Survival, growth and reproduction of the salt-marsh amphipod Uhlorchestia spartinophila reared on natural diets of senescent and dead Spartina alterniflora leaves

The talitrid amphipod Uhlorchestia spartinophila lives in close association with standing-dead leaves of the smooth cordgrass Spartina alterniflora Loisel in salt marshes along the Atlantic coast of North America. This study probed the strength of the trophic link between the amphipod population and the decomposition process in this detrital-based ecosystem. We measured survival, growth and reproductive output in groups of amphipods reared for 6 wk on five diets derived from sheath and blade portions of S. alterniflora leaves just prior to (senescent) and during (dead) decomposition. In unfed treatments, the daily specific mortality rate was 0.391 and starved amphipods survived no longer than 11 d. Among the fed treatments, a diet of senescent sheaths resulted in the lowest survival (20%) and yielded no offspring. Groups fed senescent blades, dead sheaths, dead blades and unwashed dead sheaths had survival rates of 56 to 84% and produced 5.0 to 12.5 offspring replicate⁻¹. Sex ratio usually favored females, but approached unity in treatments with high overall survival, suggesting that quality of available food resources may influence sex ratio in this species. Mean specific growth rates (mm mm⁻¹ d⁻¹) ranged from 0.013 to 0.016, and matched previous estimates of growth from field populations. Overall ecological performance (survival + growth + reproduction) was similar for all food treatments, except senescent sheaths, which yielded a final mean (±SD) dry biomass (0.4 ± 0.42 mg replicate⁻¹) of amphipods significantly lower than that of other diets (1.7 ± 0.81 to 2.6 ± 0.69 mg replicate⁻¹). Natural diets derived from decomposing cordgrass leaves can fulfill the nutritional requirements of U. spartinophila populations, but variation in initial amounts of living fungal biomass among the five experimental diets only partially explained the responses of amphipods in our experiment. Structural characteristics and variation in rates of fungal occupation within different portions of cordgrass leaves may affect the amphipod's ability to access plant production made available by decomposers. Received: 12 December 1996 / Accepted: 18 December 1996     

Light and carbon balance in the seagrass Thalassia testudinum : evaluation of current production models

The production dynamics and carbon balance of Thalassia testudinum in the lower Laguna Madre, Texas, USA, were examined during the 1995 summer period based on in situ photosynthesis vs irradiance (PI) measurements and continuous measurements of underwater photon-flux density (PFD). The validity of applying the H _sat model, used to calculate production for Zostera marina as the product of the maximum rate of photosynthesis (P _max) and daily hours of saturating irradiance (H _sat) was assessed for T. testudinum by comparison with integrated production estimates derived through numerical integration. Gross integrated production values were combined with dark-respiration measurements of photosynthetic (PS) and non-photosynthetic (NPS) tissues and areal biomass to generate daily whole-plant carbon balance. Production and whole-plant carbon balance are discussed in relation to surface and underwater PFD measurements, biomass and other physical and chemical parameters collected during a 1 yr period from January to December 1995. The H _sat model significantly underestimated production during all summer months, averaging 70% of integrated production over the entire study period. Gross integrated production ranged between 11.5 mg C g⁻¹ leaf dry wt d⁻¹ in June (during a period of unseasonably low PFDs caused by a drift-alga mat covering the seagrass bed) to 26.7 mg C g⁻¹ leaf dry wt d⁻¹ in July. Modeled net carbon gain was highest in July at 454 mg C m⁻² d⁻¹ (1.4 g dry wt m⁻² d⁻¹), sufficient to account for measured rates of leaf production in the study area and representative of T. testudinum populations of low productivity. During part of the summer period, however, the population was in negative carbon balance. The relatively low productivity of this population and the periods of negative carbon balance are attributed to low net photosynthesis:dark respiration (P _net:R _d) ratios, sporadic low-light periods, the small fraction of PS tissue relative to whole-plant biomass (5 to 13%) and nutrient limitation. Production models are sensitive to both light availability and the proportion of PS tissue supporting NPS biomass as reflected in whole-plant P _net:R _d ratios. Received: 13 August 1997 / Accepted: 6 March 1998     

Annual cycle of biomass of a threatened population of the intertidal seagrass Zostera japonica in Hong Kong

The phenology and primary productivity of a population of Zostera japonica (Aschers. & Graebn.) threatened by the construction of Hong Kong's new international airport were studied over a 12-month period. The need to conserve the population, and the small leaf size of Z. japonica rendered traditional destructive or marking techniques inapplicable for percentage cover and biomass estimation. A nondestructive method based on image analysis techniques was therefore devised for repeated estimation of percentage cover, biomass and leaf area index. This technique, which involved random quadrat sampling, photographic recording and image analysis, was able to provide data on the three parameters with acceptable precision and was cost-effective in the field. Z. japonica demonstrated a strongly seasonal cycle of vegetative growth, with different patterns for leaf density (peak in March) and overall bed area (peak in June). Total (above- and below-ground) net primary productivity was estimated at between 344 and 688 g AFDW m⁻² yr⁻¹. Percentage cover of Z. japonica was negatively correlated with total suspended solids (TSS) in the water column while total bed area was negatively correlated with water salinity. Increased sedimentation associated with the new airport project was identified as one important factor affecting the growth of the seagrass, as TSS reached the high level of ≈1 g DW l⁻¹ during the first half of the study period. Sediment traps set in the beds also recorded potential sedimentation rates at between 2.89 and 14.5 mg cm⁻² d⁻¹. This high turbidity resulted in a sharp decrease in the density of Clithon spp., the dominant grazers of epiphytic algae on Z. japonica. Effects of sedimentation and shading on growth of Z. japonica were investigated by field manipulative experiments. Experimental increase of sedimentation rate and shade both resulted in larger decreases in percentage cover and above-ground AFDW compared with the control. Received: 3 March 1997 / Accepted: 14 March 1997     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Predaceous gastropods regulate new-shell supply to salt marsh hermit crabs

Marsh hermit crabsPagurus longicarpus Say directly acquire new shells as the predatory gastropodMelongena corona Gmelin consumes marsh periwinkles,Littorina irrorata Say. The influx rate of new shells into a salt marsh hermit crab population was measured by marking live periwinkles and daily recovering the shells from hermit crabs over periods of 3 to 6 d. Average rates of new shell acquisition ranged from 4.0 to 23.3 new shells per day from salt marsh areas of 4×10 m. Such consistently high rates contrast with the negligible rates generally assumed for new shell entry into hermit crab populations. The number of new shells acquired each day varied directly with the number of the predatory gastropod,M. corona, present in each study area at both natural and manipulated predator densities. Empty shells on the substrate are usually considered as the primary source of new shells to hermit crabs. However, over 500 empty shells had to be placed on the substrate in a 4×10 m area to provide a daily rate of 20 new shells to the hermit crab population.This is the first in a new contribution series from the Florida State University Marine Laboratory No. 1001     

Heterotrophic nitrogen fixation (acetylene reduction) during leaf-litter decomposition of two mangrove species from South Florida, USA

Heterotrophic nitrogen-fixation (acetylene reduction) was measured during decomposition (under dark conditions) of Rhizophora mangle L. and Avicennia germinans (L.) Stearn leaf litter. Nitrogen-fixation rates in leaf litter increased following 24 d incubation, then decreased after ≃44 d for both species. Maximum rates of 66.2 and 64.6 nmol C₂H₄ g⁻¹ dry wt h⁻¹ were reached by R. mangle and A. germinans leaf litter, respectively. Higher fixation rates of leaf litter were associated with an increase in water content and sediment particles on leaf surfaces of both species. Rates of nitrogen fixation by diazotrophs attached to sediment particles were not significantly different from zero. With additions of d-glucose, ethylene production rates increased by factors of 625-, 34- and 7-fold for sediment, R. mangle and A.␣germinans leaf litter, respectively, compared to rates prior to enrichment. These organically enhanced rates of nitrogen fixation on leaves could be accounted for by increased activity associated with attached sediment particles and not the leaf material. Total phenolics [reported as tannic acid equivalent (TAE) units] decreased nitrogen-fixation rates when added to d-glucose-enriched sediment at >20 mg TAE l⁻¹. Phenolic compounds could explain the initial lag in rates of nitrogen fixation during leaf-litter decomposition of R. mangle (initial content of 110.8 mg TAE g⁻¹ dry wt), but not of A. germinans (initial content of 23.4 mg TAE g⁻¹ dry wt). The higher phenolic content and reportedly lower carbon substrate of R. mangle did not result in species-specific differences in either the magnitude or temporal pattern of nitrogen fixation compared to A. germinans leaf litter. We conclude that the availability of organic substrates leached from the leaf litter along with colonization by the heterotrophic diazotrophs (as indicated by sediment accumulation) controls nitrogen-fixation rates in a similar manner in the leaf litter of both species. Received: 8 August 1997 / Accepted: 4 December 1997     

Intraspecific physiological variability of the gastropod Littorina saxatilis related to the vertical shore gradient in the White and North Seas

 Physiological responses to desiccation and temperature stress as well as behavioural responses to fast and abrupt environmental changes were investigated in high- and low-shore Littorina saxatilis (Olivi) from several populations from the White and North Seas. Variations in evaporation rates, resistance to air exposure and to acute and chronic temperature stress between animals from different shore levels were similar in White and North Sea periwinkles, consistent with the adaptive nature of these variations. High-shore snails were found to be able to conserve body water reserves better, to resist higher temperatures and to survive longer under conditions of combined temperature and desiccation stress than their low-shore counterparts. In a temperature range of 25 to 35 °C, the rate of evaporative water loss was positively correlated with temperature in low-shore snails while being largely temperature-independent in high-shore snails. Median lethal time during air exposure in L. saxatilis was negatively but not linearly related to the temperature of exposure. In a temperature range of 30 to 38 °C, the resistance to heat exposure in air was only slightly dependent on the temperature, with Q ₁₀ = 1.4 for the median lethal time; the heat resistance dropped drastically at temperatures above 38 °C, with Q ₁₀ = 593.8. This suggests different mechanisms of temperature resistance in different parts of the studied temperature range. In contrast, behavioural response to extreme salinity fluctuations was not uniform in the high- and low-shore periwinkles from the White and North Seas, which may reflect specific environmental conditions at different shore levels in the two areas studied. Observed physiological and behavioural variations are discussed from the viewpoint of different adaptive strategies employed by eulittoral and eulittoral-fringe animals within populations of a single species. Received: 13 December 1999 / Accepted: 11 April 2000     

The amphipod Corophium multisetosum (Corophiidae) in Ria de Aveiro (NW Portugal). II. Abundance, biomass and production

 The abundance and biomass of Corophium multisetosum Stock, 1952 were determined from benthic corer samples collected monthly over 1 yr in the upper reaches of Canal de Mira (Ria de Aveiro, Portugal). Both density and biomass over the sampling period were negatively correlated with water temperature and positively correlated with chlorophyll a concentration in the sediment. C. multisetosum density was significantly negatively correlated with plant biomass and positively correlated with salinity. The nature of the sediment, favourable environmental conditions, high availability of food and low interspecific competition allowed the population to reach a maximal density of 200 × 10³ individuals m⁻² and a maximal biomass (ash-free dry wt, AFDW) of 62 g_AFDW m⁻². The population was highly productive, especially during the autumn/winter period. Production, estimated by two different methods (Hynes method: 251 g_AFDW m⁻² yr⁻¹; Morin–Bourassa method: 308 g_DW m⁻² yr⁻¹), was much higher than the values reported for other Corophium species. The annual P:Bˉ ratio (10) was high, but similar to values reported for Swedish populations of C. volutator and lower than the values estimated from Mediterranean populations of C. insidiosum. Received: 8 October 1999 / Accepted: 22 June 2000     

Phytoplankton blooms are strongly impacted by microzooplankton grazing in coastal North Pacific waters

Phytoplankton growth and microzooplankton grazing were measured in two productive coastal regions of the North Pacific: northern Puget Sound and the coastal Gulf of Alaska. Rates of phytoplankton growth (range: 0.09–2.69 day⁻¹) and microzooplankton grazing (range: 0.00–2.10 day⁻¹) varied seasonally, with lowest values in late fall and winter, and highest values in spring and summer. Chlorophyll concentrations also varied widely (0.19–13.65 μg l⁻¹). Large (>8 μm) phytoplankton cells consistently dominated phytoplankton communities under bloom conditions, contributing on average 65% of total chlorophyll biomass when chlorophyll exceeded 2 μg l⁻¹. Microzooplankton grazing was an important loss process affecting phytoplankton, with grazing rates equivalent to nearly two-thirds (64%) of growth rates on average. Both small and large phytoplankton cells were consumed, with the ratio of grazing to growth (g:μ) for the two size classes averaging 0.80 and 0.42, respectively. Perhaps surprisingly, the coupling between microzooplankton grazing and phytoplankton growth was tighter during phytoplankton blooms than during low biomass periods, with g:μ averaging 0.78 during blooms and 0.49 at other times. This tight coupling may be a result of the high potential growth and ingestion rates of protist grazers, some of which feed on bloom-forming diatoms and other large phytoplankton. Large ciliates and Gyrodinium-like dinoflagellates contributed substantially to microzooplankton biomass at diatom bloom stations in the Gulf of Alaska, and microzooplankton biomass overall was strongly correlated with >8 μm chlorophyll concentrations. Because grazing tended to be proportionally greater when phytoplankton biomass was high, the absolute amount of chlorophyll consumed by microzooplankton was often substantial. In nearly two-thirds of the experiments (14/23), more chlorophyll was ingested by microzooplankton than was available for all other biological and physical loss processes combined. Microzooplankton were important intermediaries in the transfer of primary production to higher trophic levels in these coastal marine food webs. Received: 12 November 1999 / Accepted: 4 October 2000     

Growth and productivity of the high Antarctic Bryozoan Melicerita obliqua

We analysed growth of the Antarctic bryozoan Melicerita obliqua (Thornely, 1924) by x-ray photography and stable isotope analysis. M. obliqua colonies form one segment per year, thus attaining maximum length of about 200 mm within 50 years. In the Weddell and Lazarev Seas, annual production/biomass ratio of M. obliqua is 0.1 yr⁻¹, which is in the range of other Antarctic benthic invertebrate populations. Production amounts to 3.34 mg C_orgm⁻² yr⁻¹ and 90.6 mg ash m⁻² yr⁻¹ on the shelf (100 to 600 m water depth), and to 0.13 mg C_orgm⁻² yr⁻¹ and 36.8 mg ash m⁻² yr⁻¹ on the slope (600 to 1250 m water depth). Received: 27 February 1998 / Accepted: 8 May 1998     

Spatial and temporal pattern in seagrass community composition and productivity in south Florida

We document the distribution and abundance of seagrasses, as well as the intra-annual temporal patterns in the abundance of seagrasses and the productivity of the nearshore dominant seagrass (Thalassia testudinum) in the south Florida region. At least one species of seagrass was present at 80.8% of 874 randomly chosen mapping sites, delimiting 12,800 km² of seagrass beds in the 17,000-km² survey area. Halophila decipiens had the greatest range in the study area; it was found to occur over 7,500 km². The range of T. testudinum was almost as extensive (6,400 km²), followed by Syringodium filiforme (4,400 km²), Halodule wrightii (3,000 km²) and Halophila engelmanni (50 km²). The seasonal maxima of standing crop was about 32% higher than the yearly mean. The productivity of T. testudinum was both temporally and spatially variable. Yearly mean areal productivity averaged 0.70 g m⁻²day⁻¹, with a range of 0.05–3.29 g m⁻² day⁻¹. Specific productivity ranged between 3.2 and 34.2 mg g⁻¹ day⁻¹, with a mean of 18.3 mg g⁻¹ day⁻¹. Annual peaks in specific productivity occurred in August, and minima in February. Integrating the standing crop for the study area gives an estimate of 1.4 × 1011 g T. testudinum and 3.6 × 10¹⁰ g S. filiforme, which translate to a yearly production of 9.4 × 10¹¹ g T. testudinum leaves and 2.4 × 10¹¹ g S. filiforme leaves. We assessed the efficacy of rapid visual surveys for estimating abundance of seagrasses in south Florida by comparing these results to measures of leaf biomass for T. testudinum and S. filiforme. Our rapid visual surveys proved useful for quantifying seagrass abundance, and the data presented in this paper serve as a benchmark against which future change in the system can be quantified. Received: 30 January 2000 / Accepted: 24 July 2000     

Availability of two forms of dissolved nitrogen to the coral Pocillopora damicornis and its symbiotic zooxanthellae

The relative contribution of dissolved nitrogen (ammonium and dissolved free amino acids DFAAs) to the nitrogen budget of the reef-building coral Pocillopora damicornis was assessed for colonies growing on control and ammonium-enriched reefs at One Tree Island (southern Great Barrier Reef) during the ENCORE (Enrichment of Nutrient on Coral Reef; 1993 to 1996) project. P. damicornis acquired ammonium at rates of between 5.1 and 91.8 nmol N cm⁻² h⁻¹ which were not affected by nutrient treatment except in the case of one morph. In this case, uptake rates decreased from 80.5 to 42.8 nmol cm⁻² h⁻¹ (P < 0.05) on exposure to elevated ammonium over 12 mo. The presence or absence of light during measurement did not influence the uptake of ammonium ions. Nitrogen budgets revealed that the uptake of ammonium from concentrations of 0.11 to 0.13 μM could completely satisfy the demand of growing P. damicornis for new nitrogen. P. damicornis also took up DFAAs at rates ranging from 4.9 to 9.8 nmol N cm⁻² h⁻¹. These rates were higher in the dark than in the light (9.0 vs 5.1 nmol m⁻² h⁻¹, P < 0.001). Uptake rates were highest for the amino acids serine, arginine and alanine, and lowest for tyrosine. DFAA concentrations within the ENCORE microatolls that received ammonium were undetectable, whereas they ranged up to 100 nM within the control microatolls. The contribution of DFAAs to the nitrogen budget of P. damicornis constituted only a small fraction of the nitrogen potentially contributed by ammonium under field conditions. Even at the highest field concentrations measured during this study, DFAAs could contribute only ≃11.3% of the nitrogen demand of P.␣damicornis. This contribution, however, may be an important source of nitrogen when other sources such as ammonium are scarce or during periods when high concentrations of DFAAs become sporadically available (e.g. cell breakage during fish-grazing). Received: 22 April 1998 / Accepted: 3 November 1998     

Root respiration and oxygen flux in salt marsh grasses from different elevational zones

Plants growing in waterlogged environments are subjected to low oxygen levels around submerged tissues. While internal oxygen transport has been postulated as an important factor governing flooding tolerance, respiration rates and abilities to take up oxygen under hypoxic conditions have been largely ignored in plant studies. In this study, physiological characteristics related to internal oxygen transport, respiration, and oxygen affinity were studied in low intertidal marsh species (Spartina alterniflora and S. anglica) and middle to high intertidal species (S. densiflora, S. patens, S. foliosa, a S. alterniflora × S. foliosa hybrid, S. spartinae, and Distichlis spicata). These marsh plants were compared to the inland species S. pectinata and the crop species rice (Oryza sativa), corn (Zea mays), and oat (Avena sativa). Plants were grown in a greenhouse under simulated estuarine conditions. The low marsh species S. anglica was found to transport oxygen internally at rates up to 2.2 μmol O₂ g fresh root weight⁻¹ h⁻¹. In contrast, marsh species from higher zones and crop species were found to transport significantly less oxygen internally, although rice plants were able to transport 1.4 μmol g⁻¹ h⁻¹. Under hypoxic conditions, low marsh species were better able to remove dissolved oxygen from the medium compared to higher marsh species and crops. The oxygen concentration at which respiration rates declined due to limited oxygen (P _crit) was significantly lower in low marsh species compared to inland and crop species; P _crit ranged from <4 μM O₂ in the low marsh species S. anglica up to 20 μM in the inland species corn. Flooding-sensitive crop species had significantly higher aerobic respiration rates compared to flooding-tolerant species in this study. Crop species took up 3.6–6.7 μmol O₂ g⁻¹ h⁻¹ while all but one marsh species took up <3.5 μmol O₂ g⁻¹ h⁻¹. We conclude that oxygen transport, aerobic demand, and oxygen affinity all play important and interrelated roles in flood tolerance and salt marsh zonation.     

A novel symbiosis between a cyanobacterium, Synechococcus sp., an aplastidic protist, Solenicola setigera, and a diatom, Leptocylindrus mediterraneus, in the open ocean

The abundance, biomass and distribution of Solenicola setigera, a colonial heterotrophic protist found only with the centric chain-forming diatom Leptocylindusmediterraneus, are reported for four major ocean basins. The distribution is cosmopolitan, and abundances and biomass are usually low (<500 colonies l⁻¹); however, in the summer of 1993, we observed a major biomass component (range = 5 to 31 μg C l⁻¹) in the surface waters of the North Atlantic attributable to S. setigera. These colonies of S. setigera were exceptionally large, and unusual in possessing high abundances of Synechococcus sp., a normally solitary cyanobacterium, embedded in the matrix covering the cells. We hypothesize that this relationship was mutually beneficial for both Solenicola setigera and Synechococcus sp. Received: 5 January 1998 / Accepted: 22 May 1998     

Effect of nutrient enrichment in the field on the biomass, growth and calcification of the giant clam Tridacna maxima

Nutrients were added separately and combined to an initial concentration of 10 μM (ammonium) and/or 2 μM (phosphate) in a series of experiments carried out with the giant clam Tridacna maxima at 12 microatolls in One Tree Island lagoon, Great Barrier Reef, Australia (ENCORE Project). These nutrient concentrations remained for 2 to 3 h before returning to natural levels. The additions were made every low tide (twice per day) over 13 and 12 mo periods for the first and second phase of the experiment, respectively. The nutrients did not change the wet tissue weight of the clams, host C:N ratio, protein content of the mantle, calcification rates or growth rates. However, ammonium (N) enrichment alone significantly increased the total population density of the algal symbiont (Symbiodinium sp.: C = 3.6 · 10⁸ cell clam⁻¹, N = 6.6 · 10⁸ cell clam⁻¹, P = 5.7 · 10⁸ cell clam⁻¹, N + P = 5.7 · 10⁸ cell clam⁻¹; and C = 4.1 · 10⁸ cell clam⁻¹, N = 5.1 · 10⁸ cell clam⁻¹, P = 4.7 · 10⁸ cell clam⁻¹, N + P = 4.5 · 10⁸ cell clam⁻¹, at the end of the first and second phases of the experiment, respectively), although no differences in the mitotic index of these populations were detected. The total chlorophyll a (chl a) content per clam but not chlorophyll a per cell also increased with ammonium addition (C = 7.0 mg chl a clam⁻¹, N = 13.1 mg chl a clam⁻¹, P = 12.9 mg chl a clam⁻¹, N + P = 11.8 mg chl a clam⁻¹; and C = 8.8 mg chl a clam⁻¹, N = 12.8 mg chl a clam⁻¹; P = 11.2 mg chl a clam⁻¹, N + P = 11.3 mg chl a clam⁻¹, at the end of the first and second phases of the experiment, respectively). The response of clams to nutrient enrichment was quantitatively small, but indicated that small changes in inorganic nutrient levels affect the clam–zooxanthellae association. Received: 2 June 1997 / Accepted: 9 June 1997     

Irradiance and temperature regulation of oxygenic photosynthesis and O2 consumption in a hypersaline cyanobacterial mat (Solar Lake, Egypt)

 Short-term effects of temperature and irradiance on oxygenic photosynthesis and O₂ consumption in a hypersaline cyanobacterial mat were investigated with O₂ microsensors in a laboratory. The effect of temperature on O₂ fluxes across the mat–water interface was studied in the dark and at a saturating high surface irradiance (2162 μmol photons m⁻² s⁻¹) in the temperature range from 15 to 45 °C. Areal rates of dark O₂ consumption increased almost linearly with temperature. The apparent activation energy of 18 kJ mol⁻¹ and the corresponding Q ₁₀ value (25 to 35 °C) of 1.3 indicated a relative low temperature dependence of dark O₂ consumption due to mass transfer limitations imposed by the diffusive boundary layer at all temperatures. Areal rates of net photosynthesis increased with temperature up to 40 °C and exhibited a Q ₁₀ value (20 to 30 °C) of 2.8. Both O₂ dynamics and rates of gross photosynthesis at the mat surface increased with temperature up to 40 °C, with the most pronounced increase of gross photosynthesis at the mat surface between 25 and 35 °C (Q ₁₀ of 3.1). In another mat sample, measurements at increasing surface irradiances (0 to 2319 μmol photons m⁻² s⁻¹) were performed at 25, 33 (the in situ temperature) and 40 °C. At all temperatures, areal rates of gross photosynthesis saturated with no significant reduction due to photoinhibition at high irradiances. The initial slope and the onset of saturation (E _k = 148 to 185 μmol photons m⁻² s⁻¹) estimated from P versus E _d curves showed no clear trend with temperature, while maximal photosynthesis increased with temperature. Gross photosynthesis was stimulated by temperature at each irradiance except at the lowest irradiance of 54 μmol photons m⁻² s⁻¹, where oxygenic gross photosynthesis and also the thickness of the photic zone was significantly reduced at 40 °C. The compensation irradiance increased with temperature, from 32 μmol photons m⁻² s⁻¹ at 25 °C to 77 μmol photons m⁻² s⁻¹ at 40 °C, due to increased rates of O₂ consumption relative to gross photosynthesis. Areal rates of O₂ consumption in the illuminated mat were higher than dark O₂ consumption at corresponding temperatures, due to an increasing O₂ consumption in the photic zone with increasing irradiance. Both light and temperature enhanced the internal O₂ cycling within hypersaline cyanobacterial mats. Received: 30 November 1999 / Accepted: 11 April 2000     

Contribution of salps to carbon flux of marginal ice zone of the Lazarev Sea, southern ocean

In order to estimate the in situ grazing rates of Salpa thompsoni and their implications for the development of phytoplankton blooms and for the sequestration of biogenic carbon in the high Antarctic, a repeat-grid survey and drogue study were carried out in the Lazarev Sea during austral summer of 1994/1995 (December/January). Exceptionally high grazing rates were measured for S. thompsoni at the onset of a phytoplankton bloom (0.2 to 0.8 μg chlorophyll a l⁻¹) in December 1994, with up to ≃160 μg of plant pigments consumed by an individual salp of 7 to 10 cm length per day. Dense salp swarms extended throughout the marginal ice zone, consuming up to 108% of daily phytoplankton production and 21% of the total chlorophyll a stock. Due to the much faster sinking rates and higher carbon content of salp faecal pellets, the efficiency of downward carbon flux through salps is much higher than through the other major grazers, krill and copepods. S. thompsoni can thus export large amounts of biogenic carbon from the euphotic zone to the deep ocean. With the observed ingestion rates during December 1994, this flux could have attained levels of up to 88 mg C m⁻² d⁻¹, accounting for the bulk of the vertical transport of carbon in the Lazarev Sea. However, in January 1995, when phytoplankton concentrations exceeded a threshold level of 1.0 to 1.5 μg chlorophyll a l⁻¹, salps experienced a drastic reduction in their feeding efficiency, possibly as a result of clogging of their filtering apparatus. This triggered a dramatic reversal in the relationship, during which a dense phytoplankton bloom developed in conjunction with the collapse of the salp population. Increases in the biomass and geographic range of the tunicate S. thompsoni have occurred in several areas of the southern ocean, often in parallel with a rise in sea-surface temperature during sub-decadal periods of warming anomalies. Received: 10 August 1997 / Accepted: 21 October 1997     

Species composition, biomass and vertical distribution of micronekton over the mid-slope region off southern Tasmania, Australia

Seasonal sampling was carried out based on day/night, vertically stratified tows (100 or 125 m strata) in the upper 900 m of the water column over the mid-slope commercial fishing grounds south of Tasmania. A large midwater trawl (105 m² mouth area) was used with an opening/closing cod-end. Subtropical convergence and subtropical species dominated the fauna, but many less abundant, more widely-distributed species were also present. Fishes, which contributed 89% of micronekton biomass and 135 of 178 species, were dominated by the Myctophidae (48% biomass and 48 species). Twenty micronekton species made up 80% of the total biomass. Overall, the micronekton fish biomass in this region was 2.2 g m⁻² wet weight. A pronounced day/night shift in the distribution of biomass was attributable to diel migratory species. During the day, <0.2% of the total micronekton biomass was found in 0 to 300 m; most biomass was below 400 m, with peaks at 400 to 525 m and 775 to 900 m. At night, 53% of the biomass was found in 0 to 300 m, with progressively less in each deeper stratum. The vertical ranges of individual species typically exceeded 400 to 500 m during the day and night and were non-coincident, although nyctoepipelagic migrators were concentrated in the surface 200 m at night. Distinct epipelagic, lower and upper mesopelagic assemblages were identified, and patterns of epipelagic migration, limited migration and non-migration were categorised for species from each of the lower and upper mesopelagic assemblages. The vertical distribution of these assemblages was coincident with the primary water masses: subantarctic mode water (∼250 to 600 m) and antarctic intermediate water (below ∼700 m). The flux of migrating micronekton, estimated at 0.94 to 3.36 g C m⁻² yr⁻¹ to the lower mesopelagic and 1.14 to 4.06 g C m⁻² yr⁻¹ to the upper mesopelagic, appeared to be considerably outweighed by the consumption needs of aggregated mid-slope benthopelagic predators. We suggest that advection of mesopelagic prey in antarctic intermediate water may sustain aggregated populations of orange roughy (Hoplostethus atlanticus) and other predators on the micronekton in mid-slope depths at this site. Received: 2 April 1997 / Accepted: 21 August 1997     

Survival and bioturbation of the amphipod Monoporeia affinis in sulphide-rich sediments

Laboratory experiments were conducted to investigate the survival rate of Monoporeia affinis in sulphide-rich sediment with oxic overlying water, and the effect of amphipod bioturbation on sulphide and oxygen profiles. As long as the oxygen content in the water is high, the amphipods seem to avoid quite high concentrations (>200 μmol l⁻¹) of sulphide in the sediment by creating microhabitats where sulphide is rapidly oxidised. In cores with amphipods, a decrease of sulphide concentration was found in upper layers, while an increase of sulphide was found in deeper layers. Aggregation of amphipods generated pockets of light-brown sediment, characterised by high oxygen concentrations and no sulphide, and their depth was clearly dependent on amphipod density. This indicates that M. affinis has a potential to recolonise sulphide-rich sediments, devoid of macroscopic life, after the overlying water column has become oxygenated. Received: 13 April 2000 / Accepted: 8 September 2000     

Photosynthetic utilisation of carbon and light by two tropical seagrass species as measured in situ

In situ measurements of seagrass photosynthesis in relation to inorganic carbon (Ci) availability, increased pH and an inhibitor of extracellular carbonic anhydrase were made using an underwater pulse amplitude modulated (PAM) fluorometer. By combining the instrument with a specially designed Perspex chamber, we were able to alter the water surrounding a leaf without removing it from the growing plant. Responses to Ci within the chamber showed that subtidal plants of the seagrasses Cymodocea serrulata and Halophila ovalis had photosynthetic rates that were limited by the ambient Ci concentration depending on the irradiance that was available during short-term photosynthesis–irradiance trials. Relative electron transport rates (RETRs) at light saturation (up to 500 μ mol photons m⁻² s⁻¹) increased by 66–100% when the Ci concentration was increased from ca. 2.2 to 6.2 mM. On the other hand, intertidal plants of the same species exhibited a much lesser limitation of photosynthesis by Ci at any irradiance (up to 1500 μ mol photons m⁻² s⁻¹). Both species were able to use HCO⁻ ₃ efficiently, and there was stronger evidence for direct uptake of HCO⁻ ₃ rather than extracellular dehydration of HCO⁻ ₃ to CO₂ prior to Ci uptake. Subtidally, H. ovalis and C. serrulata grew to 10 and 12 m, respectively, where ambient irradiances were approximately 16 and 11% of those at the surface. Maximum RETRs (at light saturation) were lower for these deep-growing plants than for the intertidally growing ones. For both species, the onset of light saturation of photosynthesis (E _k) occurred at approximately 100 μ mol photons m⁻² s⁻¹ for the deep water populations, which was four and two times lower than for the shallow populations of C. serrulata and H. ovalis, respectively. This, and the differences in maximal photosynthetic rates (RETR _max), reflects an acclimation of the deep-growing populations to the lower light environment. The results presented here show that photosynthesis, as measured in situ, was limited by the availability of Ci for the deeper growing plants in Zanzibar, while the intertidally growing plants photosynthesised at close to Ci saturation. The latter result is contrary to previous conclusions regarding Ci limitations for these intertidal plants, and, in general, our findings highlight the need for performing similar experiments in situ rather than under laboratory conditions. Received: 4 April 2000 / Accepted: 31 August 2000