Ecological energetics of the seaweed zone in a marine bay on the Atlantic coast of Canada. I. Zonation and biomass of seaweeds

An intertidal and underwater survey of the zonation of seaweed in St. Margaret's Bay, NovaScotia, Canada showed 8 major zones as one proceeded away from shore: (1) Fucus and Ascophyllum; (2) Chorda with filamentous browns; (3) Chondrus crispus; (4) Zostera marina; (5) Laminaria digitata with L. longicrusis; (6) Laminaria longicruris; (7) L. longicruris with Agarum cribrosum; (8) Agarum cribrosum with Ptilota serrata. Zostera occurred at the same level as O. crispus but replaced it in sheltered water. Ascophyllum was more abundant in sheltered water. L. digitata was confined to the more exposed, steeper shore. The average horizontal extent of the seaweed zone was 369 m, and the greatest depth of significant amounts of seaweed 20 to 30 m. Laminaria longicruris contributed 36% of the total biomass, and Laminaria spp. and Agarum together constituted 83%, while intertidal seaweeds contributed less than 10% of the biomass. The estimated average total biomass per m of shore line was 1,481 kg fresh weight, 326 kg dry weight, 98 kg carbon, or 980x10³ kcal. When averaged over the whole area of the bay, the corresponding figures were 1.38 kg/m² fresh weight, 0.30 kg/m² dry weight, 91 g/m² carbon or 912 kcal/m². Dry matter of Laminaria was 15 to 27% of fresh weight in blades, 10 to 12% in stipes. The dry matter content of blades was least in spring and highest in autumn, but carbon content and calorific value of dry matter showed little difference with species or season.Contribution to the International Biological Programme CCIBP 108.Bedford Institute Contribution BI 249.     

A comparison of macrofaunal and meiofaunal distribution and standing stock across a rocky shore,with an estimate of their productivities

In 1984 and 1985 algal, macrofaunal and meiofaunal standing stocks were estimated on a exposed rocky shore along the west coast of False Bay, South Africa, using comparable, area-based sampling techniques. The shore supported a rich growth of algae, particularly in summer, when a maximum standing crop of 403 g m^-2 was recorded in the low shore. In winter, the largest component of macrofaunal biomass comprised the filter-feeding barnacle Tetraclita serrata, which attained 75 g m^-2 in the middle balanoid; but as a result of late recruitment and high mortality of this species, the summer shore was dominated by herbivorous gastropods, particularly Patella cochlear, which attained a maximum biomass of 66 g m^-2 on the low shore. Meiofaunal numbers and biomass were closely correlated to the distribution of algal turfs and associated trapped sediments. Numerically, the most important components of the meiofauna were nematodes and copepods, while the biomass was more evenly shared among foraminifera, minute gastropods, copepods and insect larvae. Numbers and biomass peaked in the lower balanoid during winter (1.9×10⁶ individuals, or 8.5 g m^-2). Macrofauna:meiofauna numbers and biomass ratios are presented for each zone and the distribution patterns discussed in relation to the conditons in each. Numerically, meiofauna exceed macrofauna by an overall ratio of 1:391, with values ranging from 1:556 in the lower balanoid to 1:18 in the Littorina zone. Macrofaunal biomass exceeds that of meiofauna in all zones by an overall ratio of 10:1, but values range from 2.1:1 in the upper balanoid to 48:1 in the middle balanoid. By incorporating turnover ratios extrapolated from the literature, mean annual productivity ratios have been calculated. These indicate that macrofauna account for 75% of total secondary production and meiofauna for 25%. Failure to incorporate meiofauna in analyses of energy flow on rocky shore ecosystems would thus lead to considerable errors. The possible trophic role of meiofauna in such systems is discussed.     

Plankton of the fladen ground during FLEX 76 II. Population dynamics and production of Thysanoessa inermis (Crustacea: Euphausiacea)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR) and the Longhurst-Hardy Plankton Recorder (LHPR) during the Fladen Ground Experiment in 1976 (FLEX 76) are used to describe the vertical distribution and population dynamics of Thysanoessa inermis (Krøyer) and to provide estimates of the production and carbon budget of the population from 19th March to 3 June 1976. Spawning occurred in late April and early May, in near synchronisation with the start of the spring bloom of phytoplankton. Eggs, nauplii and calyptopes reached maximum abundance in succession, and furciliae were numerous when sampling ceased in early June. Adults increased in length from a mean of 12.1 mm in mid-March to 17.5 mm in early June and the estimated production was 2.40 mg m^-3 over the 74 d period. Total carbon ingested by the population of T. inermis was estimated to be 10 mg C m^-2 d^-1 in the upper 100m which was only 1.5% of the daily primary production of 0.68 gC m^-2 measured over the FLEX period 26 March to 4 June 1976. The grazing by T. inermis on the phytoplankton population was assumed to have little effect on the control and depletion of the spring phytoplankton bloom during FLEX 77.JOSDAP Contribution No. 50     

Net productivity of two cohorts of Chordaria flagelliformis (Phaeophyta) in Nova Scotia,Canada

Productivities of two cohorts of Chordaria flagelliformis (O. F. Müll.) C. Ag. were estimated from measured changes in biomass and survivorship over time. Maximum productivity during the summer growing season was 2.6 g C m^-2 d^-1. Although this figure is relatively high, the short growing season results in an annual production of only 89 g C m^-2. The significance of primary production by C. flagelliformis lies in its seasonal timing. During the summer growth period, 50% of production was recycled directly by detrital material. During the same time period, productivity and biomass losses of other seaweeds are at their lowest.     

Nutrient cycling between the water column and a marine sediment. I. Organic carbon

The carbon flow through the sediments at a station located in 18.3 m of water off the Scripps Institution of Oceanography, San Diego, California (USA) was determined. The parameters studied [and their mean rates of input (+) or output (-) to the benthos] were macro-detritus (+0.028 gC m^-2day^-1), fallout of particulate debris (+3.3 gC m^-2day^-1), benthic net photosynthesis during the day (-0.06 gC m^-2 daylight period^-1), burial (0 gC m^-2day^-1), benthic respiration at night (-0.28 gC m^-2 night period^-1), and resuspension (-3.0 gC m^-2day^-1). Resuspension of sediment at this station was found to have a controlling effect on the sediment organic carbon content. Benthic photosynthesis was able to provide 79% of the organic carbon required by the benthos for respiration during the daylight hours. A carbon-flow diagram linking together all of the above measurements is presented.     

Modelling the growth and harvest yield of the giant kelp Macrocystis pyrifera

The giant kelp Macrocystis pyrifera is one of the largest and fastest growing seaweeds and is dominant over large areas of the west coast of North America. A model of its growth has been developed which describes plant biomass and production over the course of a year as a function of environmental parameters which affect the light flux. Such parameters include water clarity, spacing between plants, bottom depth, latitude, harvesting activity, and photosynthetic response (P _max and I _k ). Model results for a standard set of conditions (latitude 33°N, 3 m plant spacing, water absorbance of 0.115 m^-1 and 12 m depth) yield a peak daily gross production of almost 6 g C m^-2 d^-1, peak daily net production of almost 3 g C m^-2 d^-1, and a peak specific growth rate of about 0.022 d^-1. Annual gross production for this case is 1 567 g C m^-2 yr^-1; annual net production is 537 g C m^-2 yr^-1. These values are comparable to those from field measurements. Size and timing of biomass and production peaks are affected by changes in the parameters describing the light field, with peaks usually occurring later in the year for more adverse circumstances. Inhigher latitudes, the seasonal variation is so extreme that the plant could not last the year at 53° N in 12 m of water, although it is able to survive the year in shallower water. Harvesting has severe effects on biomass and production. Model results suggest that light limitation is a very important constraint on kelp growth that should not be overlooked. This implies that differences in parameters describing two environments must be considered when comparing results obtained at different locales.     

Differences in productivities between the Great Australian Bight and the Gulf of Carpentaria,Australia, in summer

Phytoplankton standing crop (chlorophyll a) and primary productivity were recorded, and zooplankton biomass was estimated in the two large bays of Australia, the Great Australian Bight on the south coast (December, 1965) and the Gulf of Carpentaria on the north coast (December, 1968). In the Gulf of Carpentaria, the phytoplankton standing crop (average, 27.3 mg chlorophyll a m^-2) and primary productivity (average, 133.1 mg C m^-2 h^-1), as well as zooplankton biomass (average, 305.3 mg wet weight m^-3) are much higher than in the Great Australian Bight (12.1 mg chlorophyll a m^-2, 18.2 mg C m^-2 h^-1, 7.1 mg wet weight m^-3, respectively). The unexpectedly low productivity values in the Great Australian Bight are attributable to environmental conditions of this bay, which obtains neither replenishment of nutrients from the land nor receives upwelling of deep water.     

Influence of antecedent water temperatures on standing crop of aSargassum spp.-dominated reef flat in Hawaii

The standing crop of seaweeds was estimated monthly over a 22 mo period from 1971 to 1973 on a reef at Waikiki, Honolulu, Hawaii. Wet and dry standing crops averaged 1.2 and 0.26 kg m^–2, respectively. Approximately 70% of the total biomass was contributed by a single species,Sargassum polyphyllum, and the remainder was contributed by 29 other species. The size of the total standing crop and ofS. polyphyllum in particular was highly correlated with antecedent water temperatures. The highest correlation was with temperatures recorded 3 to 4 wk prior to the estimate of standing crop. Approximately 65% of the variability of the standing crop on the reef was explained by this single variable. Multiple linear-regression analyses did not uncover further significant environmental factors related to theS. polyphyllum standing crop. Temperature may have acted by stimulating the vegetative growth of new plants between the period of minimal abundance in February and maximal abundance in October. The population decline after November relates to the switch to reproductive growth. Total productivity on the reef was estimated to be 16 800 to 19 314 kg dry wt ha^–1 yr^–1, based on the average size of the standing crop and estimates of removal rate and turnover time, respectively.     

The trophic role of glycolic acid in coastal seawater. II. Seasonal changes in concentration and heterotrophic use in Ipswich Bay,Massachusetts, USA

Glycolic acid is a known algal excretory product which is found in marine waters and is readily metabolized by marine bacteria. The following parameters were measured over the course of a year in Ipswich Bay: chlorophyll a, temperature, viable bacteria, heterotrophic uptake of glycolate, and glycolate concentrations. The latter two were combined to give estimates of the flux of glycolate for a station 3 km out in the bay and for an inshore station. Pronounced seasonal changes were found for all parameters. Statistically significant correlations between heterotrophic V _max and glycolic acid concentrations and temperature were found, but not between planktonic chlorophyll a and any of the other parameters. Measurements of glycolic acid flux yielded an annual flux of 2.84 g m^-2 for the bay station, which is about 0.5% of the bay phytoplankton production. Glycolic acid turns over roughly 12 times per year in the bay. It contributes perhaps as much to bacterial metabolism as any other single substrate, but is apparently not of dominating importance.     

In situ water motion and nutrient uptake by the giant kelp Macrocystis pyrifera

Rates of NO ₃ ^- uptake by individual blades of Macrocystis pyrifera (L.) C. Agardh were measured at different flow rates in the laboratory. Dissolution rates of hemispherical, plaster buttons attached to the blade surface provided a relative measure of flow rates over blades used in uptake experiments and also over intact blades of adult kelp plants in situ (Laguna Beach, California, USA; 1981). Laboratory results indicated that uptake was saturated at a flow rate equivalent to 2.5 cm s^-1 current velocity. Flow rates over intact blades in situ always exceeded this uptake saturation level. Wave surge and movement of plant surfaces relative to the surrounding water provided sufficient flow to saturate uptake, even in a dense kelp canopy during low-current and calm sea-state conditions.     

Metabolism and model of an estuarine bay ecosystem affected by a coastal power plant

The effects of a coastal power plant on an outer estuarine bay ecosystem on the west coast of Florida were evaluated with measurements and an ecological model. Field measurements of community metabolism and biomass were taken from the thermally affected bay and from similar control bays. Model simulations were used to help understand these observations in terms of ecosystem structure and functioning.In the outer discharge bay the direct impact of the thermal plume was diluted and spread overlarge areas. The ecosystem developed structure and functions with lower biomass than in the control bays but with slightly faster rates of organic turnover. The productive turnover time of producer biomass during the summer was about 5 days in the discharge bay and about 6 days in the control bays. Power plant influence on total community metabolism was small with less than 10% difference in annual averages between the discharge and control bays (5.22 and 5.58 g O₂/m²/day). The selection for faster metabolic turnover rates in the discharge by was evidenced by a dominance of plankton metabolism over benthic metabolism. The annual average gross planktonic production was around 3 g O₂/m²/day in the discharge bay and around 2 g/m²/day in the control bays.In the model, temperature served as a stimulant to both productivity and respiration. When the isolated effects of increased temperature were simulated the model responded with lower producer biomass and faster rates of organic turnover, as was found in field measurements. These simulations also showed increased nutrient recycling and indicated patterns of temperature-induced migrations. Since power plant operation affected water exchange in the bays, several levels of total water exchange were simulated. These simulations indicated the importance of water exchange as a stabilizing factor, especially for sensitive compartments with rapid turnover rates (i.e. plankton and phosphorus stocks). Simulations of the effects of future power plant units on the bay ecosystem showed no large changes in total metabolism but indicated larger effects of plankton entrainment mortality and temperature-induced migrations of larger organisms.     

Carbon budget for the spring bloom in Auke Bay,Alaska

This paper describes a carbon budget for the spring phytoplankton bloom in Auke Bay, a subarctic bay in southeastern Alaska. The budget was constructed using semiweekly data on carbon production, particulate carbon in the water column, and cumulative sedimentation of carbon, chlorophyll a, and pheopigments. From these measured parameters, seasonal carbon consumption, utilization, and import/export terms were derived. The chlorophyll and pheopigment data were used to partition carbon sinking out of the photic zone between phytoplankton cells and fecal material. The difference between total carbon production and carbon available for consumption was attributed primarily to carbon import/export related to advection of water masses into and out of the bay. Separate budgets were developed for each of five sampling years (1985–1989). An average of 130±16 g C/m² were produced by phytoplankton during each spring. Our model suggests that an average of 70% of this carbon was available for consumption by grazers within the bay; the remaining 30% is assumed to have been exported from the bay by advective transport. Of the available (non-exported) carbon, an average of 55% was consumed by grazers, 34% sank out of the photic zone in the form of uneaten algae, and about 11% remained at the end of the sampling period in the form of phytoplankton standing stocks. Overall, about 27% of the carbon produced each spring in Auke Bay (35 gC/m²) was used for growth and respiration by first-order consumers within the bay.     

Factors affecting the distribution of rock-lobster larvae (Panulirus longipes cygnus), with reference to variability of plankton-net catches

One of the major problems in the quantitative study of the distribution of larvae of the western rock lobster Panulirus longipes cygnus George, is the high variability in planktonnet catches taken at night, even when replicate hauls are made at a single station. Some of the possible causes of this variability are considered. The density of early and late-stage phyllosoma larvae caught at the surface at night showed no direct relationship to either temperature or salinity. Early-stage phyllosoma larvae were equally well represented at the surface under all light conditions encountered at night. In contrast, the density of late-stage phyllosoma larvae at the surface was, on an average, four times greater on dark nights than on bright moonlight nights. The mean density for all phyllosoma stages decreased rapidly with depth, i.e., from 0.70/1000 m³ at the surface to 0.04/1000 m³ at 25 to 35 m, indicating that the vertical movements at night are not very extensive. The density of early-stage phyllosoma larvae in the surface at night showed no relationship to micronekton biomass (measured as wet weight). Density of late-stage phyllosoma larvae at the surface at night increased slightly, but significantly, with increase of micronekton biomass, possibly indicating similar behavioural responses to light or some other factor(s). The distribution of cast exuviae of final-stage phyllosoma larvae indicated that metamorphosis to the puerulus phase can occur more than 100 miles (161 km) from shore. Moreover, 80% of the puerulus larvae caught were within 80 miles (129 km) of the shore. Mean densities of puerulus larvae taken at and below the surface at night were 0.04/1000 m³ and 0.10/1000 m³, respectively, although total numbers were low. This evidence indicates that, after metamorphosis, the puerulus larvae return to the coastal environment at some subsurface depth.     

On the Pollution Caused by Organic Materials in Chinhae Bay,Korea

In August 1989, oxygen deficient conditions of bottom waters occurred in the Chinhae Bay together with a steep pycnocline. Dissolved oxygen contents were lower than 1 ml/l from 3 m depth in the inner Masan Bay and from 10 m depth in the outer Masan Bay. in Kohyonsong Bay, surface salinity was about 29%_° and an oxygen deficient condition occurred in the bottom waters. Near Somodo Island, surface waters containing more than 30 μ/l of chlorophyll α and over 50 μmol/l of nitrate could be distinguished. in Masan Bay ammonia and phosphate concentrations increased with increasing depth suggesting the active degradation of organic materials in the bottom waters and leaching from sediments.

In Kohyonsong Bay, nitrate contents ranged from 1.9 to 5.4 μmol/l in the surface waters and subsurface maximum of chlorophyll α could be observed. ETS activity was 286.1 μl O₂/l-h in the surface waters of Masan Bay and respiratory oxygen consumption is likely to proceed at a rate of 1320 ml O₂/m²-d in the bottom waters of this bay. Primary productivity was 15.60 gC/m²-d in the inner Masan Bay and 0.75 gC/m²-d in Kohyonsong Bay.

In Masan Bay, amino acids content in the sinking materials collected by the sediment trap deployed at the bottom layer was 264 mgC/m²-d. This amount is equivalent to 6.8% of the amino acids produced in the water column by primary production. in Kohyonsong Bay 95 mg/Cm²-d of amino acids was collected corresponding to 50.8% of amino acids produced in the water column. in Kohyonsong Bay large faecal pellets produced in shellfish farms were easily settled in the sediment because of the weak current regime.     

Ecological investigations of the marine nematode Metoncholaimus scissus

Significant concentrations of the large oncholaimid Metoncholaimus scissus Wieser and Hopper (1967) (>7000 animals/24 cm² area or 2.68x10⁶ specimens/m²) have been noted in soft-bottom turtle grass communities. Activities of the nematode frequently show an indirect relationship to blooms of the benthic diatom Pleurosigma balticum. Analysis of biomass data for M. scissus (as much as 28 g wet weight/m²) demonstrate the importance of microsite activity and sensitivity of the species to small significant changes in the benthic environment.This work was supported by grant GM 12824 from the National Institutes of Health to the Institute of Marine and Atmospheric Sciences, University of Miami. Contribution No. 1188 from the Institute of Marine and Atmospheric Sciences, University of Miami and from the Entomology Research Institute, Research Branch, Canada Department of Agriculture, Ottawa.     

Energy relations of the bivalve Mercenaria mercenaria on an intertidal mudflat

Measurements of the filtration rate and oxygen consumption of the bivalve Mercenaria mercenaria (L.) have been made using flowing water systems, and then combined with growth and survivorship data to construct an energy budget for a tidal-flat population in Southampton Water. Estimated consumption (1292 kcal m^-2 year^-1) is mainly deposited as faeces and pseudofaeces (759 kcal) or excreted (160 kcal); 29% (374 kcal) is assimilated, of which 241 kcal are used for respiration, 72 kcal for flesh production and 61 kcal for gamete production. M. mercenaria contributes significantly to other trophic groups in Southampton Water; predators remove an estimated 55 kcal m^-2 year^-1, and 829 kcal pass to the scavenger/decomposer chain.     

Reproduction in the seagrass Zostera novazelandica on intertidal platforms in southern New Zealand

This study investigates the reproductive periodicity and reproductive output of the seagrass Zostera novazelandica on two intertidal reefs. Peak numbers of flowering shoots occurred during March (late summer) of two years at both sites and no flowering shoots occurred during the winter months of July to September. There were greater numbers of flowering shoots in seagrass patches in the low intertidal zone (up to an average of 55 per 0.1 m²) compared to the middle (up to 20 per 0.1 m²) and upper (up to 9 per 0.1 m²) zones, and about three times greater reproductive output in patches associated with tidepools compared to those not bordering tidepools. The average number of inflorescences per shoot was 3.1 (±0.25) at one site vs 1.2 (±0.08) at the other, and showed a progressive decrease from the lower shore to the upper shore. Patches associated with tidepools had twice the number of inflorescences per shoot (2.8 ± 0.24) than patches not bordering tidepools (1.5 ± 0.16). The number of flowering shoots was highly correlated with leaf width, leaf length, and ramet density, while the leaf-area index decreased from the lower shore to the upper shore. The reproductive effort of plants, as measured by the percent biomass invested in flowering shoots during peak reproduction, was significantly different between sites, tidepool associations, and shore level. For all the variables measured, there was considerable spatial variation, with significant interaction terms between most factors investigated. In laboratory experiments, more inflorescences were produced at light intensities of 30 and 300 μE m⁻² s⁻¹ than at 100 μE m⁻² s⁻¹. At a salinity of 17‰, 1.5 × the number of flowers was produced than at 33‰, while none was formed at 70‰. Plants cultured at 5 °C had about three times the number of inflorescences than those at 15 °C, while none was formed at 25 °C. Received: 25 June 1997 / Accepted: 24 September 1997     

Effects of silicate depletion on photosynthesis by diatoms in the plume of the Hudson River

Silicate depletion was observed during a bloom of netplankton diatoms. Netplankton chlorophyll a increased over the same salinity range and at the same rate that silicate decreased. Silicate depletion coincided with a decrease in the apparent Si:N uptake ratio as well as a change in the magnitude and diurnal phasing of light saturated photosynthesis (P _m ^B ) by netplankton diatoms. Nanoplankton P _m ^B was unaffected by silicate depletion and increased with temperature. Consequently, nanoplankton P _m ^B eventually exceeded netplankton P _m ^B while netplankton biomass was still increasing relative to nanoplankton biomass.     

Trophic implications of cross-shelf copepod distributions in the Southeastern Bering Sea

Spring distributions of some numerically dominant copepods reflect associations with two distinct water masses separated along the 80- to 100-m isobaths. Seaward of this middle shelf front, the oceanic Bering Sea hosts populations of Calanus cristatus, C. plumchrus, and Eucalanus bungii bungii; Metridia pacifica, Oithona similis, and Pseudocalanus spp. are also present. The large oceanic species are much less abundant in waters shallower than 80 m where the community is seasonally dominated by smaller copepods, O. similis, Acartia longiremis, and Pseudocalanus spp. Experimental and field-derived estimates of carbon ingestion indicate that the oceanic/outer shelf copepods can occasionally graze the equivalent of the daily plant production and probably routinely remove 20–30% of the primary productivity. Conversely, stocks of middle shelf copepods rarely ingest more than 5% of the plant carbon productivity. During 45 d between mid April to late May, 1979, approximately three times more organic matter was ingested m^-2 by the outer shelf/oceanic copepod community than by middle shelf species. This imbalance in cross-shelf grazing permits middle shelf phytoplankton stocks to grow rapidly to bloom proportions, and to sink ungrazed to the seabed. Over the outer shelf and particularly along the shelf break, a much closer coupling to phytoplankton supports a large biomass of oceanic grazers. Here, copepod stocks approaching 45 g dry wt m^-2 occur in late spring as a narrow band at the shelf break.Supported by National Science Foundation Grant DPP 76-23340Contribution no. 485, Institute of Marine Science, University of Alaska, Fairbanks     

Flow,flapping, and photosynthesis ofNereocystis leutkeana: a functional comparison of undulate and flat blade morphologies

A number of species of macroalagae possess a flat, strap-like blade morphology in habitats exposed to rapidly-moving water whereas those at protected sites have a wider, undulate blade shape. We have explored the functional consequences of flat, narrow vs. wide, undulate blade morphologies in the giant bull kelpNereocystis luetkeana. Our study focused on the behavior of blades in ambient water currents and the consequences of that behavior to breakage and to photosynthesis. In flowing water, the narrow, flat blades flap with lower amplitude and collapse together into a more streamlined bundle than do wide, undulate blades, and hence experience lower drag per blade area at a given flow velocity. If the algae at current-swept sites had ruffled blades, drag forces would sometimes be sufficient to break the stipes. However, flat blades in a streamlined bundle experience more self-shading than do undulate blades, which remain spread out in water currents. Thus, there is a morphological trade-off between reducing drag and reducing self-shading. Photosynthetic¹⁴C-HCO₃ uptake rates decrease in slow flow when the boundary layer along the blade surface across which diffusion takes place is relatively thick. However, blade flapping, which stirs water near the blade surface, enhances carbon uptake rates in slow water currents for both the undulate and the flat morphologies.