Natural growth bands and growth variability in the sea urchin Echinus esculentus: results from tetracycline tagging

Growth of the European edible sea urchin Echinus esculentus L. was studied in a population held for 2 yr in cages on the sea bed, after labelling with the skeletal growth marker tetracycline. The final position of the tetracycline tag on the genital plates agreed with an annual periodicity in natural growth zones; two such growth zones appeared beyond the position of the tag on the ground surface of the plate as light-reflecting bands separated by narrow dark lines in the middle layer. Individual and group (pooled data) growth parameters were estimated from the growth increment shown in the genital plate, whose lateral growth displayed a linear relationship to the diameter of the urchin test within the size range of these measurements. Von Bertalanffy growth parameters (asymptotic size and growth-rate function K) fitted to the growth increment on each individual were highly significantly correlated to those fitted to the natural growth lines, assuming an annual periodicity. The inferred growth pattern agrees well with conclusions based on H. B. Moore's growth-band data. The caged urchins can be assumed to have experienced exactly similar conditions, yet the growth curves fitted to individuals showed considerable variability. The good agreement between estimated growth function parameters of individuals obtained by the two methods indicate that this reflects real variability in growth between individuals that probably is genetically rather than environmentally determined. The growth of E. esculentus, and the adaptational significance of high growth variability in the population is briefly discussed.     

Stability of sea urchin dominated barren grounds following destructive grazing of kelp in St. Margaret's Bay,Eastern Canada

Kelp regeneration was observed for the first time in St. Margaret's Bay, Nova Scotia, Canada in an area known to have been devoid of macroalgae for several years. The regeneration was destroyed by sea urchins within 10 months. Experimentally induced kelp regeneration met a similar fate under normal grazing pressure. At the lowest sea urchin biomass and density encountered re-establishment of mature kelp stands seems highly unlikely. The sea urchin population required to suppress kelp regeneration is fed by benthic microalgae. Diatoms and other pioneer algal community species were found in the guts of sea urchins. The mean standing crop of benthic microalgae was found to be, 2.2 g C m^-2 and production estimated as ca 15 g C m^-2yr^-1 at 8m depth. Most of the primary production of St. Margaret's Bay has been lost with the disappearing kelp populations.     

Growth,mortality and yield-per-recruit of deep-water royal red prawns (Haliporoides sibogae) off eastern Australia,using the length-based MULTIFAN method

The MULTIFAN method of Fournier et al. was used to analyse a series of monthly length-frequency distributions of deep-water royal red prawns (Haliporoides sibogae de Man, 1907, Solenoceridae) for the estimation of growth, mortality and yield-per-recruit. Length data were collected from commercial catches made between Latitude 33° and 35°S from November 1988 to May 1990. MULTIFAN distinguished six cohorts in the royal red prawn population, aged at 6 mo intervals from about 1.5 to 4 yr. Most prawns were between 2 and 3 yr of age. Females and males were fully recruited at 23.1 and 21.5 mm carapace length (CL) respectively, when about 2 yr old. Females grew faster and reached larger sizes at age than males (growth coefficient, K=0.37 yr^-1 and asymptotic length, L =48.3 mm CL for females, and K=0.49 yr^-1 and L =33.5 mm CL for males). Total mortality (Z) was estimated at 1.6 and 1.2 yr^-1 for females and males, respectively. Approximate values for natural mortality (M) ranged from 0.6 to 0.7 yr^-1 for females, and from 0.4 to 0.8 yr^-1 for males. Analysis of yield-per-recruit suggested that while the current yield could possibly be increased by decreasing the length at first capture (L _c ), there seems to be little potential for an increase in yield from increased fishing effort in the area currently fished.     

Effects of grazing by two species of sea urchins (Strongylocentrotus franciscanus and Lytechinus anamesus) on recruitment and survival of two species of kelp (Macrocystis pyrifera and Pterygophora californica)

We studied the effects of grazing by two species of sea urchins on two species of kelp (Macrocystis pyrifera and Pterygophora californica) in the San Onofre kelp bed in southern California from 1978 through 1981. Both red sea urchins, Strongylocentrotus franciscanus, and white sea urchins, Lytechinus anamesus, were abundant and lived in aggregations. The purple sea urchin (S. purpuratus) was rare at the study site and was not studied. The aggregations of red urchins were either relatively small and stationary (for over 3 yr) or relatively large and motile (advancing at about 2 m mo^–1). Both stationary and moving aggregations were observed at the same time, and within 100 m of one another. Stationary aggregations of red urchins probably subsisted mainly on drift kelp and had no effect on kelp recruitment or on adult kelp abundance. In contrast, red sea urchins in large, motile aggregations or fronts ate almost all the macroalgae in their path. The condition of their gonalds indicated that red urchins in fronts were starved relative to red urchins in the small, stationary aggregations. Large, motile aggregations developed after 2 yr of declining kelp abundance (probably due largely to storms). We propose that a scarcity of drift algae for food results in a change in the behavior pattern of the red urchins and thus leads to the formation of large, motile aggregations. The aggregations of white urchins, which occurred along the offshore margin of the kelp bed, were large, but relatively stationary. The white urchins rarely ate adult kelps, but grazed extensively on early developmental stages of kelps and evidently prevented seaward expansion of the bed. The spatial distribution of both types of red urchin aggregations appeared to be unrelated to predation pressure from fishes or lobsters.Please address all requests for reprints to the senior author at his present address.     

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.     

Bioerosion in coral reef communities in southwest Puerto Rico by the sea urchin<Emphasis Type="Italic"> Echinometra viridis</Emphasis>

Bioerosion is one of the most important structuring forces in coral reef communities. The bioerosion impact of several species of fish, sponges and sea urchins have been estimated in the Caribbean; however, there is no information for one important species, the red sea urchin Echinometra viridis. This species can be found in high densities in many localities. In this study, bioerosion rates for E. viridis were estimated in two patch reefs off La Parguera, southwest Puerto Rico, using the population size-class distribution, average densities, and the CaCO ₃ content in fecal pellets produced over 24 h. Average densities of urchins along four depth intervals were estimated using 40-m transect lines and 1-m ² quadrats. Average size and size-structure distribution were estimated by measuring the diameter of 180–220 urchins haphazardly collected at each of the four depth intervals. The ignition–loss method was used to estimate the daily rate of bioerosion. Fecal pellets produced by the urchins over a 24 h period were collected in buckets, rinsed in fresh water, dried for 24 h at 70°C, and then burned in a furnace at 550°C, first to eliminate organics, and then at 1000°C until constant weight to determine the amount of calcium carbonate (CaCO ₃) in the fecal pellets. HCl (10%) was then added to the remainder of the sample to test for presence of CaCO ₃. Average individual CaCO ₃ bioerosion rates were estimated at 0.181±0.104 g day ^-1. Average densities (0.77–62.0 ind. m ^-2), size (2.01–2.44 cm) and average bioerosion rates (0.114–4.14 kg m ^-2 year ^-1) were significantly higher in shallow areas (1–3 m) in both reefs. Bioerosion rates were low compared to those reported for parrotfish, endolithic sponges and the black sea urchin D. antillarum, but they were higher than those reported for other small-sized sea urchins in the Caribbean and the Indo-Pacific.     

Phytoplankton composition in relation to primary production in Chesapeake Bay

The dominant phytoplankton taxa during seasonal periods of peak primary productivity were identified during a 4 yr study (July 1989 to June 1993) in Chesapeake Bay. Maximum phytoplankton abundance occurred from late winter to early spring, and was dominated by a few species of centric diatoms. This development was followed by more diversified assemblages of diatoms and phytoflagellates that produced additional concentration peaks in summer and fall; all these maxima were accompanied by concurrent productivity peaks. High summer productivity resulted when the phytoplankton concentrations of diatoms and phytoflagellates were augmented by an increased abundance of autotrophic picoplankton. There was variability in both the seasonal and annual growth maxima of these algal populations and in total productivity. Higher cell concentrations and productivity were associated with higher nutrient levels on the western side of the bay, at sites adjacent to major tributaries. Periods of highest productivity were in spring and summer, ranging from 176 to 346 g Cm^-2yr^-1 over the 4 yr period, with a mean annual productivity of 255 g Cm^-2yr^-1. The bay stations rates ranged from 82 to 538 g Cm^-2yr^-1.     

Effects of Lunatia heros predation on the population dynamics of Mya arenaria and Macoma balthica in Maine,USA

The interaction between the naticid snail predator Lunatia heros and 2 iteroparous, infaunal, intertidal bivalves was investigated in Lubec, Maine, USA. The Mya arenaria population consists primarily of young, small individuals. M. arenaria survivorship is low when young (3.5% yr^-1 for the first 5 yr), then increases. M. arenaria can attain a length of 60 mm, but it is susceptible to L. heros attack only until it is 30 mm long. It delays reproduction until it is 20 mm long (3.8 yr) and diverts its resources instead into rapid early growth (4.9 mm yr^-1 for the first 5 yr). The Macoma balthica population has a larger proportion of older individuals than does that of M. arenaria. Survivorship is higher for M. balthica than for M. arenaria (76.3% yr^-1 for the first 5 yr). Unlike M. arenaria, M. balthica attains a final length of only 25 mm and all sizes are susceptible to L. heros attack. M. balthica grows slowly (2.7 mm yr^-1 for the first 5 yr) and diverts its resources into earlier reproduction at a length of 10 mm (2.9 yr). These contrasting life-history patterns and the possible relationship between bivalve resource allocation and refuges from predation are discussed.     

Seasonal growth and productivity ofMacrocystis integrifolia in British Columbia,Canada

The seasonal growth rates and nitrogen and carbon fluxes were estimated for two subtidalMacrocystis integrifolia Bory kelp forests in British Columbia, Canada from changes in population structure through time. Mean relative growth rates of the forests varied from a high of 4.3% d^-1 to a low of-3.6% d^-1. Mean net assimilatioon rates of carbon (a photosynthesis analog) varied from a high of 0.66 g C m^-2 of foliage d^-1 to a low of-0.87 g C m^-2 d^-1. The leaf area index ranged from 0.3 to 11.9. Annual carbon input on a foliage area basis was calculated at 250 g C m^-2 yr^-1. Annual carbon input to the forest was estimated at 1 300 g C m^-2 of ocean bottom yr^-1. The yearly nitrate nitrogen input to the forest was estimated at 60 g N m^-2 of ocean bottom yr^-1. The net ecosystem production varied from-520 to +31 g C m^-2 of ocean bottom yr^-1. The intra-forest, inter-forest and seasonal variabilities of these productivity parameters are discussed.     

Allocation of45calcium to body components of starved and fed purple sea urchins (Strongylocentrotus purpuratus)

Several lines of evidence in the literature indicate that environmental stress such as starvation may initiate reallocation of sea urchin endoskeletal tissue. For example, Aristotle's lantern enlarges under conditions of starvation, and sea urchins tagged with tetracycline and then fed develop a distinct growth line, while starved individuals develop a diffuse pattern. We designed anin vivo system to examine stress-related changes in calcification in the purple sea urchinStrongylocentrotus purpuratus. SmallS. purpuratus (ca. 2 cm test diam) were collected from the Mission Bay jetty or Imperial Beach (San Diego, California, USA) in 1987.⁴⁵Ca was incorporated from seawater into all body fractions including the organic tissue/coelomic fluid. In an initial experiment, sea urchins were fed or starved for 4 wk and then post-incubated in isotope. Overall, starved individuals deposited new calcite more slowly than did fed individuals; however, allocation was very different and calcification of teeth of starved sea urchins was nearly as great as in fed individuals. In a second experiment,S. purpuratus were first pre-labeled with isotope and then treated by feeding or starving. More of the labeled calcium was mobilized from the soft tissues and coelomic fluid into calcite in fed than in starved individuals. Growth of the teeth in starved sea urchins was significantly greater than in those fed. We conclude that starvation changes the metabolism of calcium in order to preferentially build teeth. However, we also found no evidence that calcium was resorbed from old skeletal calcite in order to build new skeleton.     

Life history of the red mullet Mullus surmuletus from the bottom-trawl fishery off the Island of Majorca (north-west Mediterranean)

The red mullet Mullus surmuletus is one of the main target species of the trawling fishery along the continental shelf off the Island of Majorca. The size distribution of the catches, and the reproduction, age and growth of this species have been studied based on sampling carried out from 1990 to 1992. The length range of the catches was between 10 and 32 cm, with a main distribution between 15 and 20 cm, but this included small specimens (recruits of 10 to 11 cm) from August. In the >19 cm length-class, females clearly dominated. Monthly variations in the gonadosomatic index (GSI) and in the percentage of mature specimens showed that males spawn from December to June, whereas the reproductive activity of the females centres around spring. Fifty percent of males and females mature at 15 and 16.8 cm, respectively, corresponding to 1 yr of age. Otolith age-readings indicate that the population exploited in the trawl fishery consists of six age-groups, including a very high proportion of individuals between 0 and 4 yr old. Population growth curves revealed that females grow comparatively slowly over a longer period of time and attain greater asymptotic sizes than males. The growth parameters for the whole population are: asymptotic length, L =31.28 cm; growth coefficient, K=0.211 yr^-1; theoretical age when length is zero, t ₀=-2.348 yr.     

Abundances,growth rates,and production of tropical neritic copepods off Kingston,Jamaica

Weekly samples were collected near Kingston, Jamaica in 27 m vertical hauls, using 200 and 64µm mesh plankton nets, from July 1985 to January 1987. Thirtytwo copepod species were identified; nauplii and all copepodite stages were enumerated. Total copepod abundance ranged from 2.56 to 87.3 × 10⁴ m^–2. The annual abundance cycle was bimodal with peaks in October–November and May–June corresponding to the rainy seasons. Mean annual copepodite biomass was 0.15 g AFDW m^–2 ranging from 0.03 to 0.41 g AFDW m^–2. Mean generation time (from egg to adult) at 28°C was 19.5 d for the common speciesCentropages velificatus, Paracalanus aculeatus, andTemora turbinata. Isochronal development was demonstrated for copepodites ofP. aculeatus andT. turbinata, but not forC. velificatus. Mean daily specific growth rates (G) were 0.63, 0.63, and 0.48 d^–1 forC. velificatus, P. aculeatus, andT. turbinata, respectively. In general, daily specific growth rates decreased in the later copepodite stages. Thus, it is postulated that growth of later stages and egg production may be food limited. Annual copepodite production was estimated as 419 kJ m^–2 yr^–1, while annual exuvial production and naupliar production were 35 and 50 kJ m^–2 yr^–1, respectively. Egg production was estimated as 44% (184 kJ m^–2 yr^–1) of the total copepodite production. Thus, mean total annual copepod production was 688 kJ m^–2 yr^–1. This estimate is within the range of copepod production estimates in coastal temperate regions.     

Effects of food type and ration on reproductive maturation and growth of the sea urchin Strongylocentrotus droebachiensis

We investigated the effects of food quality and quantity on reproductive maturation and growth of juveniles of Strongylocentrotus droebachiensis (Müller) in a 22 month laboratory experiment in which we fed sea urchins four diets: (1) kelp (Laminaria spp.) for 6 d wk⁻¹ and mussel (Mytilus spp.) flesh for 1 d wk⁻¹ (KM); (2) kelp for 7 d wk⁻¹ (high ration, KH); (3) kelp for 1 d wk⁻¹ (low ration, KL) and (4) no food other than encrusting coralline algae (NF). At their first and second opportunity for reproduction, all sea urchins in the KM and KH treatments, and most in the KL treatment were reproductively mature, whereas all sea urchins in the NF treatment remained immature. Gonad index differed significantly among all fed treatments at first and second reproduction, and was highest in the KM and lowest in the KL treatment. Gonad index was similar in both sexes at first reproduction, but it was higher in females than in males at second reproduction. Diet had little or no effect on the relative abundance of spermatocytes, spermatozoa, or nutritive phagocytes in testes at first and second reproduction. In ovaries, nutritive phagocytes were significantly more abundant in females in the KM and KH treatments than in the KL treatment at first reproduction, and significantly more abundant in unfed (NF) than fed (KM, KH, KL) females at second reproduction. Mean oocyte size was similar in all fed females at first reproduction, but significantly larger in fed than unfed females at second reproduction. Mean ovum size was similar in all fed females in both reproductive periods. Increase in test diameter was greatest in the KM treatment and smallest in the KL treatment; sea urchins in the NF treatment decreased slightly in size. Survival was 95 to 100% in all fed sea urchins but significantly lower in unfed ones. The feeding rate on kelp was significantly greater in the KL than the KM and KH treatments. In the KM treatment, the feeding rate on kelp increased significantly over a 6 d period after mussel flesh was provided. Our results demonstrate that a diet of high food quality and quantity accelerates reproductive maturation and growth rate, and enhances gonad production and survival in juvenile and young adult S. droebachiensis. These findings contribute to our understanding of the reproductive ecology of S. droebachiensis in habitats with differing food supplies (e.g., kelp beds and barrens). Our results also can be used to improve aquacultural practices for sea urchins. Received: 3 June 1998 / Accepted: 2 February 1999     

Oxygen uptake and comparative energetics among eggs and larvae of three subtropical marine fishes

Oxygen uptake was measured and energy budgets were determined for eggs and larvae of bay anchovy (Anchoa mitchilli), sea bream (Archosargus rhomboidalis) and lined sole (Achirus lineatus). Using an energetics model, both minimum rations and prey levels were estimated. The Q_{O 2} [μl (mg dry wt)^-1 h^-1] increased approximately five-fold for all species during development from egg to feedingstage larvae. The weight exponents in power functions, R=aW^b, relating oxygen uptake (R) to dry weight (W) did not differ significantly among species. The exponents were: bay anchovy, 0.9770; sea bream, 0.8382; lined sole, 0.9416. Oxygen uptake was generally lower for bay achovy than for the other species. In the energy budgets issimilation efficiencies of all species were low (24–75%). Gross growth efficiencies ranged from 11–41%, and net growth efficiencies from 38–57%. An estimated 32–83% of ingested energy was excreted in feces and urine. The lavae used a relatively small proportion of ingested energy in metabolism (6–31%). The anchovy had the lowest assimilation and gross growth efficiencies, and the highest excretion rate. Consistent with the better growth and survival observed in previous experiments, sea bream were the most efficient assimilators and had the lowest required minimum rations. Required minimum prey levels for first-feeding larvae ranged from 400 copepod nauplii l^-1 for sea bream and bay anchovy to 1 000 l^-1 for lined sole. The estimated required prey levels were higher than levels at which significant survival had been observed.     

Influences of habitat and natural disturbances on contributions of massive Porites corals to reef communities

We compared densities, distributions and size frequencies of massive corals in the genus Porites on five relatively exposed, mid-shelf reefs (50 km offshore) in the central Great Barrier Reef with those on a sheltered inshore reef (10 km offshore). Data included various transect and mapping studies between 1984 and 1990, estimates of size-dependent damage from the crown-of-thorns starfish Acanthaster planci, estimated densities of herbivorous sea urchins (potential predators of juveniles), and observations of size-specific effects of tropical cyclones. Assemblages of Porites spp. on mid-shelf reefs were dominated by small colonies (2 to 10 cm diam) established either from planula larvae or from small tissue remnants that had survived A. planci predation in the early to mid-1980s. Large colonies (up to 10 m diam) were scarce, except for localized aggregations on terraces at the base of reef slopes (6 to 12 m deep). Extensive space suitable for settlement by coral larvae can be attributed to recurrent cyclones and A. planci outbreaks. Despite low sea urchin predation, the slowly growing Porites juveniles are likely to die from overgrowth by numerous, much faster growing corals. On the sheltered inshore reef, the coral community was dominated by very large (>5 m diam) Porites colonies, several centuries old; recruitment was mainly by fragmentation of large colonies; there was little space available for settlement, and probabilities of juvenile mortality from grazing urchins were high. Differences in settlement and early survival of Porites spp. are exacerbated by different regimes of storm damage. A model is proposed that links wave climate with the size and age reached by corals before dislodgement by storm waves, and which is consistent with observed densities and size-frequency distributions of Porites in sheltered and exposed areas.     

Validation of otolith-increment age estimates for a deepwater fish species,the warty oreo Allocyttus verrucosus,by radiometric analysis

Otolith increment age estimates for a deepwater species, Allocyttus verrucosus, were validated by comparison with the results from ²¹⁰Pb:²²⁶Ra radiometric analysis. Transverse sectioning and subsequent grinding of otoliths to a thickness of 0.2 mm revealed increments which provided age estimates for a range of fish sizes. Age estimates ranged from 7 yr for an immature fish of 15.2 cm total length (TL) to 130 yr for a female fish of 36.5 cm TL. Age at maturity was estimated as 28 yr for females and 24 yr for males. In comparison, radiometric analysis of whole otoliths, using a single linear otolith-mass growth-rate model suggested maximum ages of 130 to 170 yr for fish of 34 to 35 cm TL. Radiometric ages were also recalculated using a two-phase otolith-mass growth-rate model in which the growth rate was assumed to slow after maturity to 90% of the pre-maturity rate. This reduced the maximum age to 132±15 yr for a mean fish length of 34.5 cm. Age at maturity for females was estimated at 34 yr. The similarity between age estimates from otolith-increment counts and radiometric analysis strongly supports the accuracy of results from both methods, and encourages further use of such comparisons as an alternative to traditional validation techniques.     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. IV. Pseudodiaptomus marinus, the egg-carrying calanoid

Population dynamics and production of the egg-carrying calanoid copepod Pseudodiaptomus marinus were studied for a year in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was perennial, with a large numerical peak in June and small peaks in September/October and November/December. During the study period, at least 11 generations could be detected. For each generation, the stage-specific survival from egg to Copepodite Stage (C) V was determined; it was very high during early life stages (egg to NIII), and gradually decreased beyond. On average, 94% of eggs recruited into NIII, which is strongly contrasted with very high (>ca. 90%) mortality during the corresponding stages for free-spawning copepods, i.e. Acartia omorii, Centropages abdominalis and Paracalans sp. This demonstrates that the egg-carrying strategy has a great advantage to reduce mortality in egg stage. The biomass of this species showed marked seasonal variations largely in parallel with numerical abundance. The instantaneous somatic growth rate increased linearly with temperature. The population production rate was estimated as the sum of somatic growth of larval stages and egg production of adult females; the annual integration was 51.0 mg C m⁻³ yr⁻¹ or 0.38 g C m⁻² yr⁻¹. Received: 11 November 1996 / Accepted: 7 December 1996     

Influence of some coral reef communities on the calcium carbonate budget of Tiahura reef (Moorea,French Polynesia)

The calcium carbonate budget of coral reefs is the result of the interaction of the processes of calcification and biological degradation, and is reflected in the chemical properties of the seawater overlying the reefs. A series of experiments at Moorea Island (French Polynesia) in 1988 monitored the diurnal and nocturnal variations in the chemical properties of seawater under field and laboratory conditions. Our results revealed that in the study area (Tiahura barrier reef flat), the calcium carbonate budget varied over space and time as a function of location in the water current. Two in-situ sites were investigated; one was situated 100 m from the algal crest of the barrier reef, the other 300 m further downstream. As a result of cumulative upstream events, the daily net calcification was ten times higher at the downstream (5.22 gm^-2 d^-1) than at the upstream (0.45 gm^-2 d^-1) site. The carbonate uptake by in situ Porites lobata in enclosures (8 kgm^-2 yr^-1) was ten times higher than the uptake by the whole community in the surrounding water (0.8 kgm^-2 yr^-1) and five times higher than that recorded for P. lobata in laboratory experiments (1.4 kgm^-2 yr^-1), where illumination levels were 10% of in situ levels. In laboratory experiments, the planktonic fraction of the seawater had no perceptible influence on the calcium carbonate budget. In the absence of bioeroders, living coral totally depleted the carbonate content of the seawater (3.7 gm^-2d^-1). Bioerosive organisms played an important role in restoring this calcium carbonate; e.g. sea urchins grazing on algal turf covering dead coral ingested CaCO₃ and released this as a carbonate powder (1.26 gm^-2d^-1); a form of carbonate which is extremely accessible to chemical dissolution.     

Growth and production of the inshore marine copepod Pseudodiaptomus marinus in the central part of the Inland Sea of Japan

The growth and production of the inshore marine copepod Pseudodiaptomus marinus was studied in the central part of the Inland Sea of Japan. The stage-specific growth rate was determined under controlled laboratory conditions by examining the length-weight relationship and development rates at various temperatures. The stage duration was short and constant from NII to CII, beyond which development was retarded. Males developed faster than females in CIV and CV. The specific growth rate was highest in copepodite stages followed by the nauplii and adult females (=egg production rate). The daily production of P. marinus was estimated from the stage-specific growth rate and stage-specific abundance in nature as the sum of the individual stages. The production changed seasonally with water temperature and population biomass. Daily production and biomass (P/B) ratios increased linearly with temperature. Total annual production was 20.7 mg C m^-3 yr^-1.     

Settlement and recruitment of sea urchins (Strongylocentrotus spp.) in a sea-urchin barren ground and a kelp bed: are populations regulated by settlement or post-settlement processes?

I sampled recruitment of very small sea urchins (Strongylocentrotus spp.) by using the anesthetic magnesium chloride to remove individuals from substrata collected in sea-urchin barren grounds (barrens) and kelp beds at Naples Reef near Santa Barbara, California, USA. Preliminary sampling found low numbers of newly settled individuals(<0.6 mm test diam) from April–July in 1984 and 1985, and in April, 1986. In early May, 1986, I found many newly settled seaurchins (0.3 to 0.6 mm, 5 to 17 d old), and I compared the densities of the cohort on several types of natural substrata in barrens and kelp-bed habitats. Newly settled individuals of both purple sea urchins (S. purpuratus) and red sea urchins (S. franciscanus) were present in similar, high densities (1 000 S. purpuratus m^-2) on foliose red algal turf, a dominant substratum ofthe kelp bed, and on crustose coralline algae, the dominant substratum of an adjacent barrens. Larvae of S. purpuratus reared and tested in the laboratory showed high rates of settlement on both red algal turf and on crustose coralline algae, but significantly lower rates on rock. Larvae also settled in response to a partiallypurified extract of coralline algae. The reduced settlement on natural rock surfaces relative to either algal treatment and the significant settlement in response to the extract of coralline algae indicate that larvae discriminate between natural substrata and probably respond to a settlement cue other than, or in addition to, a simple microbial (bacterial) film. The similar densities of young recruits of S. purpuratus on dominant substrata of barrens and kelp bed show that, at least in this case, differential settlement cannot explain the high densities of sea urchins in the barrens habitat. Movement between barrens and kelp bed is unlikely given the small sizes of the newly recruited sea urchins relative to the large distances often involved. Reduced post-settlement mortality of newly settled individuals in the barrens remains the most likely mechanism leading to the higher densities of sea urchins in barrens relative to kelp-bed habitats.