Nitrogen limitation of Codium fragile ssp. tomentosoides as determined by tissue analysis

The critical nitrogen concentration of Codium fragile ssp. tomentosoides (van Goor) Silva was 1.90% on a dry weight basis. Internal nitrogen levels of thalli collected from three Rhode Island populations ranged from 0.75±0.08 to 3.72±0.08%. Internal nitrogen content was minimal in summer and maximal in winter, indicating nitrogen limitation during the summer and nitrogen storage during the winter. Part of this nitrogen storage appeared to support new growth of C. fragile in the spring. Carbon:nitrogen (C:N) ratios were inversely related to the internal nitrogen concentration. Low internal nitrogen content indicated nitrogen limitation better than C:N ratios because the C:N ratio could be influenced by both carbon and nitrogen metabolism.     

Tidal influences on the photoperiodic induction of tetrasporogenesis in Bonnemaisonia hamifera (Rhodophyta)

Photoperiodic control of tetrasporangium formation was studied in two low intertidal populations of Bonnemaisonia hamifera (Trailliella-phase) in Galway Bay, Ireland between September 1985 and February 1986. Photoperiodic induction was affected by tides, principally by the time of day at which high and low water occurred through the spring/neap cycle. High water of spring tides falling at the beginning and end of the day reduced the effective daylength for plants, and caused an early onset of reproduction. The threshold irradiance signalling light-dark transition was estimated to lie between (1-) 5 and 10 mol photons m^-2 s^-1 (400 to 700 nm). Low water of spring tides falling in the middle of the day exposed plants to high air temperatures during periods when seawater temperatures were below the lower limit for sporangium induction. This caused reproduction to be resumed. Quantitative differences between the two populations were caused by microclimatic effects (temperature, light reduction during tidal flooding) resulting from their different vertical heights on the shore.     

Role of cold resistance and burial for winter survival and spring initiation of an Ulva spp. (Chlorophyta) bloom in a eutrophic lagoon (Veerse Meer lagoon, The Netherlands)

In the eutrophic Veerse Meer lagoon (The Netherlands) large amounts of free-floating thalli from Ulva spp. are present from May to October. In winter however, no algae seem to occur in the lagoon. Sexual reproduction appears to be negligible, as spore formation and germling growth are observed only sporadically. Results of a field survey showed that in winter, viable Ulva biomass is present buried in the sediment of the shallow parts of the lagoon. Freezing experiments demonstrated that the algae are able to survive temperatures of −5 °C for 2 weeks when kept in darkness. In spring, the buried Ulva thalli are liberated out of the sediment to initiate a bloom. A field experiment indicates that bioturbation by the lugworm Arenicola marina does not stimulate the release of the thalli. Burial and winter survival can explain the rapid increase in Ulva biomass in spring and suggest that the initial spring biomass is one of the major factors determining the maximal biomass in summer. Received: 28 October 1997 / Accepted: 20 January 1998     

Coral mortality following extreme low tides and high solar radiation

Extreme tidal events are one of the most predictable natural disturbances in marine benthic habitats and are important determinants of zonation patterns in intertidal benthic communities. On coral reefs, spring low tides are recurrent disturbances, but are rarely reported to cause mass mortality. However, in years when extremely low tides coincide with high noon irradiances, they have the potential to cause widespread damage. Here, we report on such an event on a fringing coral reef in the central Great Barrier Reef (Australia) in September 2005. Visual surveys of colony mortality and bleaching status of more than 13,000 corals at 14 reef sites indicated that most coral taxa at wave-protected sites were severely affected by the event. Between 40 and 75% of colonies in the major coral taxa (Acropora, Porites, Faviidae, Mussidae and Pocilloporidae) were either bleached or suffered partial mortality. In contrast, corals at wave-exposed sites were largely unaffected (<1% of the corals were bleached), as periodic washing by waves prevented desiccation. Surveys along a 1–9 m depth gradient indicated that high coral mortality was confined to the tidal zone. However, 20–30% of faviid colonies were bleached throughout the depth range, suggesting that the increase in benthic irradiances during extreme low tides caused light stress in deeper water. Analyses of an 8-year dataset of tidal records for the area indicated that the combination of extended periods of aerial exposure and high irradiances occurs during May–September in most years, but that the event in September 2005 was the most severe. We argue that extreme low-tide, high-irradiance events are important structuring forces of intertidal coral reef communities, and can be as damaging as thermal stress events. Importantly, they occur at a time of year when risks from thermal stress, cyclones and monsoon-associated river run-off are minimal.     

Interactions between an invasive and a native bryozoan (<Emphasis Type="Italic">Membranipora membranacea</Emphasis> and <Emphasis Type="Italic">Electra pilosa</Emphasis>) species on kelp and <Emphasis Type="Italic">Fucus</Emphasis> substrates in Nova Scotia,Canada

Most research on biological invasions to date has focused on the population dynamics of very successful and disruptive introduced species; however, additional knowledge of the biology of the native species is essential for understanding interactions between the two and may reveal factors that limit invasion success. The invasive bryozoan Membranipora membranacea interacts with native Electra pilosa on two substrates in northwest Atlantic subtidal habitats: highly dynamic and fast-growing kelps; and smaller, more stable, and slow-growing fucoid algae. We quantified the relative abundance and evaluated encounter outcomes in different seasons of these two bryozoans on kelp and Fucus at four sites in Nova Scotia. We also examined the effects of substrate (kelp, Fucus), temperature (7, 10, 13°C), and food (limited, unlimited) on growth rates of E. pilosa in laboratory experiments and using field manipulations. We compared our findings on factors affecting the growth of E. pilosa to those on M. membranacea obtained in similar and thus directly comparable experiments from a previous study. The proportional abundance of M. membranacea was greater than that of E. pilosa on kelps, but the opposite was observed on Fucus. Competitive standoffs between the two bryozoans were more frequent than expected, with no differences recorded between substrates; most encounters were won by M. membranacea. Growth of E. pilosa was faster on Fucus than kelp, decreased with increasing temperature only on Fucus, and was not affected by food. Growth rate of E. pilosa in all treatments was slower than that previously measured for M. membranacea. Faster growth and strong overgrowth abilities likely interact on kelps to ensure success of the invasive bryozoan. Success can be limited by low space availability, which in turn restricts growth rate, and consequently, colony size, such as on fucoid substrates. The incorporation of alternative contexts into invasion research can reveal factors involved in the resilience of native communities.     

Carbon emission associated with respiration and calcification of nine gastropod species from the intertidal rocky shore of Western Europe

Intertidal rocky shores are characterized by vertical zonation that results from the interplay between environmental conditions, organism physiology, and species interactions. Metabolism of intertidal organisms is highly variable between species and it changes with vertical position along the intertidal gradient. The present study aimed to quantify the carbon metabolism of nine intertidal rocky shore gastropods, in order to clarify their respective roles in carbon production during emersion and immersion. The influences of monthly temperature variation and tidal level were tested for each species. Analyses were performed in the laboratory using the infrared gas analyzer method for measuring aerial respiration rates, and the dissolved inorganic carbon and total alkalinity technique for measuring aquatic respiration rate and calcification. Hourly carbon fluxes were calculated for the mean annual temperature of 13 °C measured in both air and underwater in the study area. Respiration rates were similar for emersion (8–25 μmol CO₂ g AFDW^?1 h^?1) and immersion (10–23 μmol DIC g AFDW^?1 h^?1). For all species, underwater respiration fluxes were more influenced by monthly temperature variation than by air fluxes, probably as an adaptation to the rapid changes occurring during emersion. Calcification was an important factor influencing annual carbon fluxes for all studied species; every species showed different calcification rates according to its size and position on the intertidal zone. Annual carbon emissions were calculated using the mean immersion/emersion time of each species. Intertidal gastropod carbon emission was primarily influenced by body biomass and their vertical position within the intertidal zone.     

Diurnal and lunar patterns of larval recruitment of Brachyura into a mangrove estuary system in Ranong Province, Thailand

The present study followed the temporal recruitment pattern of brachyuran larvae in a mangrove tidal creek on the Andaman Sea coast of Ranong Province, Thailand, based on the assumption that the processes governing recruitment are important for the overall population dynamics of mangrove brachyuran crabs. Plankton net samples were taken on five occasions: on two new moon spring tides, one waxing moon neap tide, one full moon spring tide and one waning moon neap tide during October and November 1997. In addition collectors for larval crab megalopae were employed every 3 days through one dry season and one wet season (March–October 1998). Both the plankton net samples and collector samples revealed four major brachyuran groups in three families: Ocypodidae, Grapsidae and Portunidae. The grapsid group was further separated into two morphotypes which were identified as Metaplax and sesarmid species. Identified group mean numbers per cubic metre were ocypodids 3.0, sesarmids 0.8 and Metaplax 0.5, while portunid megalopae were very scarce (≪0.1 m⁻³). Further analysis of plankton net samples showed that when considering the parameters date, depth, current direction and the diel cycle, Metaplax and ocypodids distribute differently in the tidal and lunar cycle. Metaplax recruitment dominates on flood tides and on bottom layers, followed by middle and surface layers. Conversely, ocypodid abundance varied significantly with date only. Notably recruitment was not dependent on the diel cycle for either group. The collector samples of megalopae showed that recruitment of ocypodids, Metaplax and sesarmids occurred on full and new moon spring tides, while portunid megalopae preferred to settle on full moon spring tides. Since tidal currents were related to the lunar cycle megalopa groups are also cross-correlated with tidal amplitude, except for the portunid group. It is concluded that megalopae recruit in a similar manner to what has been found in other regions of the world, except that the abundance of ocypodids and Metaplax is not influenced by the diel cycle. Received: 14 February 2000 / Accepted: 24 November 2000     

Growth patterns of Codium fragile ssp. tomentosoides in response to temperature,irradiance, salinity,and nitrogen source

Seasonal patterns of growth, reproduction, and productivity of Codium fragile spp. tomentosoides (van Goor) Silva were monitored at 3 locations in Rhode Island. Maximal growth occurred during the summer and was more significantly correlated with temperature than any other factor measured in this study. Multiple correlation models suggested an interaction between temperature, irradiance, and available nitrogen. Maximal reproduction occurred in late summer and early fall. The maximal productivity, based on harvested quadrats, was 2. 10 g dry weight m^-2 day^-1. A large amount (up to 87.3%) of the annual production entered the detrital food chain during the winter by fragmentation of the thallus. Culture studies examined the effects of temperature (6° to 30°C), irradiance (7 to 140 E m^-2 sec^-1), daylength (8 h light: 16 h dark to 24 h light: O h dark) and salinity (6 to 48) on growth. Differentiated thalli grew over a broad range of experimental conditions, with maximal growth at 24°C, 24 to 30 S, a minimal irradiance of 28 E m^-2 sec^-1, and 16 h daylength. The effect of increasing daylength was due to increased total daily irradiance rather than to a true photoperiodic effect. Undifferentiated sporelings survived and grew in a narrower range of environmental conditions than thalli. c. fragile spp. tomentosoides grew equally well with nitrate, nitrite, ammonium, and urea as a nitrogen source. The addition of NaHCO₃ stimulated growth at levels of 2.4 to 4.8 mM, suggesting an inorganic carbon limitation in static cultures. This study supports the hypothesis that the in situ seasonal growth pattern of c. fragile spp. tomentosoides is primarily due to the interaction of temperature and irradiance.     

Plasticity in the temporal organization of behaviour in the limpet <Emphasis Type="Italic">Cellana grata</Emphasis>

The behaviour of intertidal consumers is often tightly constrained to tidal movements, although activity patterns can vary within these constraints. Spatio-temporal variability in behaviour of a limpet, Cellana grata, was analysed over different tidal conditions (spring and neap tides) and during different times of the year (one summer and one winter) at sites in Hong Kong. Activity was generally dictated by tidal movements, being concentrated when animals were awash. Plasticity in behaviour was observed, with some limpets anticipating activity during the summer period and delaying activity during winter time. Limpets were active for a time equal, or slightly less, than time awash. As the time awash exceeded ~14–16 h, however, activity duration decreased. Within this general pattern, tidal variation as well as variation among times of the year was noted, with the lowest dependence on time awash being recorded during winter neap tides. Limpets showed a slight preference for being active during nighttime, which was particularly evident when animals were emersed during the summer period. Although the basic activity in C. grata is constrained to a specific temporal window, this limpet is able to modulate its foraging strategies and resting height, according to local, daily changes in environmental conditions.     

No zone is always greener: variation in the performance of<Emphasis Type="Italic"> Fucus gardneri</Emphasis> embryos,juveniles and adults across tidal zone and season

The responses of the different stages of an organisms life-cycle to different environments can have important implications for their persistence in those habitats. The marine intertidal zone is considered among the most stressful of all marine environments, because daily exposure at low tide results in daily fluctuations in a range of important environmental factors. In this study we examined the link between daytime exposure and the performance of embryos, juveniles and adults of the seaweed Fucus gardneri across zones and seasons on San Juan Island, Washington, USA. Specifically we asked: (1) whether three measures of performance for F. gardneri: growth rates, surface area of reproductive tissue and survivorship differed between high- and mid-intertidal zones, and among months throughout the year and, then, (2) whether total daytime exposure within each zone each month was a good predictor of variation in these measures of performance. At our study site, seasonal differences in the timing of low tides result in long periods each day during summer of very harsh conditions (exposure to the air, high temperatures and low humidity) compared to winter when exposure to the air during the day is virtually absent. Contrary to the expectation of lower performance in the high zone, we found no consistent differences between zones over the entire year in any measure of performance for any stage. For nearly all traits, there were significant differences among months and, importantly, significant interactions between zone and month. Thus, for each stage there were some months when thalli in the high zone performed better and some months when thalli in the mid-zone performed better. Regressions between total daytime exposure and performance each month indicated some stage-specific effects. Larger thalli appeared more strongly affected by the amount of exposure per month; there were significant negative relationships between total daytime exposure and (1) growth rates of medium and large adults in the high zone and (2) the surface area of reproductive tissue of medium adults in the high zone and small adults in the mid-zone. The lowest survivorship for all stages occurred in the 2–3 months that followed summer, suggesting a cumulative effect of long daytime exposure in summer on survivorship. Post-settlement survivorship of embryos was very low throughout the year, with only 2 out of 5395 seeded zygotes surviving to a visible size. One-month post-settlement survivorship ranged from 0% to 16.7%, and there were no differences in the survivorship curves between high- and mid-intertidal zones for each cohort. Although it is often assumed that the performance of intertidal species is lower in high-intertidal zones because of exposure to environmental extremes for longer, our results do not support this paradigm. Instead, they indicate a complex and fluctuating pattern of variation in the performance of different stages of the F. gardneri life-cycle throughout the year in different intertidal zones.Communicated by M.S. Johnson, Crawley     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Tidal and circadian activity rhythms in juvenile plaice,Pleuronectes platessa

Young plaice (Pleuronectes platessa L.), caught in the intertidal zone, exhibit a short-lived tidal rhythm when their activity is recorded in darkness in the laboratory. This tidal rhythm rapidly changes to one of circadian frequency both in darkness and in light-dark cycles. It is considered that the basic rhythm is circadian in nature, but that it can be entrained to keep in phase with the tides by some, as yet unknown, Zeitgeber present under tidal conditions.     

Abundance of estuarine crab larvae is associated with tidal hydrologic variables

Abundances of brachyuran megalopae and juveniles were measured throughout consecutive tidal cycles during six 2 to 3 d sampling periods in summer 1992, and associated with rates of change of tidal hydrologic variables in the Newport River Estuary. Current speeds and rates of pressure change fitted sinusoidal (tidal) models well; however, rates of salinity and temperature change did not. Analysis of plankton samples taken during spring and neap tides showed peak abundances during nighttime rising tides for all taxonomic groups: Callinectes sapidus, Uca spp., Xanthidae, and Pinnixa spp. megalopae, and Pinnotheres spp. juveniles. Megalopal and juvenile abundances from time-intensive sampling were related to rates of changes in the hydrologic variables using stepwise logistic regression. No hydrologic variable accounted well for the presence of Uca spp. megalopae. Megalopal presence was best predicted by current speed for Pinnixa spp. megalopae, and rates of changes in pressure for xanthid megalopae and Pinnotheres spp. juveniles, and salinity for C. sapidus megalopae. These variables might act as cues causing megalopae to ascend into the water column at a particular point in the flooding tide, and subsequently descend to or near the bottom prior to ebb flow. In this way, larvae which develop on the continental shelf or lower estuary undergo transport up the estuary by behaviorally altering their swimming activity and depth concurrent with tidal changes.     

Reproductive biology of a filter-feeding amphipod, Leptocheirus pinguis , with extended parental care

Leptocheirus pinguis (Stimpson, 1853) is a widely distributed, abundant, endobenthic amphipod that engages in extended parental care, i.e. females host their juveniles in their burrows for extended time periods. I examined reproduction and population biology of L. pinguis at mean low water (MLW) in muddy sediment in Lowes Cove, Maine, USA. Cores around individual burrows were taken monthly in 1994 and 1995, and four seasonal samples were taken at different tidal heights. During the major reproductive periods in spring/early summer and in the fall, females produced several consecutive broods and hosted growing offspring in their burrows. Juveniles remained in their mothers' burrows until they reached a length of 5 mm (approximately one-third adult size) or more. At the study site, the majority of amphipods in individual burrows were adult females. Following the main reproductive periods, subadult individuals were found in their own burrows, but densities did not increase following the reproductive period in spring/early summer 1994, probably because large numbers of L. pinguis emigrated via the water column between June and December. L. pinguis is an annual species. Many members of the cohort born in spring/early summer start reproducing in the fall, and survive until the following spring when they produce several broods. Members of the cohort born in the fall start reproducing the following spring and also produce several consecutive broods. Both the spring/early summer and fall cohorts die off after the major reproductive period in the following spring/early summer. High standing stocks of microphytobenthos occur in soft-bottoms at MLW, and I conclude that L. pinguis can engage in extended parental care there because its food is abundant year-round. The limited expandability and low stability of burrows in soft-bottoms at MLW do not permit long persistence of parent–offspring groups in L.␣pinguis. Received: 27 March 1997 / Accepted: 30 July 1997     

Life history of the sandy-beach amphipod Dogielinotus loquax (Crustacea: Dogielinotidae) from the outer coast of Washington,USA

Dogielinotus loquax Barnard, a common intertidal macroinvertebrate on exposed ocean beaches near Grays Harbor, Washington, USA, is distributed from mid to mean high tidal levels, with a mean density of 1 830 m^-2 in 1975–1976. Location of maximum density within the intertidal zone is related to surf intensity and varies seasonally. The species is iteroparous and has an average sex ratio of 1:1. Temperature constraints on growth and egg development rates apparently influence the timing of reproduction. Two dominant recruitments occur per year, one in early spring (the summer generation) and another in late summer (the overwintering generation). The sampling design and density estimates permitted estimation of the mortality rate for the summer generation. Shorebird predation is suspected to affect summer generation abundance. Dogielinotus loquax will soon be elevated to the type of a new genus, Proboscinotus (Bousfield and Tzvetkova, in press)Contribution no. 1271 from the Department of Oceanography, University of Washington; Seattle, Washington 98195, USA     

Characterizing temperate rocky shores from the perspective of an early juvenile snail: the main threats to survival of newly hatched Nucella emarginata

Mortality factors most likely to constitute substantial selective pressures for early juvenile gastropods on temperate rocky shores were identified by examining the vulnerability of hatchlings of an intertidal snail, Nucella emarginata, to heat stress, desiccation, and predation in 1992 and 1993. The highest temperature of substrata measured at tidal heights colonized by N. emarginata in Barkley Sound, British Columbia, Canada, was 28.5°C. This temperature was not lethal to hatchlings in laboratory tests. In laboratory and field desiccation experiments, all hatchlings died within 6 h of emersion. Early juveniles could not survive direct exposure to even moderate drying conditions for the duration of a low tide. Hence, intertidal microhabitats which dry up even for short periods during low tides would prove lethal. Of 45 intertidal animal species to which hatchlings were exposed in the laboratory, small decapod crustaceans were the only organisms to cause substantial hatchling mortality. Of these, Pagurus hirsutiusculus and Hemigrapsus nudus were by far the most abundant in the field, and are probably the only important predators of early juvenile N. emarginata at most sites. Total predator densities in the field were as high as 438 individuals m^–2, suggesting that predation pressure may be intense. Desiccation and predation by decapod crustanceans appear to be the most significant threats to early juvenile N. emarginata. These factors commonly occur on most temperate rocky shores and undoubtedly constitute major selective agent influencing population parameters and shaping life-history strategies and early juvenile traits of intertidal invertebrates.     

Monitoring spatio-temporal variation in molluscan grazing pressure in seasonal, tropical rock pools

In Hong Kong, low-shore rock pools support a patchy mosaic of different successional stages of crustose coralline algae. Echinoid and molluscan grazers are abundant in these low-shore rock pools and spatial and temporal variation in grazing pressure is thought to play an important role in structuring these assemblages. Direct records of graze marks using a modified wax-disc method were used to monitor spatial (different pools) and temporal (periods, 3 or 4 days each; spring vs. neap tides; summer vs. winter) variations in grazing pressure. Radula marks of Chlorostoma argyrostoma, Lunella coronata and Nerita albicilla were common in all pools. Grazing pressure was highly variable with tidal state and season but also within these two factors. Lunella and Nerita exerted highest grazing pressure at neap tides during summer, whilst Chlorostoma exerted highest grazing pressure in winter. Although overall patterns of temporal variation were similar and consistent in all pools, there was between pool variation in grazing pressure and individual species showed different patterns. Within the seasonal and tidal variations, large period-to-period differences were found for all species, illustrating small-scale temporal variation. Fine-scale spatial and temporal patterns of variation in grazing pressure of different species, therefore, contribute to the highly dynamic nature of these low-shore rock pools.     

Rhythmic activity responses of the fiddler crab Uca crenulata to artificial tides and artificial light

The fiddler crab Uca crenulata, from California (USA), was exposed to artificial tides in order to differentiate between the influence of a tidal cycle from that of a light-dark cycle on its locomotor activity. Most crabs could be well synchronized by tides, but the activity patterns of other crabs was merely exogenously reinforced. Under constant conditions, after tidal treatment, crabs showed bimodal or unimodal activity patterns. When a 24 h light-dark cycle and a 12.35 h tidal cycle acted as concurrent stimuli, their effectiveness in synchronizing the crabs' activity was variable, depending on the undividual. It is likely that crabs which tend to exhibit a unimodal activity pattern are more sensitive to a light stimulus, whereas crabs with a bimodal activity pattern respond preferably to the tides. Thus, responses of endogenous activity of U. crenulata to tides and light-dark cycles are similar, suggesting that the basic oscillarory mechanism for circadian and tidal activity may be the same.     

Growth and spawning of Urechis caupo (Echiura) in Bodega Harbor,California

Growth and spawning of the large, infaunal echiuran worm Urechis caupo Fisher and MacGinitie were studied at Bodega Harbor on the coast of central California, USA, from 1978 through 1981. In situ growth rates of marked worms were negatively related to initial size. Short-term, summer growth rates (volmo^–1) of small worms (<80 ml) were greater than longer-term growth rates measured over several seasons (asesonal). Size-frequency distributions of worms sampled from two sites also suggested a seasonal growth pattern with relatively fast spring-summer growth and slower winter growth. However, larger worms sometimes lost volume during in situ growth experiments, and the loss was most pronounced during short-term, summer growth periods. It is suggested that energy used in burrow construction may have contributed to volume loss during short-term growth experiments. In contrast, longer-term, aseasonal growth rates were nearly always positive, and indicated that reproductive size (about 56 ml) could be reached within about 1.5 yr of recruitment, and a large size (about 158 ml) could be reached within about 6 yr. A seasonal pattern of spawning was observed during three consecutive years, as indicated by ripeness indices (storage organ dry weight ÷ body wall dry weight). At least two spawning episodes occurred annually: ripe gametes that accumulated in the storage organs during the summer and fall were spawned during the winter; gametes that accumulated during late winter and early spring were spawned during the spring or early summer. Worms were spawned-out by mid-summer.     

Spatial and temporal variation of resource limitation in Chesapeake Bay

We report nutrient addition bioassays at 18 stations in Chesapeake Bay (USA) to assess resources limiting phytoplankton growth. Data were pooled from several sampling programs conducted from 1989 to 1994. Spatially, light and P limitation declined from low salinity regions to high salinity regions, as N limitation increased. This spatial pattern was driven primarily by freshwater inflows with high N/P and seawater inflows with low N/P. Seasonally, there was a marked progression of winter light limitation, spring P limitation, and summer N limitation at mesohaline and polyhaline stations. The seasonal pattern appeared to be caused by temperature, mixing, river discharge, and sediment P fluxes. At high salinity stations, we also observed winter N limitation (caused by DIN depletion prior to spring nitrate delivery), and at lower salinity stations there was fall P limitation (caused by reaeration of bottom sediments). At tidal fresh stations, turbidity and nutrient concentrations resulted in continuous light limitation, except at some stations in summer. Interannual decreases in light limitation and increases in N and P limitation appear to represent improvements in water quality. Received: 31 October 1997 / Accepted: 18 December 1998