Seasonal and tidal changes in the length of the crystalline style intertidally living Macoma balthica (Mollusca,Bivalvia)

Samples of Macoma balthica were collected during 1978 and 1979 from the westernmost part of the Dutch Wadden Sea. The length of the crystalline style in the stomach of this intertidal bivalve was studied through tidal and annual cycles. The length of the style was ca. half the shell length and changed slightly with the season. Maximum lengths were recorded in spring, the season of high food intake, and minimal values in autumn and winter. Changes in the length of the style also exhibited a tidal cycle, but with two maxima and two minima. Tidal fluctuations in style length amounted to ca. 5% only. The influence of enzymes and waste products from the digestive diverticula, the food intake as well as the influence of the pH on the length of the style are discussed.Communication No. 401 of the Delta Institute     

Reproductive isolation between intertidal and subtidal<Emphasis Type="Italic"> Scoloplos armiger</Emphasis> (Polychaeta,Orbiniidae) indicates sibling species in the North Sea

Two distinct modes of development in the common polychaete Scoloplos armiger (O. F. Müller, 1776) occur in the North Sea region: holobenthic development in egg cocoons and pelagic larvae hatching from suspended eggs. In the northern Wadden Sea near the island of Sylt, we observed that egg cocoons are produced intertidally while pelagic larvae originate from the adjacent subtidal zone. A previous genetic comparison between these subtidal and intertidal populations revealed distinct gene pools, suggesting that reproductive differences are not phenotypic but heritable. In this study, crossbreeding experiments show that intertidal and subtidal populations are reproductively isolated. Couples with males and females from different habitats had no offspring. Production of egg cocoons is determined by female origin from the intertidal zone. Pelagic larvae occurred only in couples with subtidal females and subtidal males. Intertidal males have spermatozoa with heads twice as long as those from subtidal males and a significantly shorter flagellum. We suspect that deviating sperm morphology may cause the reproductive breakdown at the fertilization stage. Juveniles hatching from cocoons have shorter anal cirri compared to juveniles that metamorphosed from pelagic larvae. We conclude there to be two sympatric sibling species in S. armiger: 'type I' in intertidal areas, which have egg cocoons, no pelagic larvae, elongated sperm heads, shortened sperm flagella and anal cirri; and a subtidal 'type S', lacking egg cocoons but with pelagic larvae, short sperm heads, long sperm flagella and anal cirri.Communicated by O. Kinne, Oldendorf/Luhe     

Active pelagic migrations of the bivalve Macoma balthica are dangerous

The bivalve Macoma balthica migrates twice during the benthic part of its life cycle. During the spring migration (May-June), the newly settled spat (0-group) migrates to the nurseries in the high intertidal. Seven to nine months later, the bivalves migrate back to the low tidal flats and the subtidal (winter migration, 1-group). Both 0- and 1-group M. balthica use byssus threads for active pelagic migrations. As many M. balthica disappear during these migrations, we examined experimentally the importance of predation on 0- and 1-group M. balthica. Laboratory experiments using a circular aquarium determined predation rates on buried (no current) and drifting (current) 0- and 1-group M. balthica by several fish species (plaice, flounder, goby and whiting) and the shore crab. Under illuminated conditions, more M. balthica were consumed when migrating than when buried, whereas there was no difference between experiments in conditions of darkness. For the 0-group, predation rates on migrating and buried M. balthica in the dark were lower than in the light. The stomachs of pelagic fish in the Wadden Sea and Oosterschelde estuary did not contain M. balthica during winter migration. In the Wadden Sea, 1-group M. balthica primarily migrated at night. In conclusion, enhanced predation on drifting, as compared to buried, M. balthica may be the mechanism that explains enhanced mortality during migration in light, and may explain why M. balthica mainly migrates at night in the field. As we found no M. balthica in stomachs of pelagic fish, we do not know whether predation on byssus drifting M. balthica exists in the field. There are, however, some indications for fish predation on infaunal polychaetes during pelagic migrations.     

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     

Spatial and temporal variation in distribution and protein ubiquitination for <Emphasis Type="Italic">Mytilus</Emphasis> congeners in the California hybrid zone

Along the west coast of North America, the invasive mussel Mytilus galloprovincialis and a native congener M. trossulus overlap in range and compete for habitat in an extensive hybrid zone along central California. The two species have been shown to exhibit differential abiotic tolerances in laboratory studies, yet little is known about how such tolerances affect spatial and temporal patterns of geographic distribution, particularly in areas of competition. We examined distributions of the two congeners and their hybrids in neighboring intertidal and subtidal habitats in Bodega Bay, CA over 2 years, and compared shell length and seasonal ubiquitin (Ub) conjugates to estimate protein turnover and physiological stress for the species at each site. The two species were spatially segregated, with M. galloprovincialis dominating the subtidal habitat, and M. trossulus constituting a majority of the intertidal mussel population. Hybrid individuals appeared in low numbers at both sites. For each habitat, there was no statistical difference between shell lengths of M. galloprovincialis and hybrids but M. trossulus mussels were statistically smaller than the other two. In regards to physiological performance, ubiquitin conjugate values showed different seasonal cycles for the two species, suggesting different periods of peak environmental stress. The highest levels of Ub-conjugated proteins were observed in winter for M. galloprovincialis and in summer for M. trossulus, consistent with the respective range edges for their distributions since Bodega Bay is near the northern range edge of the invader and the southern edge of the native species. These findings suggest that future assessments of Mytilus populations along the California coast may need to consider vertical distributions and seasonal cycles as part of monitoring and research activities.     

Tisular and population level responses to habitat harshness in sandy beaches: the reproductive strategy of <Emphasis Type="Italic">Donax hanleyanus</Emphasis>

Microscopic analysis and field sampling procedures were used to compare demographic and reproductive strategies of the intertidal wedge clam Donax hanleyanus (Bivalvia: Donacidae) in two exposed sandy beaches with contrasting morphodynamics (reflective vs. dissipative) during 13 consecutive months. Histological analysis showed that: (1) the reproductive cycle of D. hanleyanus was more extended in the dissipative beach, and this was true for all the three pre-active (beginning of gonadal activity), active (maturation) and spawning stages; and (2) males and females showed significantly smaller sizes at sexual maturity at the reflective beach. Even though successive increments in proportion and mature at size were observed, the sigmoid function was significantly steeper at the reflective Arachania for both sexes, suggesting an abrupt transition to maturity. Field sampling revealed a more extended recruitment period at the dissipative beach, where recruits were also significantly more abundant than at the reflective beach. These results give support for the habitat harshness hypothesis, which predicts that in intertidal species capable of sustaining populations across a wide spectrum of physical conditions, such as D. hanleyanus, abundance, recruitment, size at maturity and extent of reproductive and recruitment seasons increase from reflective to dissipative beaches. However, a recent hypothesis suggests that reflective beaches acting as sink populations were not sustained, because mature and spawning individuals of both sexes were found in the reflective beach throughout the study period. Thus, we suggest that post-settlement processes are critical in modulating population patterns for this bivalve.     

Carrion-feeding on the sediment surface at nocturnal low tides by the polychaete<Emphasis Type="Italic"> Phyllodoce mucosa</Emphasis>

Harsh physical conditions in the intertidal zone are the cause of an ample amount of dead macroinvertebrates, which constitute a food source for carrion-feeders. In the European Wadden Sea, this trophic guild includes decapod crustaceans and fish when the tide is in, while during nocturnal low tides the polychaete Phyllodoce mucosa is attracted in large numbers by dead mollusks, crabs or worms on the sediment surface. Within 10 s worms emerged to the surface, crawled as far as 15 m on mucus trails towards the carcass, sucked in tissue up to one-third of their own weight, and then quickly retreated to below the surface. Abundance of P. mucosa was highest in the lower intertidal zone and winter. The seaward high abundance pattern, however, did not continue into the shallow subtidal. In summer, few were attracted during daytime or when the tide was in. However, up to 447 worms aggregated at a single crushed mussel within 20 min at dusk during low-tide exposure. This study suggests that during winter carrion-feeding is an important trophic niche on cold-temperate, intertidal mud flats occupied by a phyllodocid polychaete that is segregated in feeding time from most other scavengers and benefits from cold-sensitive benthic invertebrates.Communicated by O. Kinne, Oldendorf/Luhe     

Effects of Lunatia heros predation on the population dynamics of Mya arenaria and Macoma balthica in Maine,USA

The interaction between the naticid snail predator Lunatia heros and 2 iteroparous, infaunal, intertidal bivalves was investigated in Lubec, Maine, USA. The Mya arenaria population consists primarily of young, small individuals. M. arenaria survivorship is low when young (3.5% yr^-1 for the first 5 yr), then increases. M. arenaria can attain a length of 60 mm, but it is susceptible to L. heros attack only until it is 30 mm long. It delays reproduction until it is 20 mm long (3.8 yr) and diverts its resources instead into rapid early growth (4.9 mm yr^-1 for the first 5 yr). The Macoma balthica population has a larger proportion of older individuals than does that of M. arenaria. Survivorship is higher for M. balthica than for M. arenaria (76.3% yr^-1 for the first 5 yr). Unlike M. arenaria, M. balthica attains a final length of only 25 mm and all sizes are susceptible to L. heros attack. M. balthica grows slowly (2.7 mm yr^-1 for the first 5 yr) and diverts its resources into earlier reproduction at a length of 10 mm (2.9 yr). These contrasting life-history patterns and the possible relationship between bivalve resource allocation and refuges from predation are discussed.     

Seasonal and spatial differences in the fish assemblage of the non-estuarine Sulaibikhat Bay,Kuwait

In the period September 1986 to August 1987 fish were captured once a month, using an otter trawl, from the intertidal and subtidal regions of Sulaibikhat Bay, Kuwait. Correlation analysis revealed that numbers were not directly related to temperature at the time of sampling. The recruitment of large numbers of young of the year Liza carinata (Valenciennes) during March was related to low temperatures in November, the probable time of spawning of this species. Salinity at the time of sampling was inversely correlated with numbers. This result indicates that large numbers of 0+fish recruit to the Bay during the period of maximum fresh water outflow through the Shatt-al-Arab. Numbers of fish were significantly greater in the intertidal region, where they were present almost entirely as 0 group fish, than in the subtidal. The three dominant species of the assemblage are shown to use the two depth intervals in differnt ways. L. carinata was capture a almost exclusively in the intertidal region as 0+ fish. Pomadasys stridens (Forsskål) was captured as 0+ fish in both the intertidal and subtidal regions but in greater numbers in the intertidal region. Leiognathus brevirostris (Valenciennes) was captured as small, mostly 0+ individuals in the intertidal region and as larger fish in the subtidal region. The use made by the smaller fish of the intertidal region is related to the avoidance of sublittoral predators and reduction of intraspecific competition, whilst larger fish in the sublittoral region may be avoiding predation by piscivorous birds.     

Effect of periphyton grazing by Hydrobia ulvae on the growth of Zostera noltii on a tidal flat in the Dutch Wadden Sea

The decrease of the intertidal seagrass Zostera noltii in the Dutch Wadden Sea may have been the result of enhanced periphyton load due to eutrophication. Decrease of this seagrass species coincided with an increase in the mudsnail Hydrobia ulvae. Feeding of this mudsnail on periphyton may have partly counteracted an increase of periphyton biomass. Exclosure experiments on seagrass stands in the Dutch Wadden Sea in 1987 showed that density of periphyton on leaves of Z. noltii decreased significantly with increasing density of grazers. An increased density of mudsnails significantly enhanced the density and biomass of seagrass, in particular of the below ground parts. Since this seagrass species survives winter in temperate climate zones mainly by means of rhizomes, grazing may also influence the seagrass dynamics in the long term. Results of the experiment were in agreement with field observations on coinciding low densities of mudsnails and high densities of fouling of seagrass stands on the tidal flats of western Wadden Sea in the late 1970s.     

Siphon size and burying depth in deposit- and suspension-feeding benthic bivalves

This paper analyses the significance of siphon investment in the life strategy of benthic bivalves. It describes the relationships between siphon weight, burying depth and shell size in Mya arenaria, Cerastoderma edule. Scrobicularia plana and Macoma balthica. All data were collected on an intertidal flat in the Dutch Wadden Sea during seven successive winter and summer periods. The four species have in common that (1) the increase of depth in relation to size can be described with an S-curve; (2) there is a linear relationship between log siphon weight and log shell size; (3) siphon investment is maximal for the size classes with the greatest increase in their depth; (4) siphon weight, in proportion to total body weight, decreases gradually for the larger size classes whose depth does not increase; (5) burying depth increases with siphon weight if individuals within a same size class are compared, but burying depth levels off above a certain siphon weight. Macoma balthica and Scrobicularia plana live twice as deep in winter as in summer, although siphon weight for both seasons is about the same. In summer both species use a part of the siphon to graze the surface around the burrow, whereas deposit feeding does not occur in winter. This might explain the seasonal variation in burying depth. On the other hand Cerastoderma edule and Mya arenaria, which are both suspension feeders, show hardly any increase of depth in winter as compared to summer. For benthic bivalves the risk of being taken by a predator decreases with depth. The burying depth levels off where individuals reach the depth refuge (though in winter Scrobicularia plana live at greater depth). The conclusion is that siphon size is one of the main factors determining the burying depth of benthic bivalves and thus plays a critical role in their survival.     

Social spacing in the temperate marine goby Coryphopterus nicholsi

Coryphopterus nicholsi is a temperate marine goby that occupies rock rubble in protected subtidal areas along the Pacific coast of NOrth America. In field populations, fish of all sizes and both sexes are found in the same habitat. All observed fish over 25 mm standard length defended space during both the reproductive and the non-reproductive seasons, with territory size being directly correlated with fish size. There was peripheral overlap of territories, particularly between adjacent adult conspecifics of different size classes, but also with smaller fish that had territories centered in the interstices of larger territories. However, zones of overlap were used at different times by space-sharing fish so that defended areas remained temporally discrete. Access to areas of overlap appeared to be determined by dominance rank. More dominant (usually larger) fish used areas of overlap at will; the defense and use of shared areas by less dominant fish was contingent upon the absence of larger fish. Social organization in C. nicholsi appears to be defined by a combination of territorial behavior and dominance relationships among adjacent individuals, permitting the coexistence of all members of the population under circumstances of limited habitat availability.     

Clonal architecture in an intertidal bed of the dwarf eelgrass <Emphasis Type="Italic">Zostera noltii</Emphasis> in the Northern Wadden Sea: persistence through extreme physical perturbation and the importance of a seed bank

Genotypic structure and temporal dynamics of the dwarf seagrass, Zostera noltii, were studied in an intertidal meadow that has persisted since prior to 1936 near the Wadden Sea island of Sylt. Samples were collected from two 10 × 10 m plots separated by 250 m from May 2002 to June 2005 and from four 1 × 1 m plots from June 2003 to September 2004. All the samples were genotyped with nine microsatellite loci. No genotypes were shared between the plots separated by 250 m. Genetic diversity was higher in the Wadden Sea than in the other regions of its geographic range. The average clone size (genets) (SD) in the two plots was 1.38 (0.26) and 1.46 (0.4) m2, respectively, with a range up to 9 m2 and <20% persisted for >4 years. A high genetic and genotypic diversity was maintained by annual recruitment of seedlings despite a dramatic decrease in ramet density that coincided with the severe heat stress event of 2003. Fine-scale (1 m2) analysis suggested that extensive loss of seagrass cover precluded space competition among the genets, while a persistent seed bank prevented local extinction. Long-term persistence of Z. noltii meadows in the intertidal Wadden Sea was achieved by high genet turnover and frequent seedling recruitment from a seed bank, in contrast to the low diversity observed in large and long-living clones of Z. noltii and other seagrasses in subtidal habitats.     

Assemblage shift following population collapse of a non-indigenous bivalve in an urban lagoon

The quahog Mercenaria mercenaria has been introduced repeatedly to the Pacific coast of North America, but only one population is known to have become established. In the 1970s, the population of M. mercenaria at Colorado Lagoon, in Los Angeles County, California (33^o46′16″N, 118^o08′05″W), was estimated at more than 300,000 individuals. To determine the current status of this non-indigenous species (NIS), in 2009, we sampled 57 intertidal and 20 shallow subtidal plots, identifying and quantifying collected bivalves. No quahogs were found among the 2,490 living bivalves in our plots, though two were found intertidally outside of our plots. The M. mercenaria population has thus collapsed since 1980, but the native community has not recovered. Six of the fourteen living bivalve species we encountered were NIS; three are new records for the location, including the clam Venerupis philippinarum, which made up 87.6% of collected individuals. Though M. mercenaria is likely on its way to extinction on the US Pacific coast, the bivalve assemblage at this location remains heavily dominated by NIS.     

Reintroduction of eelgrass (Zostera marina) in the Dutch Wadden Sea; review of research and suggestions for management measures

Eelgrass (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal flats. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Levels of genetic polymorphism at two enzyme encoding loci in eight species of the genus Macoma (Mollusca: Bivalvia)

Levels of genetic polymorphism were surveyed at two enzyme loci (LAP, PGI) in 2 intertidal and 6 subtidal species of the bivalve genus Macoma living in the waters of the San Juan Islands (Washington and Canada). The temporal environmental variability-genetic variability hypothesis predicts that intettidal species should have greater levels of genetic polymorphism than subtidal species. This is not true for the genus Macoma. However, at the PGI locus, genetic polymorphism was proportional to niche breadth in both the intertidal and subtidal species, but only for intertidal species at the LAP locus. These results support the contention that temporal environmental variability is not necessarily important in maintaining genetic variability, but that environmental heterogeneity may select for increased polymorphism at some loci.     

Reintroduction of eelgrass ( Zostera marina ) in the Dutch Wadden Sea; Review of research and suggestions for management measures

Eelgras (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal falts. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Are Wadden Sea tidal systems with a higher tidal range more resilient against sea level rise?

Accelerated sea level rise may have serious implications for the Wadden Sea ecosystem in its present state. If sediment accumulation rates on the extensive intertidal flats stay behind sea level rise, the flats will eventually submerge. Drowning of the flats has negative consequences for nature conservation and for coastal risk management. Based upon an evaluation of steady state relations for Wadden Sea tidal basins, Hofstede (Zeitschrift für Geomorphologie 59(3): 377-391, 2015) postulated that the capacity of these basins to balance sea level rise by accumulation on intertidal flats seems positively related to mean tidal range. In the present study, morphodynamical simulations with a numerical model were performed for two tidal basins in the German Wadden Sea to verify the empirically established hypothesis. The following conclusions are established. Larger mean tidal range improves the capacity of Wadden Sea tidal basins to balance sea level rise. Wadden Sea intertidal flats are effective sediment sinks and seem quite resilient against (higher rates of) sea level rise. Finally, subtidal gullies may constitute a significant sediment source for accumulation on intertidal flats in response to sea level rise. With respect to the limited comparability of the two investigated tidal systems, morphodynamical modelling of all Wadden Sea tidal systems should be conducted.