Documentation of sites of intertidal blue mussel (Mytilus edulis L.) beds of the Lower Saxonian Wadden Sea, southern North Sea (as of 2003) and the role of their structure for spatfall settlement

Field surveys (dating back to 1950) and aerial photograph series (dating back to 1966) were evaluated to determine sites of intertidal blue mussel (Mytilus edulis) beds at the Wadden Sea coast of Lower Saxony. Maps were prepared indicating sites of blue mussel beds during the last decades. A table gives additional information on the presence (or absence) of blue mussel beds at each site at the time of large-scale surveys. Altogether 187 sites of M. edulis beds were recorded in the investigation area. In spring 1996, there were still only 19 sites where mussel beds still occurred, although at 51 sites residual mussel-bed structures were present, e.g. shell bases of former beds or protruding patches (which had been occupied by M. edulis before the beds vanished) and open spaces. At that time, the majority of the sites contained neither mussel beds nor mussel-bed structures. The analysis of recent data confirmed that mussel larvae have preferred to settle in sites of present mussel beds and sites with bases of former mussel beds. There was no preferential selection of one of these categories (settled beds vs. shell bases). On the other hand, the presence of mussel beds or mussel bed structures is not obligatory for settlement, because sites without those structures were also re-settled by the spatfall in 1996, even though on a smaller scale.     

Mussel beds on different types of structures support different macroinvertebrate assemblages

Abstract Artificial structures, such as seawalls, pilings and pontoons, are common features of urban estuaries. They replace natural structures or add to the amount of hard substratum in an area and provide habitats for many fish and invertebrates. Previous work has concentrated on fish or on the invertebrates that occupy the primary substratum of artificial structures. Mussels often grow on different types of structures (pontoons, pilings, seawalls and natural reefs) and provide a secondary substratum for other organisms to inhabit. Counting and identifying organisms associated with mussel beds is traditionally done to species level, which is very time‐consuming. To save time, organisms in this study were identified to coarse levels of taxonomic resolution (a mix of taxa, such as class, order, family and genus), which showed similar patterns to those when particularly speciose and abundant groups were identified to species. This study tests hypotheses that the distribution and abundance of mobile and sessile organisms that inhabit mussel beds will differ among natural and various types of artificial structures. When the associated assemblages of mussel beds from different types of structures and from different locations were examined, assemblages varied according to the type of structure they inhabited and its location. Assemblages associated with mussels on pontoons differed consistently from those on other types of structures. Patterns in the assemblages were also consistent through time. These data show that the types and amounts of artificial structures added to an environment can affect the types, distribution and abundances of organisms living in biogenic habitats.     

Diversity in mussel beds at deep-sea hydrothermal vents and cold seeps

Remarkably little is known about fundamental distinctions (or similarities) between the faunas of deep‐sea hydrothermal vents and seeps. Low species richness at vents has been attributed to the transient nature of vent habitats and to toxic effects of hydrogen sulphide and heavy metals in vent effluents. Seeps are arguably more stable and more chemically benign than vents. They have also been regarded as more diverse, but until now there has not been a rigorous test of this hypothesis. We evaluated diversity indices for invertebrates associated with mussel beds at six vents and two seeps and found that invertebrate diversity was significantly higher at seeps than vents, although some vent mussel beds supported nearly the same diversity as seep mussel beds. Lower diversity at vents may be a consequence of a greater physiological barrier to invasion at vents than at seeps. Diversity was lowest where spacing between vents was greatest, suggesting that risks of extinction as a result of dispersal‐related processes may contribute to the pattern of diversity observed at vents.     

Biodeposition in a juvenile mussel bed of the east Frisian Wadden Sea (Southern North Sea)

Seasonal trends of biodeposition and animal coverage in backbarrier beds of the musselMytilus edulis in the East Frisian Wadden Sea were documented and quantified to assess the effects of physical disturbance by storms and to develop a conceptual model for the interpretation of biodeposits in the stratigraphic record. Accretion and erosion of biogenic muds (faeces and pseudofaeces) were quantified by repeated precision levelling of a 12 m² test area. Animal coverage was documented by vertical photography prior to each levelling survey. Levelling and photography were conducted from a bridge and rail system to avoid physical disturbance by trampling. Four surveys, covering a period of different weather conditions, showed variable mean accretion/erosion rates of biodeposits ranging from +0.56 mm d⁻¹ (accretion) to −1.34 mm d⁻¹ (erosion) and areal coverage of mussels ranging from 39.1% to 0%. A pronounced seasonal trend in the biodepositional pattern was observed. Mean accretion rates were high (>0.5 mm d⁻¹) during the summer season, whereas in the winter season net deposition decreased to zero or showed erosional trends. Longterm sediment budgets are thus composed of highly variable mean annual rates which, in turn, comprise highly variable seasonal and even monthly rates, depending on the local weather pattern. This forms the basis for a conceptual model, in terms of which the stratigraphic record of fossil biodeposits can be meaningfully interpreted.     

Causes of species diversity differences: a comparative analysis of Markov models

While many dynamic processes have been proposed to produce diversity differences among communities, most empirical investigations focus on static system attributes. An ideal analysis would consider multiple dynamic processes and their impact on many community members, but such analyses can be logistically daunting. I compared Markov chain models of ecological communities to explore general processes leading to diversity differences of sessile species between coral reefs and rocky intertidal mussel beds. As predicted by diversity theory, high diversity coral reefs had lower species replacement probabilities and higher disturbance rates than did lower diversity mussel beds. Intransitivities in species replacements, recruitment limitation and responses to perturbing species from equilibrium (Jacobian elements) did not differ significantly between systems. Tradeoffs between the displacement risk or displacement ability of a species and either disturbance risk or colonising ability were not apparent. Manipulating the coral reef model to eliminate disturbance or intransitivities caused declines in species diversity, whereas removing recruitment limitation or increasing the probability of interspecific replacement did not. Higher overall disturbance levels can explain part of the diversity difference between systems, but much remained unexplained, indicating that details of the pattern and strengths of species interactions were probably extremely important.     

Direct and indirect effects of Littorina littorea (L.) on barnacles growing on mussel beds in the Wadden Sea

On the extensive sedimentary tidal flats of the Wadden Sea, beds of the blue mussel Mytilus edulis represent the only major hard substratum and attachment surface for sessile organisms. On this substratum, the barnacle Semibalanus balanoides is the most frequent epibiont. In summer 1998, it occurred on over 90% of the large mussels (>45 mm shell length) and the dry weight of barnacles reached 65% of mussel dry weight. However, the extent of barnacle overgrowth is not constant and differs widely between years. Periwinkles (Littorina littorea) may reach densities >2000 m^–2 on intertidal mussel beds. Field experiments were conducted to test the effect of periwinkle grazing on barnacle densities. An experimental reduction of grazing and bulldozing pressure by periwinkles resulted in increased recruitment of barnacles, while barnacle numbers decreased with increasing snail density. The highest numbers of barnacles survived in the absence of L. littorea. However, a lack of periwinkle grazing activity also facilitated settlement of ephemeral algae which settled later in the year. Field experiments showed that the growth rate of barnacles decreased in the presence of these ephemeral algae. Thus, L. littorea may reduce initial barnacle settlement, but later may indirectly increase barnacle growth rate by reducing ephemeral algae. It is suggested that periwinkle density may be a key factor in the population dynamics of S. balanoides on intertidal mussel beds in the Wadden Sea.     

Prolonged Lag in Population Outbreak of an Invasive Mussel: A Shifting-Habitat Model

Biological invasions pose a great threat to the integrity of natural communities. Some invasive species demonstrate a population explosion shortly after arrival while in other cases a prolonged lag between arrival and population outbreak is evident. This paper describes a case of a prolonged lag and explores the possible mechanism for this lag. The Red Sea mussel Brachidontes pharaonis, a Lessepsian migrant, was first recorded in the Mediterranean seven years after the opening of the Suez Canal in 1869. Since then it spread along the Israeli coast and as far northwest as Sicily. Studies conducted in the late 1970s, when B. pharaonis was still rare, predicted that it would not establish dense populations along the Israeli coast and would not outcompete the indigenous mussel Mytilaster minimus, although it has strong negative effects on survival and growth of the native species. It was attributed to the invader's low intrinsic rate of increase relative to that of the native species, and to strong density-independent mortality generated by exposure to high wave action and sedimentation. In contrast to these predictions, we found massive formations of B. pharaonis beds after lag of about 120 years. We looked for distributional patterns that may explain this lag and found no south–north gradient but a strong habitat-dependent colonization. Most apparent are dense B. pharaonis mussel beds (density >300 per 100cm²) on rocky platforms where mussel beds were absent in the past. These platforms lack the vermetid rim that is typical to this formation. In platforms protected by a biogenic rim, sediment accumulation is high and perennial algae flourish. None of the mussel species form beds in such habitats. We suggest that the delayed formation of B. pharaonis beds along the Israeli coast is a consequence of a recent shift in habitat conditions on some platforms. It is possible that receding of the biogenic rim at the edge of these platforms allowed more effective washing, reduced sediment accumulation, and reduced perennial algae cover making platforms more suitable for the mussels. Lower density-independent mortality allowed B. pharaonis to dominate on such platforms over the indigenous species. On beachrock, a habitat previously dominated by M. minimus, we recorded a rapid shift in numerical domination to B. pharaonis (from 1:7 to 1.4:1 Brachidontes/Mytilaster individuals) over a period of 4 years (1995–1999). This is probably a result of saturation of the habitat by B. pharaonis recruits originating from the established populations on platforms. Salinity changes and a potential genetic shift may also have contributed to the invasive mussel outbreak.     

Management-free techniques for restoration of Eisenia and Ecklonia beds along the central Pacific coast of Japan

Because of the importance of seaweed resources for conservation of thecoastal environment, the provision of habitats of commercially importantspecies and because of use as human food, a study was carried out on bedsof Eisenia bicyclis and Ecklonia cava. This included the mostimportant environmental factors and the development of techniques forrestoration and long-term maintenance of the beds. At a site chosen fordetailed study, the Eisenia and Ecklonia bed was influenced bymany factors, including desiccation, rainfall, low salinity, low light intensity,turbidity, accumulation of suspended sediments, cover by adhering animals,wave and current action and grazing pressure. The distribution of suchseaweed beds is also limited locally by the size and hardness of rock on thebottom, and by fluctuations in the sand level at the base of the reefs.The management-free technique of creating Eisenia and Ecklonia beds is based on an understanding of the factors limiting the localdistribution of the beds. Conditions for seaweed growth can then beartificially enhanced by coastal engineering. Suggestions to enhance Eisenia and Ecklonia bed restoration using management-freetechniques include raising the bottom, elevating substrata above the sandybottom, and providing substrata of a shape suitable for the attachment ofkelp.     

Patterns of settlement within a restored mussel bed site

The failure of depleted bivalve populations to recover naturally is often due to a breakdown in recruitment processes, such as a lack of larvae or suitable settlement substrate. The identification of these recruitment limitations are a critical initial step for determining an appropriate strategy for restoration. A lack of larval supply and settlement substrate have both been suggested as possible causes of the failure of natural recovery in the green‐lipped mussel population in the Hauraki Gulf, New Zealand, and were therefore examined in this study. Larval mussels settled throughout most of the year directly onto artificial collectors placed within restored adult mussel beds and in the immediate vicinity of the beds at comparatively lower levels than for remnant populations throughout New Zealand. This low settlement coupled with a lack of recruitment to the restored mussel beds suggests larval supply may be limited in the Hauraki Gulf and warrants further examinations into larval dispersal and retention. Larval settlement was also higher on collectors within mussel beds, suggesting the presence of adults enhances larval settlement and highlights the importance of transplanting adult mussels for reestablishing mussel populations.     

Bed-building in captive chimpanzees (Pan troglodytes): the importance of early rearing

In the wild, great apes sleep in beds they make by successively bending branches into an interwoven platform. These beds are functionally more closely related to human beds than they are to the nests and tree-holes used by other primate species. The idea that bed-building by great apes is learned behavior that is dependent on appropriate early experiences has been proposed but never fully tested. In the present study this hypothesis was indirectly tested in 73 captive adult chimpanzees (Pan troglodytes; 27 wild-born and 46 captive-born). Bed-building and use were tested experimentally by the introduction of three sets of bedding materials. Over 200 hr of scan-sampling data were collected during 45-min observations following introduction of the bedding material. The wild-born subjects built and used beds significantly more often than the captive-born subjects. Also, wild-born subjects used more complex techniques during construction. Captive-born subjects that were mother-reared through early adolescence spent more time building and using beds than their nursery-reared counterparts. These differences remained consistent even when previous adult experience with bedding (measured as naturalistic vs. traditional housing) was accounted for. These results suggest that bed-building is a learned behavior that requires early experience and practice for acquisition.     

The role of hoplonemerteans in the ecology of seagrass communities

Seagrasses of the world harbor a rich and varied fauna, but a review of the literature revealed that little has been done to evaluate the ecological importance of nemerteans in such communities. Monostiliferous hoplonemerteans are common inhabitants of some seagrasses, e.g. eelgrass (Zostera), but generally they are seldom collected or identified or are apparently absent in other species such as schoalgrass (Halodule) or turtlegrass (Thalassia). Nineteen species of hoplonemerteans (four families) have been identified from eelgrass beds around the world; they exist mainly as epifauna, and all except two species are probably suctorial feeders. Some palaeonemerteans (2 species) and heteronemerteans (4 species) are also associated with eelgrass, but mainly as infauna. Suctorial nemerteans (4 species in 3 families) from eelgrass beds located along the mid-Atlantic coast of the United States feed in the laboratory on a variety of amphipod species that inhabit eelgrass. Tubicolous species (e.g. Corophium) seem to be preferred. Zygonemertes virescens feeds on nine species of amphipods belonging to six families, and is the only species to feed on isopods (3 species). Analyses of field studies on the occurrence of hoplonemerteans in eelgrass beds in Virginia and New Jersey, along with available information on the food habits of these worms, were used as a basis for demonstrating their potential importance as predators of peracarids in seagrass systems. More careful methods for collecting and identifying worms, continued studies on food preferences and rates of predation, and emphasis on the population dynamics of worms and prey, are recommended in order to evaluate the role of suctorial hoplonemerteans in the ecology of seagrasses.     

Influence of permanent raised bed planting and residue management on physical and chemical soil quality in rain fed maize/wheat systems

Densely populated, intensively cropped highland areas in the tropics and subtropics are prone to erosion and declining soil fertility, making agriculture unsustainable. Conservation agriculture in its version of permanent raised bed planting with crop residue retention has been proposed as an alternative wheat production system for this agro-ecological zone. A five years field experiment comparing permanent and tilled raised beds with different residue management under rainfed conditions was started at El Batán (Mexico) (2,240 m asl; 19.31°N, 98.50°W; Cumulic Phaeozem) in 1999. The objective of this study was to determine the soil quality status after five years of different management practices. The K concentration was 1.65 times and 1.43 times larger in the 0–5 cm and 5–20 cm profiles, respectively, for permanent raised beds compared to conventionally tilled raised beds. The Na concentration showed the opposite trend. Sodicity was highest for conventionally tilled raised beds and for permanent raised beds it increased with decreasing amounts of residue retained on the surface. Permanent raised beds with full residue retention increased soil organic matter content 1.4 times in the 0–5 cm layer compared to conventionally tilled raised beds with straw incorporated and it increased significantly with increasing amounts of residue retained on the soil surface for permanent raised beds. Soil from permanent raised beds with full residue retention had significantly higher mean weight diameter for wet and dry sieving compared to conventionally tilled raised beds. Permanent raised beds with full residue retention had significantly higher aggregate stability compared to those with residue removal. A lower aggregation resulted in a reduction of infiltration. Principal component analysis (PCA) was performed using these soil physicochemical variables that were significantly influenced by tillage or residue management. The PC1 and PC2 separated the conventionally tilled raised beds from the permanent raised beds and PC3 separated permanent raised beds with at least partial residue retention from permanent raised beds with no residue retention. These clear separations suggest that tillage and residue management have an effect on soil processes. The research indicates that permanent raised bed planting increases the soil quality and can be a sustainable production alternative for the (sub)tropical highlands. Extensive tillage with its associated high costs can be reduced by the use of permanent raised beds while at least partial surface residue retention is needed to insure production sustainability.     

Distribution and Recent Reduction of Gelidium Beds in Toyama Bay, Japan

The distribution and recent reduction of Gelidium beds, i.e. mat-like beds dominated by the agarophyte G. elegans Kützing in Toyama Bay (Sea of Japan), in which 95% of the coastline is protected artificially, are reported. Gelidium beds were common in shallow waters (usually < 10 m deep); most of the large beds (> 1 ha) were restricted to the inner coasts of the bay. In calm and eutrophic areas, however, G. elegans was heavily colonized by epiphytes. In the last decade, two beds were buried in situ and beds in their vicinity were damaged by the stagnation of coastal water and/or sedimentation by silts which accompanied land reclamation. At the other two beds monitored since 1988, Gelidium declined a few times but most prominently in 1998, when episodic long summer rain was recorded. This is the first report, not only on the current status of Gelidium beds other than for the central Pacific Coast of Honshu in Japan, but also concerning reduction of the beds caused by both anthropogenic and natural events.     

Morphological Adaptations of the Mussel Crenomitilus grayanus (Bivalvia) to Attached Life

The morphology of the mussel Crenomytilus grayanus from different biotopes manifesting the adaptation of this mollusk to attached life on the bottom surface was studied. It was established that colonization of rocky coastal areas with active hydrodynamics by mussels is related to retardation of linear growth rates, the formation of convex shells, and vigorous development of retractor muscles of the foot and byssus. An environment protected from the effect of surfs on silted bottoms facilitated the fast growth of mussels, which resulted in the formation of an elegant flat shell. In that biotope mussels are specified by a rather poor development of the muscles responsible for its attachment to the substrate.     

海草床育幼功能及其机理

海草床是近岸海域中生产力极高的生态系统,是许多海洋水生动物的重要育幼场所。从生物幼体的密度、生长率、存活率和生境迁移4个方面阐述海草床育幼功能,并从食源和捕食压力两个方面探讨海草床育幼功能机理。许多生物幼体在海草床都呈现出较高的密度、生长率和存活率,并且在个体发育到一定阶段从海草床向成体栖息环境迁移。丰富的食物来源或较低的捕食压力可能是海草床具有育幼功能的主要原因,但不同的生物幼体对海草床的利用有差异,海草床育幼功能的机理在不同环境条件下也存在差异。提出未来海草床育幼功能的重点研究方向:(1)量化海草床对成体栖息环境贡献量;(2)全球气候变化和人类活动对海草床育幼功能的影响;(3)海草床育幼功能对海草床斑块效应和边缘效应的响应,以期为促进我国海草床育幼研究和海草床生态系统保护提供依据。     

Seasonal changes in abundance and shifts in dominance of life history stages of the carrageenophyte Sarcothalia crispata (Rhodophyta, Gigartinales) in south central Chile

The population dynamics of the carrageenophyte Sarcothalia crispatais described from subtidal beds at two localities in south-central Chile. Seasonal fluctuations in total density and biomass were not evident. Frondswere identified to phase by the presence of reproductive structures and theresorcinol reaction. The monthly changes in abundance of each kind offrond were determined. Permanent gametophytic or sporophyticdominance was not evident: the more exposed site showed a seasonal shiftfrom sporophytic dominance in summer to gametophytic dominance inwinter, whereas the more protected site showed an interannual shift fromgametophytic to sporophytic dominance. The differences between localitiesand years suggest a very local population dynamics with large contributionof self-seeding to the maintenance of the S. crispata beds.     

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     

Experimental confirmation of multiple community states in a marine ecosystem

Small changes in environmental conditions can unexpectedly tip an ecosystem from one community type to another, and these often irreversible shifts have been observed in semi-arid grasslands, freshwater lakes and ponds, coral reefs, and kelp forests. A commonly accepted explanation is that these ecosystems contain multiple stable points, but experimental tests confirming multiple stable states have proven elusive. Here we present a novel approach and show that mussel beds and rockweed stands are multiple stable states on intertidal shores in the Gulf of Maine, USA. Using broad-scale observational data and long-term data from experimental clearings, we show that the removal of rockweed by winter ice scour can tip persistent rockweed stands to mussel beds. The observational data were analyzed with Anderson’s discriminant analysis of principal coordinates, which provided an objective function to separate mussel beds from rockweed stands. The function was then applied to 55 experimental plots, which had been established in rockweed stands in 1996. Based on 2005 data, all uncleared controls and all but one of the small clearings were classified as rockweed stands; 37% of the large clearings were classified as mussel beds. Our results address the establishment of mussels versus rockweeds and complement rather than refute the current paradigm that mussel beds and rockweed stands, once established, are maintained by site-specific differences in strong consumer control. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The tip of the tail: molecular identification of seahorses for sale in apothecary shops and curio stores in California

Millions of seahorses a year are traded internationally, with many eventually sold at retail shops as curios, as aquarium pets, and especially for use in traditional medicine. The entire genus is now protected by CITES, but conservation measures have been limited in part by difficulties with species identification and incomplete understanding of trade patterns. In this study, we use molecular techniques to make species- and population-level identifications of 56 seahorses sampled from both traditional medicine and curio shops in San Francisco and central California. Seahorses from medicinal sources included unexpectedly large numbers of the eastern Pacific Hippocampus ingens, a species poorly protected by current CITES export recommendations. Curio shops were dominated by H. barbouri, a spiny species that has been reportedly confused in trading records as H. histrix. Specific populations of origin could be inferred for several species through comparison with publicly available phylogeographic data. Our results underscore the strengths and weaknesses of current recommended export regulations, and we suggest that molecular forensics can help in verifying trade documentation and developing more effective conservation measures.     

Community dynamics of intertidal soft-bottom mussel beds over two decades

Macrozoobenthos communities in the North Sea showed pronounced changes over the past decade in relation to an increasing number of invasive species and climate change. We analysed data sets spanning 22 years on abundance, biomass and species composition of intertidal soft bottom mussel beds near the island of Sylt (German Bight) in the Northern Wadden Sea, based on surveys from 1983/1984, 1990, 1993 and from 1999 to 2005. Mussel bed area and blue mussel biomass decreased, and a change in the dominance structure in the associated community comparing 1984 to mid-1990s with the period from 1999 to 2005 was observed. Coverage of the mussel beds with the algae Fucus vesiculosus decreased since the end of the 1990s. Within the study period biomass and densities of the associated community increased significantly. Dominance structure changed mainly because of increasing abundances of associated epibenthic taxa. Apart from the Pacific oyster Crassostrea gigas all other alien species were already present in the area during the study period. Community changes already started before Pacific oysters became abundant. An attempt is made to evaluate effects on the observed changes of decreasing mussel biomass, ageing of mussel beds, decreasing fucoid coverage and increasing abundances of invader. All four factors are assumed to contribute to changing community structure of intertidal mussel beds.