On predation of pelagic larvae and early juveniles of marine bottom invertebrates by adult benthic invertebrates and their passing alive through their predators

The dynamic quantitative balance between prey and predator invertebrate species inhabiting the same shallow-shelf (sublittoral level bottom) benthic communities was first discussed by Thorson (1953). Thorson considered the exact timing of larval settlement of prey and predator species possessing pelagic development and temporal supression of the adult predators' feeding activities during reproduction at the time of the preys' settlement to constitute the major factors which facilitate survival of the prey species in such communities. However, information obtained demonstrates that Thorson's “mechanism of balance between predator and prey species of benthic communities” is not always effective in securing survival from predation not only of the prey's spat but even sometimes of the predator's spat also. Because of this, the “mechanism” can not be rated as universally effective in all situations. Analysis of the data so far published demonstrates that, in marine benthic communities, especially in shallow-shelf waters, it is not uncommon for gametes, larvae, or early juveniles of different prey species to pass alive through suspension (filter)-feeding and deposit-feeding adult invertebrates preying on them. Sometimes development can even continue after excretion by predators. The hypothesis of Voskresensky (1948) and Goycher (1949) of the importance of this phenomenon for the maintenance and recruitment of the mussel Mytilus edulis and other filter-feeding lamellibranchs of nearshore waters preying on their own and other lamellibranch pelagic larvae must be rejected on the basis of accumulated data on their feeding and general biology and on the adverse influence of the mucous of their faecal pellets and pseudofaeces on the larvae excreted by them alive. The data considered here demonstrate that, although the passing alive of larvae and spat of benthic invertebrates through benthic predators is not uncommon in shallow-shelf bottom-communities, it plays no important role in the processes of maintenance and recruitment of the species and communities involved nor of the marine benthos as a whole. The actual ecological significance of predation on pelagic larvae and bottom spat of benthic invertebrate prey species by all three main trophic groups of marine benthos (suspension or filter-feeders, deposit-feeders, carnivores) and its importance to predator-prey dynamics in marine benthic communities remains open to debate until more reliable quantitative data become available.     

Yolk resorption,onset of feeding and survival potential of larvae of three tropical marine fish species reared in the hatchery

This paper provides basic early life-history information on milkfish (Chanos chanos), seabass (Lates calcarifer) and rabbitfish (Siganus guttatus) which may explain in part the observed differences in their survival performance in the hatchery. Egg size, larval size, amount of yolk and oil reserves and mouth size are all greater in milkfish than in seabass, and greater in the latter than in rabbitfish. During the first 24 h after hatching, rabbitfish larvae grow much faster than milkfish and seabass larvae at similar ambient temperatures (range 26°–30°C, mean about 28°C). The eyes become fully pigmented and the mouths open earlier in seabass and rabbitfish (32–36 h from hatching) than in milkfish (54 h). Seabass larvae learn to feed the earliest. Yolk is completely resorbed at 120 h from hatching in milkfish, and yolk plus oil at 120 h in seabass and 72 h in rabbitfish at 26° to 30°C. Milkfish and seabass larvae have more time than rabbitfish to initiate external feeding before the endogenous reserves are completely resorbed. Delayed feeding experiments showed that 50% of unfed milkfish larvae die at 78 h and all die at 150 h from hatching. Milkfish larvae fed within 54 to 78 h after hatching had improved survival times: 50% mortality occurred at 96 to 120 h, and 10 to 13% survived beyond 150 h. Unfed seabass larvae all died at 144 h, while 6 to 13% of those fed within 32 to 56 h after hatching survived beyond 144 h and well into the subsequent weeks. Unfed rabbitfish larvae all died at 88 h, while 7 to 12% of those fed within 32 to 56 h after hatching survived beyond 88 h. A delay in initial feeding of more than 24 h after eye pigmentation and opening of the mouth may be fatal for all three species.Contribution No. 167 from the SEAFDEC Aquaculture Department     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Laboratory study of predation by Aurelia aurita on larvae of cod,flounder, plaice and herring: development and vulnerability to capture

Capture success of the medusa Aurelia aurita preying on various developmental stages of fish larvae was measured together with larval reactivity and escape speed after being stung. These experiments were conducted in the spring of 1983 with A. aurita medusae collected from Loch Etive, Scotland and laboratory-reared larvae of Gadus morhua L., Platichthys flesus L., Pleuronectes platessa L. and Clupea harengus L. Capture success of the medusae increased with medusa size, but decreased with advancing larval development. Smaller species of larvae were more vulnerable to capture. Larval reactivity to encounters with medusae increased with advancing development, and larger species of larvae were more reactive to encounters. Larval escape swimming speeds also increased with advancing larval development and size. These results indicate that earlier stages of larvae within a species and smaller species of larvae at a given stage are more vulnerable to predation by medusae since they are less reactive to encounters. Apparently they are more susceptible to the effects of neurotoxins. Predation rates on different developmental stages of herring larvae are documented and compared with rates predicted by a predation model. Predictions fell within the range of observed predation rates, but tended to overestimate rates by larger medusae feeding on larger herring larvae. This indicates the possibility of predator satiation and/or behavioural avoidance.     

A laboratory study of predation by Aurelia aurita on larval herring (Clupea harengus): Experimental observations compared with model predictions

Predation by the medusa Aurelia aurita L. on early first-feeding stage larvae of the herring clupea harengus L. was studied in the laboratory. The medusae were captured in Loch Etive, Scotland. Herring larvae were reared from the extificially fertilized eggs of spawning Clyde herring caught in March, 1982. Swimming speeds, volume searched”, capture efficiency and predation rates increased as medusa size increased. Predation rates on fish larvae increased with prey density, but appeared to approach a maximum at high prey densities; in 1 h experiments, a maximum rate of predation of 6.64 larvae h^-1 was estimated by fitting an Ivlev function. A model to predict predation rates was constructed from swimming speeds, sizes and densities of medusae and larvae, and capture efficiency. The rates of predation predicted from the model fell within the range of experimental data, but tended to underestimate rates and did not account for saturation of medusae. Swimming patterns of medusae changed after prey capture: (a) before capture, encounter rates were low and medusae were relatively less active; (b) after capture of 1 larva, encounter rates doubled, with the stimulated medusae exhibiting increased activity and an aftered “searching” path; and (c) after capture of many larvae, swimming speeds and encounter rates of medusae decreased.     

Osmoregulation, nutritional effects and buoyancy of marine larval fish: a bioassay for assessing density changes during the earliest life-history stages

It has been hypothesized that marine fish larvae in the advanced stages of starvation would show increased density (ρ = mass volume⁻¹) from water loss due to osmoregulation failure. Changes in larval buoyancy are currently attributed to swim bladder regulation and protein synthesis or catabolism. Osmoregulation-related changes in density is an alternative mechanism, the importance of which remains untested in the laboratory and the influence of which on vertical distributions is unknown. We provide evidence that loss of osmotic control is a plausible mechanism for increased density of larval cod (Gadus morhua L.). Furthermore, our results show that this mechanism is not restricted to larvae in the advanced stages of starvation. “Relative” larval densities are estimated using a modified density gradient. We use a gravimetric method to separate the effects of nutrition from osmoregulation failure. We assessed the importance of sampling strata on estimates of larval density. Proportional sampling within three depth strata (stratified sample) produced the least biased method for determining the “average” density of a population of larvae in laboratory culture. Larvae sampled from the bottom third of the culture tank were significantly more dense then those sampled from the surface. This was true for larvae of all ages. The average change in density from hatching till death from starvation for larvae sampled in the surface stratum was nominal (Δρ = 5.0 × 10⁻⁴ g cm⁻³), while the change for those sampled from the bottom stratum was large (Δρ = 3.8 × 10⁻³ g cm⁻³). These large density differences suggest that larvae sampled from the bottom stratum were either osmotically stressed or were facultatively changing their density via regulatory pathways. Preliminary observations suggest that vitality is lower amongst those larvae which are sampled near the bottom. The small change in average density of larvae sampled from the surface stratum was due to starvation. The density differences we observed between “osmotically stressed” and “starving” larvae could readily have been misconstrued as differences in feeding and growth experienced by individual larvae. The potential bias of increased density from osmoregulation failure must be considered as a factor in experimental designs developed to assess the effect of fed and starved treatments on buoyancy for larvae of all ages. The simple bioassay we describe may prove useful both as a means of assessing larval condition and as a mechanism for evaluating factors affecting larval vertical distributions in the field. Received: 13 January 1997 / Accepted: 3 February 1997     

Zooplankton growth rates: the influence of female size and resources on egg production of tropical marine copepods

Egg production was measured in 17 species of copepods from the genera Acartia, Calanopia, Centropages, Clausocalanus, Corycaeus, Eucheata, Euterpina, Oithona, Oncaea, Paracalanus, Parvocalanus, Temora and Undinula in Jamaican waters. At the high local temperatures (∼28 °C), mean egg production ranged from 3.2 to 88 eggs female^–1 d^–1, and instantaneous female growth (g, as egg production) ranged from 0.04 to 0.87 d^–1. Female growth was positively related to ambient chlorophyll concentration (r ² = 0.44) and negatively to female body size (r ² = 0.29). Together these two variables explained 60% of the variation in growth. When quadratic terms for chlorophyll and a term for interaction of body size and chlorophyll were introduced, 82% of the variance in growth rate was explained. Egg production rates represent an extension of the resource and size-dependent relationship established for copepodites. In smaller species (<3.5 μg), egg production was comparable to prior copepodite somatic growth; in larger species (>3.5 μg), egg production is compromised at lower resource concentrations than copepodite somatic growth. Thus, it appears that egg production in tropical copepods may be frequently limited by resources in a size-dependent manner. Under conditions where growth is resource limited, we caution against the application of egg production rates for the calculation of total copepod production. Received: 30 May 1997 / Accepted: 13 May 1998     

Salinity effects on the coexistence of cryptic species: a case study on marine nematodes

The coexistence of four cryptic species of Rhabditis (Pellioditis) marina (Nematoda: Rhabditidae) at small geographical scale challenges ecological competition theory and was therefore studied in the laboratory at two different salinities, where their performance in combined cultures was compared with that in monospecies cultures. We found that three of the four cryptic species were able to coexist, but that interspecific interactions (competition and facilitation) were common. Salinity had an effect on these interactions, with a shift from contest to scramble competition. This shift may result from an increased population development of two of the four species at the lower salinity in the monospecific cultures. This experiment demonstrates that abiotic conditions may play an important role in achieving coexistence between cryptic species and can alter the interspecific interactions between them.     

Venoms from six species of marine fish: lethal and hemolytic activities and their neutralization by commercial stonefish antivenom

A specimen of the stonefish Synanceja verrucosa was captured in Okinawa in March 1988. Live specimens of the scorpionfish Inimicus japonicus were purchased from the Tokyo Central Wholesale Market in November 1988 and those of four species of zebrafish Pterois lumulata, P. volitans, P. antennata and Dendrochirus zebra from an aquarium in December 1988. Crude venoms were extracted from dorsal spines of the six species. All venoms exhibited lethal activity against mice and hemolytic activity specific for rabbit erythrocytes. The lethal activity (or hemolytic activity) of each venom was very unstable to freezing, lyophilization and heating. Both lethal and hemolytic activities of S. verrucosa venom were remarkably neutralized by the commerical stonefish (Synanceja trachynis) antivenom. This antivenom was also effective, to some extent, in counteracting the other venom activities. The neutralizing capacities of the antivenom were calculated to be 7310 LD₅₀ ml^-1 for S. verrucosa venom and 1 220 to 2 990 LD₅₀ ml^-1 for the other venoms. Results of neutralization tests suggest that venoms from the six species were comparable in terms of antigenecity.     

The effects of three sympatric ladybird species on oviposition by Menochilus sexmaculatus (Coleoptera: Coccinellidae)

Oviposition site selection has a strong impact on the spatial distribution and sustenance of a species. This site selection by ladybirds (Coleoptera: Coccinellidae) is known to be well mediated amongst other factors by the oviposition deterring pheromones present in conspecific larval tracks. It would be also useful to the organism to be sensitive to heterospecific larval tracks, to protect from intraguild predation; however, this has not yet garnered much attention and there are only a few scattered studies with ambiguous results. The present study thus investigates the oviposition deterring effects of heterospecific presence and semiochemicals of heterospecific life stages (eggs, fourth instar larvae, pupae, adult males and females) of ladybird species, viz. Coccinella septempunctata, C. transversalis, Propylea dissecta on females of Menochilus sexmaculatus. Results reveal that the presence of all heterospecific life stages inhibits oviposition of M. sexmaculatus and the inhibition is density dependent. Semiochemicals of eggs, larval and adult tracks also show density-dependent inhibitory effects. The semiochemical-induced oviposition inhibition was stage specific with respect to the degree of inhibition. Of all heterospecific species tested, maximum and minimum oviposition deterrence of M. sexmaculatus was against C. septempunctata and P. dissecta, respectively. Comparison of our previously published results on oviposition inhibition in M. sexmaculatus due to conspecific presence with the current ones revealed increased inhibition in conspecific presence than heterospecific indicating ability to differentiate.     

Effects of fish feeding by snorkellers on the density and size distribution of fishes in a Mediterranean marine protected area

Although there is a great deal of evidence to show that supplementary feeding by humans in terrestrial environments causes pronounced changes in the distribution and behaviour of wild animals, at present very little is known about the potential for such effects on marine fish. This study evaluated the consequences of feeding by snorkellers on fish assemblages in the no-take area of the Ustica Island marine protected area (MPA; western Mediterranean) by (1) determining if reef fish assemblage structure is affected in space and time by tourists feeding the fish; (2) assessing the effects of feeding on the abundance of the most common fish species; and (3) assessing the effects of feeding on the size structure of the two most numerically dominant ones. In particular, we hypothesised that both the abundance and the size structure of some fish species would increase at the study site following supplementary feeding, since the additional food provided by humans would make the site more appealing to them. Fish feeding influenced the fish assemblages within the Ustica MPA, and significant spatio-temporal changes occurred. While fish feeding appeared to have no effect on the ornate wrasse Thalassoma pavo, there was a noticeable increase in the number of Oblada melanura and Epinephelus marginatus in the impacted location after feeding. It is very likely that aggregations of fishes that evolve as a result of fish feeding by the public may have negative effects on local populations of fishes and invertebrates that make up their prey. Recreational use of coastal areas and MPAs is increasing elsewhere, making fish feeding a generalised human activity. Accurate information about its effect on the fish assemblage is essential to make responsible management decisions.Communicated by R. Cattaneo-Vietti, Genova     

The effect of food availability,age or size on the RNA/DNA ratio of individually measured herring larvae: laboratory calibration

RNA/DNA ratios in individual herring (Clupea harengus) larvae (collected from Kiel Bay, Baltic Sea, in 1989) were measured and proved suitable for determining nutritional status. Significant differences between fed and starving larvae appeared after 3 to 4 d of food deprivation in larvae older than 10 d after hatching. The RNA/DNA ratio showed an increase with age or length of the larvae and was less pronounced in starving larvae compared to fed larvae. The individual variability of RNA/DNA ratios in relation to larval length of fed larvae and of larvae deprived of food for intervals of 6 to 9 d is presented. Based on the length dependency and the individual variability found within the RNA/DNA ratios, a laboratory calibration is given to determine whether a larva caught in the field has been starving or not. An example for a field application is shown.     

Detection of pollution effects on marine macrobenthos: further evaluation of the species abundance/biomass method

Further empirical data are presented to evaluate the method of detecting pollution-induced disturbance in marine benthic communities by a comparison of the distribution of numbers of individuals among species with the distribution of biomass among species. A suggested abbreviated name for the technique is the ABC method (abundance biomass comparison). Application of the technique to new data shows that it is a sensitive indicator of natural physical and biological disturbance as well as pollution-induced disturbance over both spatial and temporal scales. Changes in the configuration of ABC plots during ecological succession are the reverse of those resulting from increased pollution levels. The technique should also be applicable to intertidal sediments, where physical disturbance of the sediment by waves does not appear to preclude its utility.     

A response-surface approach to the combined effects of temperature and salinity on the larval development of Adula californiensis (Pelecypoda: Mytilidae). I. Survival and growth of three and fifteen-day old larvae

Laboratory experiments of a factorial design were used to examine the combined effects of temperature and salinity on the survival and growth of early and late-stage larvae of Adula californiensis (Phillippi, 1847). Response-surface curves were generated to predict optimal conditions for survival and growth in order to better understand the successful recruitment of this species within the Yaquina Bay estuary (Oregon, USA). Three-day old cultured larvae were more sensitive to reduced salinity than were 15-day old larvae. However, the 15-day old larvae showed a narrower temperature tolerance than the 3-day old larvae. A. californiensis larvae survived over a wider range of temperatures near optimum salinities than at salinities near their lower tolerance limit, and conversely. Temperature and salinity ranges for maximum survival (10° to 15°C, 31 to 33) were narrower than the ranges which occur within the estuary where the adult populations exist. Larval size did not increase markedly during the 15-day rearing period, and was not greatly affected by temperature or salinity. No statistically significant temperature-salinity interaction was found for either survival or growth.     

Seaweed secondary metabolites as antifoulants: effects of Dictyota spp. diterpenes on survivorship, settlement, and development of marine invertebrate larvae

Summary. A recent investigation showed that the brown seaweed Dictyota menstrualis was unfouled relative to co-occurring seaweeds, and that larvae of fouling invertebrates avoided settling on D. menstrualis due to chemicals on its surface. The secondary metabolites dictyol E and pachydictyol A are among the compounds found on this alga's surface. In the present study, we tested the effects of specific diterpenes from Dictyota on the survivorship, growth, and development of invertebrate larvae and developing juveniles that could foul seaweeds. Exposure to dictyol E, dictyol B acetate, pachydictyol A, and dictyodial from Dictyota menstrualis and D. ciliolata caused significant larval mortality, abnormal development, and reduce growth rates for three species of co-occurring invertebrates when their larvae were forced into contact with these metabolites. Larvae were damaged at metabolite concentrations as low as 5% of maximum possible surface concentrations of these compounds for the populations of Dictyota we studied. The negative effects of these secondary metabolites on potential foulers, in conjunction with data demonstrating larval avoidance of dictyol-covered surfaces, suggest that these compounds could function as chemical defenses against fouling, and could select for larvae that avoid hosts producing these metabolites. Received 25 May 1998; accepted 22 June 1998.     

Effect of low dissolved oxygen concentrations on behavior and predation rates on red sea bream Pagrus major larvae by the jellyfish Aurelia aurita and by juvenile Spanish mackerel Scomberomorus niphonius

A shift in outcomes of predator-prey interactions in plankton community may occur at sublethal dissolved oxygen concentrations that commonly occur in coastal waters. Laboratory experiments were conducted to investigate how a decline in dissolved oxygen concentration alters the predation rate on fish larvae by two estuarine predators. Behavior and consumption of larval fish by moon jellyfish Aurelia aurita (103.1±12.4 mm in bell diameter) and by a juvenile piscivore, Spanish mackerel Scomberomorus niphonius (30.1±2.1 mm in standard length: SL), were observed under four oxygen concentration treatments (1, 2 and 4 mg l^–1 and air-saturated: 5.8 mg l^–1). Larvae of a coastal marine fish species, red sea bream Pagrus major (7.21±0.52 mm SL), were used as prey for the experiment. Bell contraction rate of the jellyfish did not vary among the oxygen concentrations tested, indicating a tolerance to low oxygen concentration. Gill ventilation rate of the Spanish mackerel increased and swimming speed decreased as the oxygen concentration decreased, indicating that oxygen concentrations 4 mg l^–1 are physiologically stressful for this species. The number of larvae consumed in 15 min. by jellyfish increased whereas those consumed by Spanish mackerel decreased with the decrease in oxygen concentration. Low oxygen concentrations that are commonly observed in coastal waters of Japan during summer have the potential to increase the relative importance of jellyfish as predator of fish larvae and to change the importance of alternative trophic pathways in estuarine ecosystems.Communicated by T. Ikeda, Hakodate     

Products of photosynthesis by marine phytoplankton: the effect of environmental factors on the relative rates of protein synthesis

A method is presented by which the gross pattern of photosynthetic carbon-dioxide fixation in marine phytoplankton can be determined. It depends on differential solvent extraction yielding an ethanol-soluble, a hot TCA-soluble (polysaccharide) and a residue (protein) fraction. Using this fractionation technique, the effects of various environmental factors on the pattern of photosynthesis by the marine diatom Phaeodactylum tricornutum (Bohlin) have been investigated. Low light intensities and increasing degrees of nitrogen limitation in a chemostat increase markedly the relative rates of protein synthesis. Growth of the alga at lower temperatures also increases the proportion of carbon incorporated into the protein fraction. This increased protein syntheses is generally at the expense of the polysaccharide fraction. Preliminary experiments have established the suitability of this fractionation method for natural populations of phytoplankton and have shown similar effects of light intensity on the relative rates of protein synthesis.     

The effect of laboratory conditions on the extrapolation of experimental measurements to the ecology of marine zooplankton. IV. Changes in respiration and excretion rates of boreal zooplankton species maintained under fed and starved conditions

Herbivorous zooplankton species (Calanus plumchrus, Paracalanus parvus and Euphausia pacifica) and carnivorous species (Parathemisto pacifica and Pleurobrachia pileus) collected from Saanich Inlet, British Columbia, Canada, were maintained in the laboratory under fed and starved conditions. Respiration rate and excretion rates of ammonia and inorganic phosphate were measured successively on the same batch populations of each species in different feeding conditions. Respiration rate remained at a constant level or increased during the feeding experiment but decreased progressively in starved individuals. Herbivorous, but not carnivorous, species showed a rapid decrease in both excretion rates for the first few days of an experiment irrespective of feeding conditions. However, the general level of excretion rates of fed specimens was higher than that of starved ones. The O:N, N:P and O:P ratios were calculated from respiration, ammonia excretion and phosphate excretion and discussed in relation to metabolic substrates of animals during the experiment. A marked difference was shown in the O:N ratio between fed hervivores (>16) and fed carnivores (7 to 19), suggesting highly protein-oriented metabolism in the latter. One unknown factor causing variation in excretion rates is speculated to be the physiological stress on animals during sampling from the field. It is suggested that the laboratory measurement of realistic excretion rates of zooplankton is difficult owing to their large fluctuations, but this is not the case with respiration rate.     

Influence of site,season and year on contributions made by marine,estuarine, diadromous and freshwater species to the fish fauna of a temperate Australian estuary

Catches obtained at regular intervals by beach seining, gill netting and otter trawling at ten, four and six sites, respectively, have been used to determine the contribution of the different species and life-cycle categories of fish to the ichthyofauna of the large Swan Estuary in temperate south-western Australia between February 1977 and December 1981. These data were also examined to investigate the influence of site, season and year on the densities of the more abundant species. A total of 630 803 fish, representing 36 families and 71 species, were caught in the shallows using beach seines during this 5 yr study. Although the majority of these species were marine teleosts that were caught infrequently (marine stragglers), representatives of 7 of the 15 most abundant species were marine teleosts which entered the estuary regularly, and in large numbers (marine estuarine-opportunists). Of the remaining 8 most abundant species in the shallows, 7 completed their life cycle within the estuary (estuarine species) and 1 (Nematalosa vlaminghi) was anadromous, feeding for a period at sea and spawning in the upper reaches of the estuary. The contribution of individuals of the marine estuarine-opportunist category to catches in the shallows declined from nearly 95% in the lower estuary, to 17% in the middle estuary and 6% in the upper estuary. The estuarine and anadromous groups made a considerable contribution to the catches in both the middle and upper estuaries. By contrast, the contribution of freshwater species was small and even in the upper estuary accounted for only 0.2% of the catch. Site within the estuary generally influenced the catches of individual species to a greater extent than either season or year, or the interactions between these factors. When seasonal effects were strong, they could be related to summer spawning migrations into the upper estuary (Nematalosa vlaminghi, Amniataba caudavittatus), spring immigrations into the lower estuary (Mugil cephalus), or winter movements into deeper and more saline waters (Apogon rueppellii). Annual variations in the density of Torquigener pleurogramma were related to marked annual differences in the recruitment of the 0+age class.