Depth as a factor in abundance and size of juvenile penaeid shrimps in the absence of estuaries and marshes

Triweekly trawling around low tide during daylight at 1, 2, 3, and 4 m depths along north Kuwait Bay's extensive intertidal mudflats from 1 April 1985 through 9 December 1986 showed the catchability ofMetapenaeus affinis (H. Milne-Edwards, 1837),M. stebbingi Nobili, 1904,Parapenaeopsis stylifera (H. Milne-Edwards, 1837), andPenaeus semisulcatus De Haan, 1844 differed significantly with depth and season. All species were significantly more catchable in 1 and 2 m depths than in 3 and 4 m depths.M. affinis showed the clearest and most consistent relationship with depth: catchability in 1 m depths was significantly greater than that in 2 m depths (24.9 vs 5.4 per 5-min tow), and catchability in 1 and 2 m depths was significantly greater than that in 3 and 4 m depths (15.2 vs 1.4 per 5-min tow).M. affinis also showed the clearest relationship of increasing size with depth. Shrimp captured in 1 m depths were significantly smaller (~ 14 mm carapace length, CL) than those captured in 2 m depths in the spring (~23 mm CL) and summer (~ 18 mm CL). In the absence of low-salinity waters and intertidal vegetation, it is believed that the edge of the advancing and receding tide over the mudflats, i.e. the shallowest waters, provides juvenile shrimp, particularlyM. affinis, protection from predation.     

Seasonal variation of membrane fluidity of the naturally acclimatized Baltic Sea amphipods Gammarus spp. and Monoporeia affinis

The membrane fluidity of the Baltic Sea crustaceans Gammarus spp. and Monoporeia affinis was studied in different seasons. Gammarus spp. were collected at a location with stable salinity and with temperature fluctuations from about 0 to 20 °C, and M. affinis at a deep location with stable salinity and a constant low temperature of about 1.5 to 4.5 °C. The membrane fluidity was measured from preparations enriched with plasma and mitochondrial membranes employing a fluorescence polarization technique using 1,6-diphenyl-1,3,5-hexatriene (DPH) as a probe. The measurements were carried out without any preceding acclimation of the animals. In Gammarus spp., the membrane fluidity increased in the order: summer animals < autumn animals < spring animals. The differences between seasons were significant, with pronounced homeoviscous adaptation, and correlated well with the water temperature. It is suggested that in Gammarus spp., temperature is the determining factor for membrane fluidity. In M. affinis, the membranes of summer animals were the most fluid, and there was a significant difference only between summer and spring animals. In M. affinis the difference was not connected to the water temperature and no homeoviscous adaptation was noticed. The deep-water M. affinis experience a long period of fresh food deficiency, which probably affects the membrane fluidity. Although there are seasonal differences in fluidities between Gammarus spp. and M. affinis, it is interesting to note that all data obtained from M. affinis settled between the polarization lines of spring and summer Gammarus spp., regardless of the prevailing temperatures. Received: 30 June 1999 / Accepted: 7 June 2000     

Deep-water decapod crustacean communities in the Northwestern Mediterranean: influence of submarine canyons and season

The specific composition and abundance of bathyal decapods in the Catalan Sea were investigated. A total of 109 bottom trawls were effected at depths ranging from 141 to 730 m on the continental slope in the Catalan Sea (northwestern Mediterranean) during two sampling cruises in spring and autumn 1991. Multivariate analysis of the samples revealed four groups of the decapod crustacean communities: (1) A shelf-slope transition-zone group at depths between 146 and 296 m, primarily characterized by the presence of Plesionika heterocarpus; (2) an upper-slope community between 245 and 485 m, characterized by the presence of the mesopelagic species Pasiphaea sivado and Sergestes arcticus, with Processa nouveli, Solenocera membranacea and Nephrops norvegicus as secondary species; (3) a middle-slope community below 514 m, with Aristeus antennatus and Calocaris macandreae as the most abundant species; (4) a group at 430 to 515 m, comprising all samples collected exclusively within or in the vicinity of submarine canyons. Mesopelagic decapods were predominant on the slope, while benthopelagic fishes (Merluccius merluccius, Micromesistius poutassou, Gadiculus argenteus) replaced mesopelagic decapods on the shelf. There were seasonal variations, with higher densities of mesopelagic species in spring, which were probably related, among other factors, to variations in the photoperiod. Our surveys also revealed higher species richness in the canyons together with seasonal changes in the megafaunal biomass. Generally, the upper and middle-slope communities both displayed seasonal changes in the composition and abundance of megabenthos.     

Temporal succession and spatial segregation of clupeoid larvae in the coastal waters off the Tanshui River Estuary, northern Taiwan

In the coastal waters off the Tanshui River Estuary in northern Taiwan, eight species of clupeoid larvae were observed. They exhibited a distinct temporal succession in association with seasonal temperature changing. The time of peak abundance of Etrumeus teres was in January, Engraulis japonicus in late April, Sardinella spp. in mid-May, Dussumieria elopsoides in early June, Thryssa dussumieri in late June, Stolephorus insularis in mid-September, Encrasicholina heteroloba in early October and E. punctifer in mid-November. The time intervals of the temporal succession of the fishes were approximately 15 to 25 d in the spring/summer and 25 to 35 d in the autumn/winter fishing seasons. Also, they showed spatial segregation by distributing in areas with different water depths: Sardinella spp. at a water depth of 10 to 20 m, T. dussumieri at less than 10 m, E. heteroloba at 20 to 50 m and E. punctifer at 10 to 40 m. The larvae of these sympatric clupeoid species segregated their nursing periods and areas apparently to reduce competition for habitat and thus to maximize the utilization of resources. Received: 3 March 1997 / Accepted: 19 March 1997     

Dominance behaviour,body weight and fat variations,and partial migration in European blackbirds Turdus merula

Summary A partially migratory population of European blackbirds Turdus merula was studied in central Europe during the non-breeding season. Body weight and the amount of subcutaneous fat deposits showed seasonal variations typical for each age and sex category. Juveniles were fattest in autumn and spring and both juveniles and females lost fat and body weight in mid-winter. Adults and males reached their maximum body weight in mid-winter. These findings together with data from Schwabl (1983) suggest that juveniles and females constitute the main fraction of those birds migrating from the area in autumn. In winter mainly adults and males have access to food resources. These patterns may be regulated by dominance behaviour, since adults and males are usually dominant over juveniles and females throughout the non-breeding season. Subordinate birds that stay in the area probably suffer from a higher winter mortality but have the advantage of early breeding.     

Growth and breeding of meso- and bathypelagic organisms of the Rockall Trough,northeastern Atlantic Ocean and evidence of seasonality

Seasonality of growth and reproduction of a variety of epimeso-and bathypelagic organisms living in the Rockall Trough is examined in samples collected between 1973 and 1978. Rates of growth change seasonally. Epipelagic and shallow-living mesopelagic migrators tend to grow most rapidly in the spring and summer while deeper-living mesopelagic migrators tend to accelerate their growth in the summer and autumn when seasonal water temperatures at these depths are at their seasonal maxima. Mesopelagic migrators all breed seasonally. Rates of growth of bathypelagic non-migrating species could not be determined through analyses of body length/frequency histograms because of the continuous (aseasonal) breeding and recruitment of juveniles to the adult populations. An exception was the mysid crustacean Boreomysis microps that had a seasonal period of spring recruitment of juveniles to the population. Several of the mesopelagic crustaceans and fish have extended periods of approximately linear growth in body length. There is no evidence that organisms living in the mesopelagic or bathypelagic environments have rates of growth that are markedly slower than those of congeners in shallower environments.     

Behavioural response to gradually declining oxygen concentration by Baltic Sea macrobenthic crustaceans

Behavioural responses to decreasing oxygen concentrations were studied in the amphipods Monoporeia affinis (Lindström) and Pontoporeia femorata Krøyer and the isopod Saduria entomon (L.), all benthic macrofauna species from the Baltic Sea. Infrared time-lapse video recording was used. Seven oxygen concentrations were studied, starting near saturation (about 90%), followed by about 65, 49, 33, 16, 8 and 5% oxygen saturation, with each oxygen level maintained for 24?h. M. affinis was more active than P. femorata. The duration of swimming activity of M. affinis was shorter at ≤33% oxygen saturation compared with at the two highest levels. The duration of swimming by P. femorata was longest at 90, 65 and 5% oxygen saturation. For both amphipods the frequency of swimming activity was highest at 5% oxygen saturation. Above 7% oxygen saturation, S. entomon spent most of its time in the sediment, whereas at levels ≤7% it was most often found on the sediment surface. Both amphipods were predominantly night active (nocturnal), whereas no diurnal variation in behaviour was detected for S. entomon. It is suggested that the amphipods minimise the costs associated with obtaining oxygen as well as their risk of predation by moving little on or above the sediment until a lower critical oxygen level is reached. Below this level, they show an oxygen-seeking behaviour. S. entomon also hid in the sediment for as long as possible, suddenly coming to the surface once the oxygen concentration became intolerably low. Its behaviour is also indicative of a trade-off between predation risk and obtaining sufficient oxygen. It is suggested that the change in behaviour shown at about 33% oxygen saturation could lead to a long-term alteration in community structure in the deeper parts of the northern Baltic proper, whereas the change at about 8% oxygen saturation reflects a more acute, short-term response.     

Cephalopod hatchling growth: the effects of initial size and seasonal temperatures

Temperature is known to have a strong influence on cephalopod growth during the early exponential growth phase. Most captive growth studies have used constant temperature regimes and assumed that populations are composed of identically sized individuals at hatching, overlooking the effects of seasonal temperature variation and individual hatchling size heterogeneity. This study investigated the relative roles of initial hatchling size and simulated natural seasonal temperature regimes on the growth of 64 captive Octopus pallidus over a 4-month period. Initial weights were recorded, and daily food consumption and fortnightly growth monitored. Two temperature treatments were applied replicating local seasonal water temperatures: spring/summer (14–18°C) and summer/autumn (18–14°C). Overall octopuses in the spring/summer treatment grew at a rate of 1.42% bwd⁻¹ (% body weight per day) compared to 1.72% bwd⁻¹ in the summer/autumn treatment. Initial size influenced growth rate in the summer/autumn treatment with smaller octopuses (<0.25 g) growing faster at 1.82% bwd⁻¹ compared to larger octopuses at 1.68% bwd⁻¹. This was opposite to individuals in the spring/summer treatment where smaller octopuses grew slower at 1.29% bwd⁻¹ compared to larger octopuses at 1.60% bwd⁻¹. Initial size influenced subsequent growth, however, this was dependent on feeding rate and appears to be secondary to the effects of temperature.     

Food and feeding habits of the king crab Paralithodes camtschatica near Kodiak Island,Alaska

Stomach contents from 809 king crabs, Paralithodes camtschatica (Tilesius), from 6 areas near Kodiak Island, Alaska, and 9 sampling periods (1978–1979) were exammed quantitatively; 713 (88%) contained food. Mollusca (mainly the bivalves Nuculana spp., Nucula tenuis, and Macoma spp.) and Crustacea (mainly barnacles) were the dominant food groups in terms of percentage wet weight and frequency of occurrence; fishes were the next most important group of prey. No significant differences in feeding between sexes occurred; however, significant differences were apparent in the quantity of food consumed from different sampling periods, areas, depths, size groups, and crab molt-classes. Consumption was greater in spring and summer and in offshore locations at depths of 126 to 150 m. In addition, king crabs <140 mm carapace length (CL) consumed more food than crabs 140 mm CL. Adult, newshell (individuals that molted during the last molting period) females greater than 95 mm CL, and newshell males greater than 100 mm CL, each contained more food than did juvenile, newshell females <120 mm CL.Contribution No. 449, Institute of Marine Science, University of Alaska, Fairbanks, Alaska 99701, USA     

Rates of primary productivity and growth in<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> measured at different depths,over an annual cycle,at West Island,South Australia

The pattern of growth (biomass accumulation) in Ecklonia radiata throughout the year and across a depth profile was investigated using the traditional hole-punch method, and the information presented in context with concurrently measured in situ net productivity rates. The rate of net daily productivity showed a lack of consistent seasonal variability, remaining constant throughout the year at two of the four depths measured (3 m and 12 m), and becoming higher during winter at another (5 m). Throughout the year, rates of net daily productivity differed significantly across the depth profile. Net daily productivity rates averaged 0.017 g C g^–1 dwt day^–1 and 0.005 g C g^–1 dwt day^–1 at a depth of 3 m (1,394 mol O₂ g^–1 dwt day^–1) and 10 m (382 mol O₂ g^–1 dwt day^–1) respectively. In contrast, the biomass accumulation rate of E. radiata was highly seasonal, with low rates of growth occurring in autumn (0.002 g dwt g^–1 dwt day^–1 at both 3 and 10 m) and summer (0.007 and 0.004 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively) and higher rates in spring (0.016 and 0.007 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively) and winter (0.015 and 0.008 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively). The proportion of assimilated carbon used for biomass accumulation varied throughout the year, between 5% and 41% at 3 m and between 28% and 128% at 10 m. The rates of biomass accumulation at all depths represented only a small proportion of the amount of carbon assimilated annually.Communicated by P.W. Sammarco, Chauvin     

Diurnal patterns in photosynthetic capacity and depth-dependent photosynthesis-irradiance relationships inSynechococcus spp. and larger phytoplankton in three water masses in the Northwest Atlantic Ocean

The photosynthetic characteristics of prokaryotic phycoerythrin-rich populations of cyanobacteriaSynechococcus spp. and larger eukaryotic algae were compared at a neritic frontal station (Pl), in a warm-core eddy (P2), and at Wilkinson's Basin (P3) during a cruise in the Northwest Atlantic Ocean in the summer of 1984.Synechococcus spp. numerically dominated the 0.6 to 1 m fraction, and to a lesser extent the 1 to 5 m size fractions, at most depths at all stations. At P2 and P3, all three size categories of phytoplankton (0.6 to 1 m, 1 to 5 m, and >5 m) exhibited similar depth-dependent chages in both the timing and amplitude of diurnal periodicities of chlorophyllbased and cell-based photosynthetic capacity. Midday maxima in photosynthesis were observed in the upper watercolumn which damped-out in all size fractions sampled just below the thermocline. For all size fractions sampled near the bottom of the euphotic zone, the highest photosynthetic capacity was observed at dawn. At all depths, theSynechococcus spp.-dominated size fractions had lower assimilation rates than larger phytoplankton size fractions. This observation takes exception with the view that there is an inverse size-dependency in algal photosynthesis. Results also indicated that the size-specific contribution to potential primary production in surface waters did not vary appreciably over the day. However, estimates of the percent contribution ofSynechococcus spp. to total primary productivity in surface waters at the neritic front were significantly higher when derived from short-term incubator measurements of photosynthetic capacity rather than from dawn-to-duskin situ measurements of carbon fixation. The discrepancy was not due to photoinhibitory effects on photosynthesis, but appeared to reflect increased selective grazing pressure onSynechococcus spp. in dawn-to-dusk samples. Low-light photoadaptation was evident in analyses of the depth-dependency ofP-I parameters (photosynthetic capacity,P _max; light-limited slope, alpha;P _max alpha,I _k ; light-intensity beyond which photoinhibition occurs,I _b ) of the > 0.6 m communities at all three stations and was attributable to stratification of the water column. There was a decrease in assimilation rates andI _k with depth that was associated with increases in light-limited rates of photosynthesis. No midday photoinhibition ofP _max orI _b was observed in any surface station. Marked photoinhibition was detected only in the chlorophyll maximum at the neritic front and below the surface mixed-layer at Wilkinson's Basin, where susceptibility to photoinhibition increased with the depth of the collected sample. The 0.6 to 1 m fraction always had lower light requirements for light-saturated photosynthesis than the > 5 m size fraction within the same sample. Saturation intensities for the 1 to 5 m and 0.6 to 1 m size fractions were more similar whenSynechococcus spp. abundances were high in the 1 to 5 m fraction. The > 5 m fraction appeared to be the prime contributor to photoinhibitory features displayed in mixed samples (> 0.6 m) taken from the chlorophyll maxima. InSynechococcus spp.-dominated 0.6 to 1 and 1 to 5 m size fractions, cellular chlorophylla content increased 50- to 100-fold with depth and could be related to increases in maximum daytime rates of cellularP _max at the base of the euphotic zone. Furthermore, the 0.6 to 1 m and > 5 m fractions sampled at the chlorophyll maximum in the warm-core eddy had lower light requirements for photosynthesis than comparable surface samples from the same station. Results suggest that photoadaptation in natural populations ofSynechococcus spp. is accomplished primarily by changing photosynthetic unit number, occuring in conjuction with other accommodations in the efficiency of photosynthetic light reactions.     

Stable isotope analysis of two species of anguilliform leptocephali (<Emphasis Type="Italic">Anguilla japonica</Emphasis> and <Emphasis Type="Italic">Ariosoma major</Emphasis>) relative to their feeding depth in the North Equatorial Current region

Anguilla japonica leptocephali are transported from their offshore spawning area to their recruitment areas in East Asia, but their depth distributions, food sources and feeding are still poorly known. This study analyzed carbon and nitrogen stable isotope ratios of leptocephali of A. japonica, Ariosoma major and Ariosoma spp., and of particulate organic matter (POM), their likely food source, at five different depths in 2004–2009. We used mixing models to show that A. japonica appeared to be feeding at depths between 5 and 50 m, but sometimes deeper. A. major appeared to have a tendency of mostly feeding at depths of 50 m or shallower. Although the A. japonica and Ariosoma spp. collected in the same area during the leptocephalus stage appeared to have different feeding ecologies possibly related to different types of POM, their different depth distributions, sizes and transport histories may also help explain these differences.     

Seasonal occurrence of the ctenophore Pleurobrachia pileus in the western Dutch Wadden Sea

Due to the short residence time of the water in the western Dutch Wadden Sea the ctenophore Pleurobrachia pileus has to be considered as an allochthonous species, which enters the estuary passively from the North Sea. The seasonal occurrence of this species in the Dutch Wadden Sea is a reflection of its abundance in the coatal zone of the North Sea and is characterized by a pronounced spring peak of up to 17000 individuals per 1000 m³ and a lower autumn peak with numbers up to 400 individuals per 1000 m³. Similar to other areas, predation by Beroë gracilis, another ctenophore species, may be a factor in controlling the P. pileus population in this region, although B. gracilis abundance is low. P. pileus is only important as a predator for a short time, i.e. the month of May, when it reaches maximum biomass values of up to 12 00 mgC per 1000 m³. Both the estimated potential consumption rates in comparison with the available food as well as the observed suboptimal growth rate of the population suggest that in the coastal zone of the North Sea area food conditions may not be optimal for P. pileus.     

The spiny lobster Palinurus elephas in Scottish waters

Observations have been made on the biology of spiny lobsters, Palinurus elephas Fabr., caught by divers on the west coast of Scotland from 1972 to 1975. P. elephas in this Scottish population show the same length/carapace length relationship as spiny lobsters taken from the west of Ireland. It occurs inshore during the period April to October on shallow rocky reefs in depths of 5 to 20 m. In females, a pre-mating moult occurs in the inshore areas in late summer. Soon after, mating takes place and the spermatophores are deposited on the sternum of the female. The eggs, which are shed within 1 week to 10 days, are then carried by the female throughout the winter, and hatch in the following spring. Offshore migration takes place in the autumn, after mating and egg laying, and inshore migration takes place in the spring and summer.     

Effect of light and turbulent mixing on the growth of<Emphasis Type="Italic"> Skeletonema costatum</Emphasis> (Bacillariophyceae)

Skeletonema costatum was grown in an outdoor mesocosm to test the hypothesis that fluctuations in irradiance brought about by changes in mixing time and depth can reduce diatom growth and biomass in the turbulent mixed layer. The light environment and mixing regime within the mesocosm were comparable to those in shallow lakes and coastal waters. Experiments showed no significant differences for 24-h mean and 7-day mean chlorophyll a and carbon-specific growth for mixed depths of 1 m and 3 m, and mixing times between 4 min and 65 min. Fluctuations in irradiance brought about by turbulent mixing had no significant effect on specific growth. The relationship between mixed depths and critical depths for S. costatum was therefore independent of fluctuations in irradiance in the turbulent mixed layer. The results indicated that to control growth of S. costatum mixed depths would have to exceed photic depths by a factor of 15, instead of the conventionally accepted factor of 5. Thus, it is likely that artificial mixing of shallow (<10 m) eutrophic waters will be more effective in controlling slow-growing summer biomass than fast-growing spring blooms dominated by diatoms.Communicated by P.W. Sammarco, Chauvin     

Natural diet of larval Penaeus merguiensis (Decapoda: Penaeidae) and its effect on survival

The relationship between Penaeus merguiensis protozoea larvae and their phytoplankton diet was examined using seasonal plankton surveys and in situ rearing experiments. Larval abundance, phytoplankton community structure, and chlorophyll a concentration in Albatross Bay, Gulf of Carpentaria, were monitored monthly for 2 yr. Larval abundance peaked in November (spring) and March (autumn), at which times diatoms were the most abundant group in net samples of phytoplankton and in the guts of larvae. During November 1989 and March 1990, larvae were reared in nylon mesh enclosures positioned throughout the water column at three depths: 0 to 3 m, 3 to 6 m and 6 to 9 m. Overall, larval survival and gut fullness were both higher in November than in March. In both months, larval survival was lower at the surface than at other depths. This correlated with lower chlorophyll a concentrations, but lower total cell densities were not detected. During the in situ experiments, diatoms were the most abundant phytoplankton group in the water column and in the guts of larvae and, therefore, appeared to be the principal diet of larvae. Pigment analysis demonstrated that while gut contents generally reflected the composition of the phytoplankton community, the larvae were not feeding exclusively on diatoms. They also ingested green algae and possibly seagrass detritus. The in situ experiments demonstrated that the predominantly diatom flora in Albatross Bay can provide a nutritionally adequate environment for prawn larvae even at seasonally low levels. It is unlikely, therefore, that starvation is a major cause of mortality of P. merguiensis larvae during either of the biannual peaks in their abundance in Albatross Bay, Gulf of Carpentaria.     

Effects of season and maternal food on<Emphasis Type="Italic"> Calanus finmarchicus</Emphasis> reproduction,with emphasis on free amino acids

Effects of maternal food environment and season were examined during spring and autumn on females, eggs and nauplii of Calanus finmarchicus, in different natural prey suspensions or cultures of Rhodomonas baltica. Females sampled in spring were in general larger, had higher protein content, and showed higher egg production and hatching rates, than in autumn. The cumulative egg production was almost double in spring compared to autumn (females fed R. baltica). Females had higher content of free amino acids (FAA) and free essential amino acids (EAA) in autumn than in the spring. Also, the FAA contents in eggs and nauplii were higher in autumn than in spring. In contrast, the composition of EAA in eggs was constant between seasons, indicating maternal regulation. The highest cumulative egg production was correlated with a high similarity in the free pool of EAA in the food suspension and the female copepod. Thus, the data support the hypothesis that similarity in the free pool of EAA of diet and female promotes high fecundity and egg hatching success in C. finmarchicus.     

Spatio-temporal changes in diversity and community structure of planktonic copepods in Sagami Bay, Japan

Seasonal changes in diversity and community structure of planktonic copepods at a shelf site in Sagami Bay, Japan was studied in relation to cross-shelf interaction of species components. Seasonal mesozooplankton samples were collected from the shelf station (St. M) of the north-west part of Sagami Bay from 1995 to 1997. Vertical multi-layered samples were collected near the center of Sagami Bay (St. P) in June 1996. A total 185 copepod species were identified from the two stations. We observed a clear seasonal succession in calanoid diversity and community structure at St. M from a simple shelf water community (>11 species) during spring blooming periods to highly diverse and mixed communities (ca 20–30 species) of shelf water species coupled with various Kuroshio Current species during late summer to autumn. Cluster and non-metric multidimensional scaling ordination analyses showed two distinct calanoid community groups. One group, which included samples of St. M and the surface layer of St. P, consisted of shelf water species, such as Calanus sinicus, Ctenocalanus vanus, Paracalanus spp., and Kuroshio species, such as, Canthocalanus pauper, Scolecithrix danae, etc. The other cluster was restricted to the samples collected from mid and deep layers at St. P, which consisted of meso- and bathypelagic species and Oyashio species (cold-current species, such as Neocalanus cristatus, Pseudocalanus spp., Eucalanus bungii and Metridia pacifica). In the mid and deep layers at St. P, the population of dormant copepodid stage V (CV) of Eucalanus californicus and C. sinicus were dominant. The deep CV population of C. sinicus might be ecologically discriminated from the surface and shelf water population due to their larger body length and dormant life cycle. E. californicus was also collected at the shelf site during each spring bloom period, whereas the population might descend into the mid- and deep-layers of the central bay before summer. Our results suggest that the seasonal fluctuation of community structure in the shelf water was controlled by both physical (Kuroshio Current) and biological factors, i.e., spring bloom and ontogenetic vertical migration of E. californicus. In particular, transport and diffusion processes of Kuroshio Current in Sagami Bay played a key role in controlling the shelf water calanoid community.     

Skeletal growth zones as age-markers in the sea urchinPsammechinus miliaris

Growth of the inshore sea urchinPsammechinus miliaris (Gmelin) was studied in Loch Creran, western Scotland, using the skeletal growth-marker tetracycline in order to test the validity of natural growth-banding in the coronal test plates as annual age-markers. In order to test whether tetracycline affected the growth ofP. miliaris, an injected and a control group of urchins were held in identical conditions in running sea water aquaria for 21 mo from 1989 to 1991 and measured periodically. A small but significant difference in mean size of injected compared to controls was recorded at 12 mo, but none during subsequent measurements. Size measurements during the trial were consistent with an annual growth cycle, with a maximum in spring and slowing or cessation of growth during autumn/winter. Tetracycline-labelled juveniles were recovered up to 18 mo after initial tagging in mark/recapture experiments undertaken from 1987 to 1989 at two intertidal marked quadrats in Loch Creran. Large numbers ofP. miliaris were also marked with tetracycline and held for 1 yr at 10 m depth in seabed cages in Loch Creran during 1988–1990. All of the intertidal recoveries, and about 69% of the caged specimens that had been successfully labelled, showed a consistent relationship between the position of the tetracycline tag and the pattern of natural growth zones. The remainder were mostly large, slow-growing urchins with the tag positioned near the plate margin. In these the outer growth bands were closely spaced and, particularly if major growth bands seemed to be broken up into double or multiple lines, the major bands were impossible to resolve at the margin. The results support the assumption that in wild populations the dark bands visible with reflected light (translucent in transmitted light) after charring the plate are formed when skeletal growth has stopped or slowed in winter. These lie between wider, lighter coloured (opaque in transmitted light) zones of active plate growth in spring/summer. The dark band formed beyond the tag usually was made up of several closely spaced fine lines, or sometimes of two closely spaced dark bands. The wide growth zones beyond the tag, like those formed previously, usually were broken by fine, dark lines that may represent brief discontinuities in growth. From tagging, the double dark bands can be related to growth over one year; but such anomalous bands, along with the general presence of fine, dark lines interrupting the growth zones, make it difficult reliably to estimate age from the closely spaced peripheral banding on older, slow-growing urchins.