Comparison of a current eelgrass disease to the wasting disease in the 1930s

A comparison is made of the wasting disease that struck the whole Atlantic population of Zostera marina L. in the 1930s and a current outbreak of a rather similar disease in Z. marina beds along the north-eastern coasts of the U.S.A. Although the disease phenomena on the plants appear to be very similar, disease-related declines of Z. marina are at present still very local. In Europe, diseased plants have been found, but no declines have been observed.

The wasting disease in the 1930s was not investigated before the epidemic reached a devastating stage. Present observations may indicate that a new widespread die-off may be developing. In order to facilitate the study of the current epidemic, a scenario of disease and related decline, with several variants, has been elaborated, based on the existing knowledge of the epidemic of the 1930s, but also clearly showing the gaps in this knowledge.     

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.     

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     

From introduced species to invader: what determines variation in the success ofCodium fragile ssp.tomentosoides (Chlorophyta) in the North Atlantic Ocean?

The green algaCodium fragile ssp.tomentosoides (Chlorophyta) has been introduced accidentally and successfully from Japan to many shores of the northern and southern hemispheres, including those of the Northeast and Northwest Atlantic Ocean. On most European coasts,Codium occurs regularly but at low abundances in the intertidal zone and is absent from subtidal habitats. In contrast,Codium is extremely abundant in subtidal kelp beds in the Northwest Atlantic Ocean where it often reaches nuisance proportions. This differential success cannot be accounted for by either the properties of the invader or by physico-chemical differences between invaded coasts. A theoretical comparison between two regions on opposite sides of the Atlantic Ocean, i.e. Eastern Nova Scotia, Canada, and south central Britain, illustrates how the resident benthic community may determine the difference in relative abundance ofCodium in subtidal habitats between northeast America and Europe. In this review, low floral species diversity, biological disturbance and facilitation by a previous species invasion are suggested as potential factors for the establishment, success and abundance ofCodium in the Northwest Atlantic Ocean, but these require testing in field experiments.     

A review of mangrove rehabilitation in the Philippines: successes,failures and future prospects

From half a million hectares at the turn of the century, Philippine mangroves have declined to only 120,000 ha while fish/shrimp culture ponds have increased to 232,000 ha. Mangrove replanting programs have thus been popular, from community initiatives (1930s–1950s) to government-sponsored projects (1970s) to large-scale international development assistance programs (1980s to present). Planting costs escalated from less than US$100 to over $500/ha, with half of the latter amount allocated to administration, supervision and project management. Despite heavy funds for massive rehabilitation of mangrove forests over the last two decades, the long-term survival rates of mangroves are generally low at 10–20%. Poor survival can be mainly traced to two factors: inappropriate species and site selection. The favored but unsuitable Rhizophora are planted in sandy substrates of exposed coastlines instead of the natural colonizers Avicennia and Sonneratia. More significantly, planting sites are generally in the lower intertidal to subtidal zones where mangroves do not thrive rather than the optimal middle to upper intertidal levels, for a simple reason. Such ideal sites have long been converted to brackishwater fishponds whereas the former are open access areas with no ownership problems. The issue of pond ownership may be complex and difficult, but such should not outweigh ecological requirements: mangroves should be planted where fishponds are, not on seagrass beds and tidal flats where they never existed. This paper reviews eight mangrove initiatives in the Philippines and evaluates the biophysical and institutional factors behind success or failure. The authors recommend specific protocols (among them pushing for a 4:1 mangrove to pond ratio recommended for a healthy ecosystem) and wider policy directions to make mangrove rehabilitation in the country more effective.     

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.     

A quantitative real-time polymerase chain reaction assay for the seagrass pathogen Labyrinthula zosterae

The protist Labyrinthula zosterae (Phylum Bigyra, sensu Tsui et al. 2009) has been identified as a causative agent of wasting disease in eelgrass (Zostera marina), of which the most intense outbreak led to the destruction of 90% of eelgrass beds in eastern North America and western Europe in the 1930s. Outbreaks still occur today, albeit at a smaller scale. Traditionally, L. zosterae has been quantified by measuring the necrotic area of Z. marina leaf tissue. This indirect method can however only lead to a very rough estimate of pathogen load. Here, we present a quantitative real-time polymerase chain reaction (qPCR) approach to directly detect and quantify L. zosterae in eelgrass tissue. Based on the internal transcribed spacer (ITS) sequences of rRNA genes, species-specific primers were designed. Using our qPCR, we were able to quantify accurately and specifically L. zosterae load both from culture and eelgrass leaves using material from Europe and North America. Our detection limit was less than one L. zosterae cell. Our results demonstrate the potential of this qPCR assay to provide rapid, accurate and sensitive molecular identification and quantification of L. zosterae. In view of declining seagrass populations worldwide, this method will provide a valuable tool for seagrass ecologists and conservation projects.     

Multiple introductions promote range expansion of the mollusc Cyclope neritea (Nassariidae) in France: evidence from mitochondrial sequence data

Since the 1970s, the nassariid gastropod Cyclope neritea has been extending its range north along the French Atlantic coasts from the Iberian Peninsula. This may be due to natural spread because of the recent warming of the northeastern Atlantic. However, human-mediated introductions related to shellfish culture may also be a probable explanation for this sudden range expansion. To examine these two hypotheses, we carried out a comprehensive study based on mitochondrial gene sequences (cytochrome oxidase I) of the five recently colonized French bays as well as 14 populations located in the recognized native range of the species. From a total of 594 individuals, we observed 29 haplotypes to split into three divergent clades. In the native range, we observed a low molecular diversity, strong genetic structure and agreement between geography and gene genealogies. Along the French coasts, we observed the opposite: high genetic diversity and low genetic structure. Our results show that recurrent human-mediated introductions from several geographical areas in the native range may be a source for the French Atlantic populations. However, despite the low dispersal ability of C. neritea, the isolation-by-distance pattern in France suggested that this gastropod may have been present (although unnoticed) on the French Atlantic coasts before the 1970s. As C. neritea shows characteristics of a cryptogenic species, the classification of Atlantic populations as either native or introduced is not straightforward. Cryptogenic species should be studied further to determine the status of new populations close to their recognized native range.     

On the population development of the introduced razor clamEnsis americanus near the island of Sylt (North Sea)

The American razor clamEnsis americanus (=E. directus) was introduced into the eastern North Sea in the late 1970s. By larval and poslarval drifting the species rapidly extended its distribution, now ranging from the English Channel to the Kattegat. Near the island of Sylt in the eastern North Sea it has been recorded since 1979. Recruitment was rather irregular, with about six strong year-classes within two decades. Growth seems comparable with populations in its native range (Atlantic North America). Although present in the lower intertidal zone, maximum densities occurred in shallow subtidal sand with a biomass similar to that of dense beds of native cockles and mussels in the adjacent intertidal zone.Ensis americanus established in otherwise sparsely faunated sand (channels exposed to strong currents) as well as in dense infaunal assemblages (lower intertidal and subtidally). There were no significant interactions with resident species. In dense beds of razor clams, however, fine sediment particles accumulated which may have altered abundances of polychaetes. In spite of high annual variability,E. americanus has become a prominent component of the macrobenthos in shallow subtidal sands of the North Sea.     

Modelling past and present geographical distribution of the marine gastropod Patella rustica as a tool for exploring responses to environmental change

A climate envelope approach was used to model the distributions of the intertidal gastropod Patella rustica , to test the robustness of forecast responses to climate change. The model incorporated variables that were likely to determine the abundance and the northern range limit of this species in the NE Atlantic. The model was built using classification and regression tree analysis (CART) trained with historical distribution data from the mid 1950s and a set of corresponding climatic and oceanographic variables. Results indicated air and sea temperature, in particular during the reproductive and settlement periods, as the main determinants of the Atlantic distribution of P. rustica . The model was subsequently fed with contemporary climatic data and its output was compared with the current distribution and abundance of P. rustica , assessed during a 2002–2003 survey. The model correctly hindcasted the recent collapse of a distributional gap in northern Portugal, as well as an increase in abundance at locations within its range. The predicted northward expansion of the northern range limit did not occur because the absence of the species was confirmed in a survey encompassing the whole Atlantic French coast up to Brest. Stretches of unsuitable habitat too long to be overcome by dispersal are the likely mechanism controlling the northern limit of the distribution of this intertidal species.     

Surface and submarine pollination in the seagrass Zoster a marina L

COX, P. A., LAUSHMAN, R. H. & RUCKLESHAUS, M. H., 1992. Surface and submarine pollination in the seagrass Zostera marina L. Hydrophilous plants can be divided into three ecological categories depending upon whether their pollen is transported above, on, or under the water surface. A mixed mode of submarine and surface hydrophilous pollination occurs in the seagrass Zostera marina L. In the surface mode of pollination, pollen rafts or 'search vehicles' which superficially resemble snowflakes, form at low tide and are transported on the surface of the sea by winds and water currents. Some of the search vehicles collide with the floating female stigmas, effecting pollination. In the submarine mode of pollination, small pollen masses resembling whisk brooms travel beneath the surface of the water. Although we failed to observe a submarine pollination event, SEM analysis of stigmas from subtidal populations confirms that submarine pollination does occur in Z. marina . However, observations of stigmas positioned at and below the surface of the water show surface pollination to be highly efficient. Electrophoretic evaluation of both subtidal and intertidal populations indicates significant genetic variation between populations. Given the high flux rates of surface-borne pollen and pollen viability in excess of 5 h, it is likely that surface-borne pollen is a major source of gene flow in Zostera marina populations.     

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     

The American slipper limpet <Emphasis Type="Italic">Crepidula fornicata </Emphasis>(L.) in the northern Wadden Sea 70 years after its introduction

In 1934 the American slipper limpet Crepidula fornicata (L.) was first recorded in the northern Wadden Sea in the Sylt-R?m? basin, presumably imported with Dutch oysters in the preceding years. The present account is the first investigation of the Crepidula population since its early spread on the former oyster beds was studied in 1948. A field survey in 2000 revealed the greatest abundance of Crepidula in the intertidal/subtidal transition zone on mussel (Mytilus edulis) beds. Here, average abundance and biomass was 141 m^–2 and 30 g organic dry weight per square metre, respectively. On tidal flats with regular and extended periods of emersion as well as in the subtidal with swift currents in the gullies, Crepidula abundance was low. The main substrate of attachment was live mussels. Compared with the years following their initial introduction, Crepidula is more abundant today and has shifted from the now extinct oyster beds to the epifaunal community of the mussel beds. Their present abundance is considerably lower than at more southern European coasts where the species may dominate the epifauna. Low winter temperatures are suggested to have limited the population expansion in the northern Wadden Sea until now. Electronic Publication     

Competition, mortality and diversity in South Atlantic coastal boulder communities

Boulder shores are common at all latitudes and dominate the intertidal and subtidal zones of sub-Antarctic coastlines. The encrusting benthos of boulders was examined on similar shore types at four locations: Tierra del Fuego, East Falkland, West Falkland and Bird Island (off South Georgia). Bird Island is unusual in experiencing high trampling and organic enrichment from fur seals. The results were compared to a Patagonian site and a non-trampled South Georgia site (Husvik) and other sites taken from the literature. Principal Component Analysis revealed South Atlantic/Southern Ocean encrusting faunas formed a distinct cluster when compared to assemblages from elsewhere at similar latitudes. Bray Curtis cluster analysis of the South Atlantic-Southern Ocean sites showed the major division was between Southern Ocean and South Atlantic Ocean sites, beyond which there were three distinct clusters centred around Patagonia (Magellanic), the Falklands and Southern Ocean sites. The organisation of competitive interactions between species was mostly determinate and transitive (essentially hierarchical). The transitivity index scores were higher than most similar assemblages studied to date. The diversity of encrusting assemblages ranged from Shannon Weaver H values of 2.38–0.77 (East Falkland and Bird Island, respectively) in the intertidal to 1.27–0.73 (Patagonia and South Georgia, respectively) in the subtidal zone. Annual mortality (of bryozoan colonies) varied from 85–97% in the intertidal to 65–92% in the subtidal, being higher in the Southern Ocean than South Atlantic sites, largely due to ice scour and wave action. The Bird Island mortality values may be high even for a Southern Ocean site. Accepted: 5 October 2000     

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.     

Sequencing type material resolves the identity and distribution of the generitype Lithophyllum incrustans,and related European species L. hibernicum and L. bathyporum (Corallinales,Rhodophyta)

DNA sequences from type material in the nongeniculate coralline genus Lithophyllum were used to unambiguously link some European species names to field‐collected specimens, thus providing a great advance over morpho‐anatomical identifi‐cation. In particular, sequence comparisons of rbcL, COI and psbA genes from field‐collected specimens allowed the following conclusion: the generitype species, L. incrustans, occurs mostly as subtidal rhodoliths and crusts on both Atlantic and Mediterranean coasts, and not as the common, NE Atlantic, epilithic, intertidal crust reported in the literature. The heterotypic type material of L. hibernicum was narrowed to one rhodolith belonging in Lithophyllum. As well as occurring as a subtidal rhodolith, L. hibernicum is a common, epilithic and epizoic crust in the intertidal zone from Ireland south to Mediterranean France. A set of four features distinguished L. incrustans from L. hibernicum, including epithallial cell diameter, pore canal shape of sporangial conceptacles and sporangium height and diameter. An rbcL sequence of the lectotype of Lithophyllum bathyporum, which was recently proposed to accommodate Atlantic intertidal collections of L. incrustans, corresponded to a distinct taxon hitherto known only from Brittany as the subtidal, bisporangial, lectotype, but also occurs intertidally in Atlantic Spain. Specimens from Ireland and France morpho‐anatomically identified as L. fasciculatum and a specimen from Cornwall likewise identified as L. duckerae were resolved as L. incrustans and L. hibernicum, respectively.     

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.     

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.     

Ecological Wood Anatomy of Avicennia marina

The soil physicochemical indexes including edaphic pH values, edaphic salinity and soil nutrient indexes were determined in 8 Avicennia marina populations located in national nature reserve of Shenzhen of Guangzhou Province. The morphological characters in secondary xylem of Amarina populations corresponding to soil sampling which represented different habitats were surveyed by means of light microscopy and electronic microscopy. Some common specialized wood structures in Amarina growing in different habitais can be observed which suggest they function as safely conducting water under high negative pressure and are thus adaptive to intertidal habitats．These characteristics include the occurrence of: (1) some fibriform vessel elements; (2) vessel vestures and helical structures on vessel walls; (3) parenchyma cells contain lots of starch grains;(4) included phloem. Soil organic matter and soil total salt have significant effect on quantitative wood anatomical features by the stepwise regression analysis.With the increase of soil salinity and soil organic matter content which suggest occurrence of more and more serious “physiological drought”, Amarina is characterized by a higher porosity.and double thickness of vessel wall making contact with an adjacent vessel. The trends inferred that variations in the quantitative wood anatomical features of Amarina growing at different sites are adaptive to heterogenous habitats in the intertidal areas.     

A new species of Euzonus (Polychaeta: Opheliidae) from subtidal zones in Japan

A new opheliid polychaete, Euzonus japonicus sp. nov., is described. This species was collected from subtidal zones in Japanese coasts, while most Euzonus species inhabit intertidal sandy beaches. E. japonicus sp. nov. is morphologically most similar to another subtidal species, E. flabelliferus (Ziegelmeier, 1955) collected from northern Europe, but different from that in the form of branchiae. They share a unique characteristic on setiger 10, i. e., a pair of lateral transverse rows of conical cirri, instead of a pair of lateral smooth ridges that are common to all intertidal Euzonus species.