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     

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.     

Distribution,life history,and production of three species of <Emphasis Type="Italic">Neomysis</Emphasis> in Akkeshi-ko estuary,northern Japan

Spatial and seasonal distribution pattern, life history and production of three species of Neomysis (Mysidacea) which commonly occur in northwestern subarctic Pacific coastal waters, were investigated throughout the year in the Akkeshi-ko estuary, northern Japan. The most abundant species Neomysis awatschensis (annual mean density: 179.8 inds. m⁻², biomass: 108.8 mg DW m⁻²) occurred at the inner part of the estuary including low salinity areas with no clear preference for the seagrass bed. The second most abundant Neomysis mirabilis (mean density: 95.8 inds. m⁻², biomass: 90.1 mg DW m⁻²) occurred at relatively saline seagrass site throughout the year. Occurrence of Neomysis czerniawskii in the estuary was limited to the seagrass bed during summer when their population mainly consisted of juveniles, suggesting that this species is a seasonal migrant between the estuary and the marine environment. Both N. awatschensis and N. mirabilis populations were composed of two generation types, a larger sized overwintering and smaller sized spring/summer generations; however, each species had a different reproductive strategy. N. awatschensis was characterized by fast growth to maturity at a smaller size than N. mirabilis with a relatively high fecundity during warm season, suggesting that this species is an r-strategist which can utilize opportunistically a wide variety of habitats. In contrast, the seagrass bed resident N. mirabilis was a K-strategist which matures at a larger size producing fewer but larger offspring. The annual production of N. awatschensis (0.57–0.70 g DW m⁻², mean of the whole estuary) and N. mirabilis (0.58–0.68 g DW m⁻², mean of the seagrass bed) at their respective habitats was comparable. Consequently, species-specific life history and distribution pattern are concluded to allow Neomysis spp. to coexist in the estuary and the high carrying capacity of seagrass bed is suggested to contribute to maintain their high biomass level.     

Clonal extent,apical dominance and networking features in the phalanx angiosperm <Emphasis Type="Italic">Zostera noltii</Emphasis> Hornem.

Disaggregating seagrass meadows and studying its components separately (clones, ramets, shoots) can provide us insights on meadow dynamics and growth patterns. The clonal growth, dependent upon clonal rules may regulate and impose constraints to plant architecture and, therefore, determine how individual clones evolve into the environment. In order to investigate the relationship between clonal growth rules and clone architecture, the belowground network architecture of single-clones of the seagrass Zostera noltii was studied. Networks were traced in situ after washing out the overlying sediment, and network characteristics were measured using digital analysis: area covered by clone, total rhizome length, type of rhizomatic axes (main, secondary, tertiary, quaternary), number and length of the internodes, branching angles and branching frequencies. This approach revealed that Z. noltii is able to develop into large clones integrating up to 300 internodes, 676 cm of rhizome, 208 shoots and 4,300 cm² of plant area. Internodal length depended on both, the distance to the apical shoot (time effect) and the axes type (apical dominance effect). However, average branching angle was independent of axis type (average 58.3 ± 0.75), but varied significantly depending on the distance from the apical shoot. This average branching angle allows Z. noltii maximize the rate of centrifugal expansion, maintaining a high density in colonized areas to produce close stands but also minimizing the investment in belowground biomass and ramets overlapping. The clonal architecture of Z. noltii seems to be regulated by the interaction of both, apical dominance strength and clonal integration distance. Moreover, clonal growth rules and growth pattern seem to constrain clonality through (clonal) plant architecture regulations (i.e. branching is restricted in secondary axes, similar average branching angles regardless the axes, the higher the distance to the apex the higher the number of internodes in secondary axes, shorter internodes in secondary and tertiary axes). Future research efforts should focus on how these complex relationships between apical dominance and clonal integration interact to elucidate the temporal (seasonal) and spatial scales of both processes and the outcome at the plant architectural level.     

Testing the Stochastic Dynamic Methodology (StDM) as a management tool in a shallow temperate estuary of south Europe (Mondego, Portugal)

A long-term monitoring program has been carried out since the early 1990s in the Mondego estuary, on Portugal's west coast, which is presently under heavy human pressure. In this shallow warm-temperate estuary, a significant macroalgal proliferation has been observed, which is a clear sign of nutrient enrichment. As a result of competition with algae, the extension of the seagrass meadows (mainly Zostera noltii) has been reduced. The present paper examined the applicability of a holistic Stochastic Dynamic Methodology (StDM) in predicting the tendencies of trophic key-components (macrophytes, macroalgae, benthic macroinvertebrate and wading birds) as a response to the changes in estuarine environmental conditions. The StDM is a sequential modelling process developed in order to predict the ecological status of changed ecosystems, from which management strategies can be designed. The data used in the dynamic model construction included true gradients of environmental changes and was sampled from January 1993 to September 1995 and from December 1998 to December 2005. The dynamic model developed was preceded by a conventional multivariate statistical procedure performed to discriminate the significant relationships between the selected ecological components. The model validation was based on independent data collected from January 1996 to January 1997 and from February 1999 to April 2000 for all the state variables considered. Overall, the simulation results are encouraging since they seem to demonstrate the StDM reliability in capturing the trophic dynamics of the studied estuary, by predicting the behavioural pattern for the most part of the components selected, with a focus on the Zostera noltii meadows recovery after the implementation of important management measures.     

Food sources used by sediment meiofauna in an intertidal Zostera noltii seagrass bed: a seasonal stable isotope study

In an intertidal Zostera noltii Hornem seagrass bed, food sources used by sediment meiofauna were determined seasonally by comparing stable isotope signatures (δ¹³C, δ¹⁵N) of sources with those of nematodes and copepods. Proportions of different carbon sources used by consumers were estimated using the SIAR mixing model on δ¹³C values. Contrary to δ¹⁵N values, food source mean δ¹³C values encompassed a large range, from −22.1 ‰ (suspended particulate organic matter) to −10.0 ‰ (Z. noltii roots). δ¹³C values of copepods (from −22.3 to −12.3 ‰) showed that they use many food sources (benthic and phytoplanktonic microalgae, Z. noltii matter). Nematode δ¹³C values ranged from −14.6 to −11.4 ‰, indicating a strong role of microphytobenthos and/or Z. noltii matter as carbon sources. The difference of food source uses between copepods and nematodes is discussed in light of source accessibility and availability.     

Crustacean epifauna of seagrass and macroalgae in Apalachee Bay,Florida, USA

Epifaunal crustaceans on turtlegrass (Thalassia testudinum) and five dominant macroalgae (Anadyomene stellata, Digenia simplex, Halimeda incrassata, Laurencia poitei and Penicillus lamourouxii) were quantitatively sampled bimonthly over a one-year period from September 1979 to September 1980 in a subtropical seagrass meadow in Apalachee Bay, Florida (northeastern Gulf of Mexico). These plant species exhibited a wide range of morphologies, with surface area-to-biomass ratios differing by over 2.5 times. A similar suite of crustaceans occurred on all macrophytes despite differences in shape or architecture among plant species. Relative abundances of many crustaceans, however, varied among plant hosts. Similarity analysis indicated that the epifaunal associates of T. testudinum were distinct from those of the macroalgae. Species richness was generally higher on turtlegrass than on any of the macroalgae. Abundances of total crustaceans per plant biomass or per plant surface area, on the other hand, were greater on all macroalgal species compared to the seagrass. Abundances (per plant biomass or plant surface area) of 14 of the 16 numerically dominant epifaunal species differed significantly among macrophytes. Twelve of the 16 species had greater abundance on one or more macroalgae, while only two species were more abundant on T. testudinum. Almost half of the dominant species had greatest abundances on the branching red alga L. poitei. Although abundances per plant biomass and plant surface area were greater on macroalgae relative to turtlegrass, densities (individuals per meter square of bottom) of animals associated with T. testudinum were significantly greater than those associated with macroalgae, primarily because of the greater abundance of turtlegrass in the grass bed. Both surface area-to-biomass ratios and degree of branching were poorly correlated with epifaunal abundance and number of species. Neither structural feature is an adequate predictor of faunal abundance and species richness among plant species, especially when macrophytes with very different morphologies are compared.     

Short- and long-term effects of eutrophication on the secondary production of an intertidal macrobenthic community

Secondary production of a macrobenthic community at an intertidal mudflat was estimated for 33 successive months. Sampling was carried out along a eutrophication gradient, including non-eutrophied Zostera meadows, an intermediate muddy area, and a strongly eutrophied sand-muddy flat, where macroalgal blooms of Enteromorpha spp. usually occur. The Zostera meadows were always the most productive habitat (145–225 g ash-free dry weight m^–2 year^–1). In the short term, the macroalgal bloom benefited the total estuarine production by enhancing the annual production in the eutrophied area. Nevertheless, our results show that this increase was short lived and in no way sufficient to match the production in the Zostera meadows. In the long term, the present study provides evidence that the disappearance of macrophyte beds, as a result of ongoing eutrophication, constitutes a major threat to the sustainability of the estuarine ecosystem.Communicated by S.A. Poulet, Roscoff     

Modelling the effects of green macroalgae blooms on the population dynamics of Cyathura carinata (Crustacea: Isopoda) in an eutrophied estuary

A population dynamics model was developed to simulate the effects of benthic macroalgae blooms (mostly Enteromorpha spp.) on the productivity of Cyathura carinata (Crustacea: Isopoda), a possible keystone species in the benthic communities of the Mondego estuary. The model describes C. carinata population dynamics, as well as the relationships between Enteromorpha biomass, Enteromorpha decaying rates, organic matter content in the sediments and detritus consumption by C. carinata, a detritic feeder. Model results support the idea that seasonal blooms of Enteromorpha determine a significant increase of organic matter content in the sediments, due to macroalgae decay, which initially contributes to enhance C. carinata consumption and growth rates, determining a significant increase in the biomass. Nevertheless, later, following the algae bloom, C. carinata biomass decreases, and reaches its lowest value, close to 0, when the algae crash. This effect is probably related with strong anoxic conditions, especially during night, due to high algal decomposition rates. In accordance with the model, migration of new individuals from adjacent areas must occur in order to recolonise the area affected by the algae bloom. Therefore, it seems reasonable to conclude that macroalgae blooms that are limited in space may favour C. carinata populations, but extensive blooms affecting the whole area of distribution of this species will determine its disappearance.     

Seasonal variations in nitrogen-fixation (acetylene reduction) and sulphate-reduction rates in the rhizosphere of Zostera noltii: nitrogen fixation by sulphate-reducing bacteria

Nitrogen-fixation (acetylene reduction) rates were measured over an annual cycle in meadows of the seagrass Zostera noltii Hornem in the Bassin d'Arcachon, south-west France, between March 1994 and February 1995, using both slurry and whole-core techniques. Measured rates using the slurry technique consistently overestimated those determined on whole cores, probably due to the release of labile organic carbon sources as a result of root damage during preparation of the slurries. Thus, the whole-core technique may provide a more accurate estimate of in situ activity, since disturbance of physicochemical gradients of oxygen, sulphide, nutrients and the relationship between the plant roots and the rhizosphere microflora is minimised. Rates measured by the whole-core method were 1.8- to 4-fold greater (dependent upon season) in the light than those measured during dark incubations, indicating that organic carbon diffusing from the plant roots during photosynthesis was an important factor in regulating nitrogen fixation in the rhizosphere. Additions of sodium molybdate, a specific inhibitor of sulphate-reducing bacteria (SRB) inhibited acetylene-reduction activity by >80% as measured by both the slurry and whole-core techniques throughout the year, inferring that SRB were the dominant component of the nitrogen-fixing microflora. A mutualistic relationship between Z. noltii and nitrogen-fixing SRB in the rhizosphere, based on the exchange of organic carbon and fixed nitrogen is proposed. Acetylene- and sulphate-reduction rates showed distinct summer peaks which correlated with a reduced availability of ammonium in the sediment and the annual growth cycle of Z. noltii in the basin. Overall, these data indicate that acetylene reduction (nitrogen fixation) activity in the rhizosphere of Z. noltii was regulated both by the availability of organic carbon from the plant roots and maintenance of a low NH ₄ ⁺ concentration in the vicinity of the plant roots due to efficient assimilation of NH ₄ ⁺ by Z. noltii during the growth season. Nitrogenfixation rates determined from acetylene-reduction rates measured using the whole-core technique ranged from 0.1 to 7.3 mg N m^-2d^-1, depending on season, and were calculated to contribute between 0.4 and 1.1 g N m^-2yr^-1, or 6.3 to 12% of the annual fixed nitrogen requirement of Z. noltii.     

Feeding ecology of the grooved tiger shrimp <Emphasis Type="Italic">Penaeus semisulcatus</Emphasis> De Haan (Decapoda: Penaeidae) in inshore waters of Qatar,Arabian Gulf

The feeding ecology of the green tiger shrimp Penaeus semisulcatus was studied in inshore fishing grounds off Doha, Qatar, using a combination of stable isotope (δ¹³C and δ¹⁵N) analysis and gut contents examination. Samples of post-larvae, juvenile and adult shrimp and other organisms were collected from intertidal and subtidal zones during the spawning season (January–June). Shrimp collected from shallow water seagrass beds were mostly post-larvae and juveniles and were significantly smaller than the older juveniles and adults caught in deeper macroalgal beds. Gut content examination indicated that post-larvae and juvenile shrimp in seagrass beds fed mainly on benthos such as Foraminifera, polychaetes, benthic diatoms and small benthic crustaceans (amphipods, isopods and ostracoda), whereas larger shrimp in the macroalgal beds fed mainly on bivalve molluscs and to a lesser extent polychaetes. In shrimp from both seagrass and algal beds, unidentifiable detritus was also present in the gut (18, 32%). δ¹³C values for shrimp muscle tissue ranged from −9.5 ± 0.26 to −12.7 ± 0.05‰, and δ¹⁵N values increased with increasing shrimp size, ranging from 4.1 ± 0.03 to 7.7 ± 0.11‰. Both δ¹⁵N values and δ¹³C values for shrimp tissue were consistent with the dietary sources indicated by gut contents and the δ¹³C and δ¹⁵N values for primary producers and prey species. The combination of gut content and stable isotope data demonstrates that seagrass beds are important habitats for post-larvae and juvenile P. semisulcatus, while the transition to deeper water habitats in older shrimp involves a change in diet and source of carbon and nitrogen that is reflected in shrimp tissue stable isotope ratios. The results of the study confirm the linkage between sensitive shallow water habitats and the key life stages of an important commercially-exploited species and indicate the need for suitable assessment of the potential indirect impacts of coastal developments involving dredging and land reclamation.     

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.     

The role of the amphipod <Emphasis Type="Italic">Gammarus locusta</Emphasis> as a grazer on macroalgae in Swedish seagrass meadows

Mesograzers are thought to play a critical role in seagrass beds by preventing overgrowth of ephemeral algae. On the Swedish west coast, eelgrass Zostera marina has decreased in recent decades as a result of eutrophication and increased growth of macroalgal mats (mainly filamentous Ulva spp. and Ectocarpales), with no indication of grazer control of the algae. The aim of this study was to investigate the ability of the amphipod Gammarus locusta to control algal blooms during nutrient-enriched and ambient conditions, using a combination of laboratory, field and model studies. Laboratory experiments demonstrated that juvenile and adult G. locusta could consume both Ulva spp. and Ectocarpales, but that consumption of Ulva spp. was significantly higher. Cannibalism was common in individual treatments involving multiple size-classes of G. locusta, but only large, male gammarids consumed smaller juveniles in the presence of Ulva spp. as an alternative food source. However, no negative effects of cannibalism were found on total grazing impact. A model using size-specific grazing rates and growth rates of G. locusta and of Ulva spp. suggests that approximately 62 young juvenile, or 27 adult G. locusta are needed per gram DW of Ulva spp. to control the algal growth during ambient nutrient conditions, and approximately 2.6 times as many gammarids during enhanced nutrient conditions. On the Swedish west coast, densities and mean sizes of G. locusta in eelgrass beds are below these critical values, suggesting that the gammarids will not be able to control the growth of the filamentous macroalgae. However, in the field cage experiment, immigration of juveniles and reproduction of encaged adult G. locusta resulted in unexpectedly high densities of G. locusta (>4,000 individual m⁻²), and very low biomass of Ulva spp. in both ambient and nutrient-enriched treatments. Although the high numbers of juveniles in all cages precluded any significant treatment effects, this suggests that in the absent of predators, the population of G. locusta can grow significantly and control the biomass of Ulva spp. Furthermore, low grazing of Ectocarpales in the laboratory and high biomass of these filamentous brown algae in the field indicate a preference for the more palatable green algae Ulva spp. This study indicates that the high grazing capacity of G. locusta, in combination with high reproduction and growth rates, would allow the amphipod to play a key role in Z. marina ecosystems by controlling destructive blooms of filamentous green algae. However, high predation pressure appears to prevent large populations of G. locusta in eelgrass beds on the Swedish west coast today.     

Modelling seasonal dynamics of biomasses and nitrogen contents in a seagrass meadow (Zostera noltii Hornem.): application to the Thau lagoon (French Mediterranean coast)

Anumerical deterministic model for a seagrass ecosystem (Zostera noltii meadows) has been developed for the Thau lagoon. It involves both above- and belowground seagrass biomasses, nitrogen quotas and epiphytes. Driving variables are light intensity, wind speed, rain data and water temperature. This seagrass model has been coupled to another biological model in order to simulate the relative contributions of each primary producer to: (i) the total ecosystem production, (ii) the impact on inorganic nitrogen and (iii) the fluxes towards the detritus compartment. As a first step in the modelling of seagrass beds in the Thau lagoon, the model has a vertical structure based on four boxes (a water box on top of three sediment boxes) and the horizontal variability is neglected until now. This simple box structure is nevertheless representative for the shallow depth Z. noltii meadows, spread over large areas at the lagoon periphery.After calibration, simulation results have been compared with in situ measurements and have shown that the model is able to reproduce the general pattern of biomasses and nitrogen contents seasonal dynamics. Moreover, results show that, in such shallow ecosystems, seagrasses remain the most productive compartment when compared with epiphytes or phytoplankton productions, and that seagrasses, probably due to their ability in taking nutrients in the sediment, have a lower impact on nutrient concentration in the water column than the phytoplankton. Furthermore, in spite of active mechanisms of internal nitrogen redistribution and reclamation, the occurrence of a nitrogen limitation of the seagrass growth during summer, already mentioned in the literature, have also been pointed out by the model. Finally, simulations seems to point out that epiphytes and phytoplankton could compete for nitrogen in the water column, while a competition for light resources seems to be more likely between epiphytes and seagrasses.     

Geographic and ontogenetic variation in the diet and daily ration of the bonnethead shark, <Emphasis Type="Italic">Sphyrna tiburo</Emphasis>, from the eastern Gulf of Mexico

To examine variation in diet and daily ration of the bonnethead shark, Sphyrna tiburo (Linnaeus 1758), animals were collected from three areas in the eastern Gulf of Mexico: northwest Florida (∼29°40′N, 85°13′W), Tampa Bay near Anclote Key (∼28°10′N, 82°42.5′W), and Florida Bay (∼24°50′N, 80°48′W) from March through September, 1998–2000. In each area, diet was assessed by life stage (young-of-the year, juveniles, and adults) and quantified using five indices: percent by number (%N), percent by weight (%W), frequency of occurrence (%O), index of relative importance expressed on a percent basis (%IRI), and %IRI based on diet category (%IRI_DC). Diet could not be assessed for young-of-the-year in Tampa Bay or Florida Bay owing to low sample size. Diet analysis showed an ontogenetic shift in northwest Florida. Young-of-the-year stomachs from northwest Florida (n = 68, 1 empty) contained a mix of seagrass and crustaceans while juvenile stomachs (n = 82, 0 empty) contained a mix of crabs and seagrass and adult stomachs (n = 39, 1 empty) contained almost exclusively crabs. Crabs made up the majority of both juvenile and adult diet in Tampa Bay (n = 79, 2 empty, and n = 88, 1 empty, respectively). Juvenile stomachs from Florida Bay (n = 72, 0 empty) contained seagrass and a mix of crustaceans while adult stomachs contained more shrimp and cephalopods (n = 82, 3 empty). Diets in northwest Florida and Tampa Bay were similar. The diet in Florida Bay was different from those in the other two areas, consisting of fewer crabs and more cephalopods and lobsters. Plant material was found in large quantities in all stomachs examined from all locations (>15 %IRI_DC in 6 of the 7 life stage-area combinations, >30 %IRI_DC in 4 of the 7 combinations, and 62 %IRI_DC in young-of-the-year diet in northwest Florida). Using species- and area-specific inputs, a bioenergetic model was constructed to estimate daily ration. Models were constructed under two scenarios: assuming plant material was and was not part of the diet. Overall, daily ration was significantly different by sex, life stage, and region. The bioenergetic model predicted increasing daily ration with decreasing latitude and decreasing daily ration with ontogeny regardless of the inclusion or exclusion of plant material. These results provide evidence that bonnetheads continuously exposed to warmer temperatures have elevated metabolism and require additional energy consumption to maintain growth and reproduction. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Stable C and N isotope analysis elucidated the importance of zooplankton in a tropical seagrass bed of Santiago Island,Northwestern Philippines

The role of zooplankton in a tropical seagrass ecosystem was investigated in milkfish farms pollution-impacted and -unimpacted seagrass beds in Santiago Island coral reefs, Northwestern Philippines. The aim was to compare between the two sites: (1) abiotic factors and zooplankton community parameters, and (2) the trophic structure using C and N stable isotopes. Low water (98–119?mV) and sediment (–121 to ?138?mV) Oxidation Reduction Potential values indicated a reducing environment in the impacted site. Zooplankton in the impacted site showed the typical community response to eutrophication (low diversity, but high total abundance due to the dominance of the cyclopoid copepod Oithona oculata), generally few elevated δ¹⁵N values, but a significant shift towards depleted ¹³C due to the organic enrichment of fish-farm feeds. Apart from suggesting a highly complex food web with POM and zooplankton as main food sources in the unimpacted site, the Bayesian mixing model simulation generated reduced complexity in feeding interactions between basal sources, zooplankton, and fish including adults of a key fish species, Siganus fuscescens, in the impacted sites. In this study, C and N stable isotope analysis has clarified the importance of zooplankton as fish prey in a seagrass bed food web.     

Food web analysis in intertidal <Emphasis Type="Italic">Zostera marina</Emphasis> and <Emphasis Type="Italic">Zostera noltii</Emphasis> communities in winter and summer

The food web of two intertidal seagrass (Zostera marina and Zostera noltii) beds that may be influenced by the seasonal variation in food source abundance was studied in winter and in summer with δ¹³C and δ¹⁵N analysis. In spite of high relative variation of abundance of main primary producers at the two sites, the food web did not vary between winter and summer. The δ¹³C range of primary producers was wide. Zostera leaves, the most ¹³C-enriched source, were not consumed directly by grazers. Deposit and filter feeders have a similar δ¹³C and could use a mix of suspended and sedimented organic particulate matter, largely composed of detritus from macroalgae to seagrass. This trophic pathway allows the local incorporation of the high biomass produced by seagrasses. The wide δ¹⁵N range of predators was linked either to a large variety from omnivore to carnivore predators or to the also wide ranges of δ¹⁵N of primary consumers.     

Fish assemblages in seagrass beds are influenced by the proximity of mangrove forests

Mangrove forests and seagrass beds frequently occur as adjacent habitats in the temperate waters of southeastern Australia. At low tide when fish cannot occupy mangroves they might utilise adjacent habitats, including seagrass. We first sampled small fish from seagrass beds close to and far from mangroves in the Pittwater estuary, NSW, Australia. Seagrass beds close to mangroves had a greater density of fish species than beds far from mangroves (close: mean 16.0 species net⁻¹, SE 1.0; far: 13.2, 1.3; P < 0.05). In particular, juvenile fish were in greater densities near to than far from mangroves (close: 5.3, 0.4; far: 3.1, 0.4; P < 0.05). We then sampled the mangrove forests during the high tide and seagrass beds during the low tide, in beds along a continuum of distances from mangroves. Multivariate analysis showed that fish assemblages differed with distance from mangroves, and the differences were attributed to the composition of the fish assemblage (i.e. presence/absence of fish species), not the abundances of individual species. In particular, fish that utilise mangrove forests at high tide were found in greater species densities and species richness in seagrass nearer to mangroves. A negative relationship was found between the density of mangrove-utilising fish species and the distance of the bed from mangroves (R ² = 0.37, P < 0.05). This confirms the important connectivity between mangroves and seagrass for fish in temperate Australian waters.     

Comparison of mussel bed community between two intertidal mytilids Septifier virgatus and Hormomya mutabilis

Mussel bed community structure of two intertidal mytilids, Septifer virgatus (Wiegmann) and Hormomya mutabilis (Gould), whose beds were contiguous vertically on a rocky intertidal shore (Wakayama Prefecture, Japan), was compared between 1982 and 1983. In the upper S. virgatus bed, crustaceans and bivalves were dominant in terms of both number of individuals and biomass. There were three barnacle species representative of the epizoans, two isopods and one amphipod as mobile fauna, and two bivalves as infauna. The lower H. mutabilis bed supported virtually no epizoans or mobile fauna. Infaunal free-ranging polychaetes and sipunculids were dominant in terms of both number of individuals and biomass. The H. mutabilis bed contained a much greater amount of sediment than did the S. virgatus bed, and the interstices among individual H. mutabilis and among their byssal threads were filled with sediment. The biomass of six of nine species dominant in the S. virgatus bed was negatively correlated with the amount of sediment. Recruitment of these faunal assemblages into artificial mussel clumps was examined in mussel enclosure experiments, and a negative sediment effect in H. mutabilis clumps was detected for one isopod and one limpet species. Factors causing the differences between two mussel bed communities are discussed, focusing on the effects of sediment.     

Eutrophication and trophic structure in response to the presence of the eelgrass <Emphasis Type="Italic">Zostera noltii</Emphasis>

In estuaries, eelgrass meadows contribute to fundamental ecosystem functions of estuaries, providing food to several predators and buffering the negative effects of eutrophication. We asked whether the presence of the eelgrass Zostera noltii decreased the nitrogen concentration in the overlying water, affected the sources of nitrogen sequestrated by primary producers and changed the benthic and pelagic food web structures. We also studied the importance of these food webs in providing food to fish. We compared bare sediment to sediment covered by a Z. noltii meadow, and examined nutrient concentrations in the water column and δ¹⁵N in primary producers as indicators of anthropogenic inputs of nutrients. We then measured both δ¹³C and δ¹⁵N in the tissues of plants and consumers to establish food web structures. There were no differences in the concentrations and sources of nitrogen between sites. Rather, δ¹⁵N values indicated anthropogenic inputs of N (e.g. sewage discharges, agriculture) in both sites. There were no major differences in the structure of the planktonic food web, which was in part sustained by particulate organic matter and supported most predator fish, and in the structure of the benthic food web. Nonetheless, there were differences in the sources of food for omnivore consumers and for the detritivore Scrobicularia plana. Overall, the benthic food web did not use food derived from the eelgrass or macroalgae deposited on the substratum. Suspension feeders used particulate and sediment organic matter, whereas the δ¹³C and δ¹⁵N values of the other consumers indicated a likely contribution of benthic microalgae. Furthermore, in both habitats we found large variability in the isotope signatures of benthic macrofauna consumers, which did not allow distinguishing clearly different trophic groups and indicated a high level of omnivory and a mixed diet opportunistically making use of the availability of food in the surroundings.