt (German Bight, North Sea), Semibalanus balanoides dominated intertidally and Balanus crenatus subtidally. Field experiments were conducted to test the effects of predation on the density of barnacle recruits. Subtidally, predator exclusion resulted in significantly increased abundances of B. crenatus, while predator exclusion had no significant effects on the density of S. balanoides intertidally. It is suggested that recruitment of B. crenatus to subtidal mussel beds is strongly affected by adult shore crabs (Carcinus maenas) and juvenile starfish (Asterias rubens), whereas recruits of S. balanoides in the intertidal zone are mainly influenced by grazing and bulldozing of the very abundant periwinkle Littorina littorea, which is rare subtidally. Thus, not only do the barnacle species differ between intertidal and subtidal mussel beds, but the biotic control factors do so as well. Electronic Publication     

Historical changes in the benthos of the Wadden Sea around the island of Sylt in the North Sea

The situation regarding the distribution and abundance of seagrass, macroalgae and benthic fauna near the island of Sylt in the south-eastern North Sea during the period 1923 to 1940 is compared with that of the 1980s. Evidence of organic enrichment in recent times is provided by (1) massive growth of green algal mats on sheltered tidal flats, (2) a decline of red algae in the subtidal zone, (3) an expansion of mussel beds along low water line and down to 20 m depth, (4) increased abundance of polychaetes inhabiting intertidal and subtidal sandy bottoms. Seagrass beds have undergone complex changes which remain unexplained. Intensified erosion has contributed to the loss of habitats in the intertidal zone, and probably affected sessile epifauna in the deep channels. Here, direct removal and disturbance by the bottom-trawling fishery may also have contributed to the observed species impoverishment. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Embryonic and larval development of Ensis arcuatus (Jeffreys, 1865) (Bivalvia: Pharidae)

The razor clam Ensis arcuatus (Jeffreys, 1865) is distributedfrom Norway to Spain and along the British coast, where it livesburied in sand in low intertidal and subtidal areas. This workis the first study to research the embryology and larval developmentof this species of razor clam, using light and scanning electronmicroscopy. A new method, consisting of changing water levelsusing tide simulations with brief dry periods, was developedto induce spawning in this species. The blastula was the firstmotile stage and in the gastrula stage the vitelline coat waslost. The shell field appeared in the late gastrula. The trochophoredeveloped by about 19 h post-fertilization (hpf) (19°C).At 30 hpf the D-shaped larva showed a developed digestive systemconsisting of a mouth, a foregut, a digestive gland followedby an intestine and an anus. Larvae spontaneously settled after20 days at a length of 378 µm. (Received 5 December 2006; accepted 19 November 2007)     

Histological survey of symbionts and other conditions in razor clam Ensis arcuatus (Jeffreys, 1865) (Pharidae) of the coast of Galicia (NW Spain)

Symbionts and abnormal conditions of razor clam Ensis arcuatus were surveyed in three commercially important natural beds of Galician estuaries (NW Spain). Samples of 15-20 E. arcuatus were collected every 2 months from January 2003 until July 2004 and processed for histological examination. Prokaryote-like colonies, renal coccidians, gregarines, Trichodina sp. ciliates, haplosporidian-like plasmodia, turbelaria, trematode metacercariae, cestode-like larvae and basophilic inclusion bodies were observed in razor clam tissues without causing host damage. Bucephalid digenean sporocysts and germinoma were seen in some samples causing moderate or severe damage to the host depending on the intensity of infection and both could be a cause for concern if prevalence reached epizootic levels in Galician E. arcuatus populations. None of the parasites detected is OIE notifiable and, in general, the commercially exploited beds studied seem to be devoid of serious pathogens.     

How subtidal were the ’subtidal beds’ ofZostera marina L. before the occurrence of the wasting disease in the early 1930's?

In the available literature about eelgrass beds in the former Zuiderzee, now western Dutch Wadden Sea, it is suggested that these stands mainly occurred in the subtidal zone. There is, however, no evidence that this suggestion is right because the relative distribution of eelgrass beds over the intertidal zone and the subtidal zone in the former Zuiderzee was never studied. A detailed mapping of eelgrass beds was carried out in 1869 and another one in 1930. The 1869 map is combined with available nautical charts by application of GIS packages and the distribution of eelgrass beds calculated. It was necessary to carry out corrections because of inaccuracies both in the eelgrass mapping and in the old nautical charts. It is concluded that in 1869 45%±4 of the eelgrass beds were present in the intertidal zone and that 55%±4 of the beds were present in the subtidal zone. In 1930 the distribution amounted to 56% in the intertidal zone versus 44% in the subtidal zone. These values illustrate that in the period prior to the wasting disease and the closure of the former Zuiderzee, eelgrass occurred in both the intertidal zone as well as in the subtidal zone.     

Subtidal-intertidal trophic links: American lobsters [Homarus americanus (Milne-Edwards)] forage in the intertidal zone on nocturnal high tides

Homarus americanus (Milne-Edwards), the American lobster, is a predator in New England subtidal communities, feeding on ecologically important grazers (sea urchins), mesopredators (crabs), and basal species (mussels). In this study, we provide the first report of adult American lobsters foraging in rocky intertidal habitats during nocturnal high tides. Censuses by SCUBA divers in the low intertidal (Chondrus crispus Stackhouse) zone showed mean densities of 2.2 lobsters/20 m² on nocturnal high tides, with contrasting low densities of 0.18/20 m² during diurnal high tides. Nocturnal high-tide intertidal densities were 62% of those reported in a previous study of lobsters in nearby subtidal rocky areas (Novak, 2004). The average carapace length of lobsters in the intertidal at night was > 50 mm. These lobsters were actively foraging in the intertidal with collected individuals having a mean stomach fullness of 67%. Prey found in the stomach contents primarily consisted of crabs, mussels and snails. Field experiments showed that lobsters rarely fed on medium to large size individuals of the common intertidal snail, Littorina littorea (L.). In contrast, experiments with local crab species demonstrated that lobsters actively and readily prey on Cancer irroratus (Say) and Carcinus maenas (L.), but were significantly less likely to consume Cancer borealis (Stimpson). The abundance of Carcinus maenas and blue mussels (Mytilus edulis L.) in the intertidal zone may explain the upshore movement of lobsters. Since nocturnal migration of Homarus americanus into the intertidal zone has not been documented before, our understanding of the dynamics of New England intertidal communities needs to be expanded to include this predator.     

Disturbance and organization of macroalgal assemblages in the Northwest Atlantic

Large seaweeds are often structurally dominant in subtidal and intertidal rocky shore benthic communities of the N.W. Atlantic. The mechanisms by which these algal assemblages are maintained are surprisingly different in the two habitats. In the subtidal community, kelps are dominant space competitors in the absence of strong grazing interactions. In contrast, the large perennial seaweeds of intertidal zones (fucoids and Chondrus crispus) are competitively inferior to both sessile filter feeders and ephemeral, pioneer algal species. Intertidal seaweed beds are maintained by carnivory of whelks, which reduces filter feeder populations, and by herbivorous periwinkles which reduce ephemeral algal populations. Through most of the intertidal zone, disturbance, both biological and physical, dictates which species shall compete and equilibrium conditions obtain subsequently.The roles of subtidal consumers are quite different. Sea urchins are the major algal herbivores and these voracious animals maintain an equilibrium state in which large tracts of subtidal coralline pavement are kept free of kelp forests. Urchins do not seem to play a successional facilitative role for kelps in the way that periwinkles do for fucoids in the intertidal. Control of herbivore populations is thus a key to the maintenance of subtidal foliose algal beds. It is clear that parasitic amoebas can decimate sea urchin populations so that kelp forest dominance is assured. However, the importance of carnivory in limiting urchins in the subtidal community is unclear in the absence of appropriate manipulation experiments. It is possible that carnivorous decapods and fin fish control sea urchin populations and hence foliose algal abundance, but this must remain speculative. The seaweed-dominated state of the subtidal system is an alternative equilibrium condition to the urchin/coralline alga configuration. The structure of the kelp beds is relatively uniform in responding to frequent small-scale, infrequent large-scale, or no, disturbance.     

Effects of the severe winter 1995/96 on the benthic macrofauna of the Wadden Sea and the coastal North Sea near the island of Sylt

The development of benthic macrofauna in the Wadden Sea and in the coastal North Sea after the severe winter of 1995/96 is compared with the preceding years with mild to moderate winters. In the intertidal of the Wadden Sea, ice-drift and low temperature caused the expected changes in species composition by increasing winter mortality in sensitive species, and by exceptionally high recruitment of some species during the succeeding summer. In the shallow subtidal (10–20 m depth), similar winter effects were observed. However, recovery of many subtidal populations was still incomplete until the summer of 1997. It is suggested that this was due to hydrographic conditions that carried many larvae or drifting juveniles into more distant offshore areas. This may have limited larval supply and may have delayed recovery at the onshore sites. Since in the eastern North Sea severe winters are accompanied by frequent easterly winds, it is not clear whether decreasing winter abundances in some species were due to increased mortality, or to a seaward dislocation of organisms. Received in revised form: 7 May 2001 Electronic Publication     

Material exchange and food web of seagrass beds in the Sylt-Rømø Bight: how significant are community changes at the ecosystem level?

Material exchange, biodiversity and trophic transfer within the food web were investigated in two different types of intertidal seagrass beds: a sheltered, dense Zostera marina bed and a more exposed, sparse Z. noltii bed, in the Northern Wadden Sea. Both types of Zostera beds show a seasonal development of above-ground biomass, and therefore measurements were carried out during the vegetation period in summer. The exchange of particles and nutrients between seagrass beds and the overlying water was measured directly using an in situ flume. Particle sedimentation [carbon (C), nitrogen (N) and phosphorus (P) constituents] from the water column prevailed in dense seagrass beds. In the sheltered, dense seagrass bed, a net particle uptake was found even on windy days (7–8 Beaufort). Dissolved inorganic N and orthophosphate were mainly taken up by the dense seagrass bed. At times of strong winds, nutrients were released from the benthic community to tidal waters. In a budget calculation of total N and total P, the dense seagrass beds were characterised as a material sink. The seagrass beds with sparse Z. noltii were a source of particles even during calm weather. The uptake of dissolved inorganic N in the sparse seagrass bed was low but significant, while the uptake of inorganic phosphate and silicate by seagrasses and their epiphytes was exceeded by release processes from the sediment into the overlying water. Estimates at the ecosystem level showed that material fluxes of seagrass beds in the Sylt-Rømø Bight are dominated by the dense type of Zostera beds. Therefore, seagrass beds act as a sink for particles and for dissolved inorganic nutrients. During storms, seagrass beds are distinct sources for inorganic nutrients. The total intertidal area of the Sylt-Rømø Bight could be described as a sink for particles and a source for dissolved nutrients. This balance of the material budget was estimated by either including or excluding seagrass beds. Including the subtidal part, the function of the ecosystem as a source for particles increased, supposing that all seagrass beds were lost from the area. During the vegetation period, seagrass beds act as a storage compartment for material accumulated in the living biomass of the community. There was great biodiversity among the plant and animal groups found in intertidal seagrass beds of the Sylt-Rømø Bay, representing 50–86% of the total number of species investigated, depending on the particular group. Since most species are not exclusively seagrass residents, the loss of intertidal seagrass beds would be of minor importance for biodiversity at the ecosystem level. Food web structure in seagrass beds is different from other intertidal communities. Primary production and detritus input is high, but secondary production is similar to that of unvegetated areas, although the relative importance of the trophic guilds is different. The loss of seagrass beds leads to profound alterations in the food web of the total ecosystem. Historical as well as recent changes in material fluxes and energy flow due to man-made alterations to the ecosystem are discussed.     

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     

Effects ofFucus vesiculosus covering intertidal mussel beds in the Wadden Sea

The brown algaFucus vesiculosus formamytili (Nienburg) Nienhuis covered about 70% of mussel bed (Mytilus edulis) surface area in the lower intertidal zone of Königshafen, a sheltered sandy bay near the island of Sylt in the North Sea. Mean biomass in dense patches was 584 g ash-free dry weight m^?2 in summer. On experimental mussel beds, fucoid cover enhanced mud accumulation and decreased mussel density. The position of mussels underneath algal canopy was mainly endobenthic (87% of mussels with >1/3 of shell sunk into mud). In the absence of fucoids, mussels generated epibenthic garlands (81% of mussels with <1/3 of shell buried in mud). Mussel density underneath fucoid cover was 40 to 73% of mussel density without algae. On natural beds, barnacles (Balanidae), periwinkles (Littorina littorea) and crabs (particularly juveniles ofCarcinus maenas) were significantly less abundant in the presence of fucoids, presumably because most of the mussels were covered with sediment, whereas in the absence of fucoids, epibenthic mussel clumps provided substratum as well as interstitial hiding places. The endobenthic macrofauna showed little difference between covered and uncovered mussel beds. On the other hand, grazing herbivores — the flat periwinkleLittorina mariae, the isopodJaera albifrons and the amphipodsGammarus spp. — were more abundant at equivalent sites with fucoid cover. The patchy growth ofFucus vesiculosus on mussel beds in the intertidal Wadden Sea affects mussels and their epibionts negatively, but supports various herbivores and increases overall benthic diversity.     

The fate of an immigrant: Ensis directus in the eastern German Bight

We studied Ensis directus in the subtidal (7–16?m depth) of the eastern German Bight. The jack-knife clam that invaded in the German Bight in 1978 has all characteristics of a successful immigrant: Ensis directus has a high reproductive capacity (juveniles, July 2001: Amrumbank 1,914?m^?2, Eiderstedt/Vogelsand: 11,638?m^?2), short generation times and growths rapidly: maximum growth rates were higher than in former studies (mean: 3?mm?month^?1, 2nd year: up to 14?mm?month^?1). Ensis directus uses natural mechanisms for rapid dispersal, occurs gregariously and exhibits a wide environmental tolerance. However, optimal growth and population-structure annual gaps might be influenced by reduced salinity: at Vogelsand (transition area of Elbe river), maximum growth was lower (164?mm) than at the Eiderstedt site (outer range of Elbe river, L _∞?=?174?mm). Mass mortalities of the clams are probably caused by washout (video inspections), low winter temperature and strong storms. Ensis directus immigrated into the community finding its own habitat on mobile sands with strong tidal currents. Recent studies on E. directus found that the species neither suppresses native species nor takes over the position of an established one which backs up our study findings over rather short time scales. On the contrary, E. directus seems to favour the settlement of some deposit feeders. Dense clam mats might stabilise the sediment and function as a sediment-trap for organic matter. Ensis directus has neither become a nuisance to other species nor developed according to the ‘boom-and-bust’ theory. The fate of the immigrant E. directus rather is a story of a successful trans-ocean invasion which still holds on 23?years after the first findings in the outer elbe estuary off Vogelsand.     

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.     

The expanding range of Undaria pinnatifida in southern New Zealand: distribution, dispersal mechanisms and the invasion of wave-exposed environments

Very few studies have addressed how the invasive kelp Undaria pinnatifida (Harvey) Suringar spreads beyond initial founding populations in harbours. Surveys of the harbours and accessible areas of open coast throughout southern New Zealand were conducted to determine how far U. pinnatifida populations had extended since initial incursions. Our findings clearly demonstrate that U. pinnatifida is capable of invading native communities and can establish reproductive populations in locations subjected to significant and consistent wave action. The extent of spread from source populations differs between harbours in which it has established. Dispersal is greatest in harbours with long established populations, those where populations have not been strategically managed, harbours with high water exchange with surrounding coastal waters, and where prevailing currents allow establishment of U. pinnatifida on suitable substrata close to harbour entrances. Dispersal along the open coast is primarily achieved by drifting adult sporophytes that are washed up in the rocky intertidal zone. Founding populations are most often found in the intertidal zone, primarily within rockpools. Subtidal transects and observations indicate that U. pinnatifida is well adapted to invade exposed coastlines and can establish within a broad range of niches in wave-exposed areas including rockpools, the low intertidal, shallow subtidal, Macrocystis pyrifera kelp forests, and in low light areas beyond the vertical extent of large native macroalgae. The current range of U. pinnatifida is much greater than expected and appears to be expanding. Due to its ability to grow in a broad range of environments and to form dense monospecific stands, U. pinnatifida has the potential to strongly modify almost all rocky subtidal and intertidal communities in temperate locations.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

Production and population ecology ofPhyllospadix iwatensis Makino. II. Comparative studies on leaf characteristics,foliage structure and biomass change in an intertidal and subtidal zone

A seagrass in Japan,Phyllospadix iwatensis Makino, is distributed in the lower intertidal zone and upper subtidal zone making a dense population on the Choshi coast, Japan. IntertidalP. iwatensis is able to receive sufficient light for photosynthesis but experienced severe exposure to the air, which decreased a large amount of aboveground biomass in April to June (i.e. the daytime exposure season). SubtidalP. iwatensis was never exposed throughout the year and the aboveground biomass increased gradually over the daytime exposure season. However, the maximum aboveground biomass and shoot density of the subtidal plant never exceeded that of the intertidal plant. The dense foliage, large aboveground biomass and high shoot density of both intertidal and subtidal plants is likely to be an adaptation to heavy water movement, but the subtidal plants always received insufficient light for photosynthesis as a result of having dense foliage, particularly in turbid water. In choppy and swell sea,P. iwatensis did not seem to be adapted to growing in the subtidal zone where there was shortage of light.