Investigations into the diet and feeding ecology of the bearded goby Sufflogobius bibarbatus off Namibia

The bearded goby Sufflogobius bibarbatus has come to replace sardine Sardinops sagax in the diets of many top predators within the marine environment off Namibia, and it is playing a key role within the region's foodweb. Previously published information on the diet of the bearded goby has been derived from small samples and has been contradictory. We investigated the gut contents of over 3 500 specimens of S. bibarbatus from two separate studies. The results indicate that the species has a fairly generalised diet, feeding mostly on benthic and pelagic animals (phytoplankton was rare), in a way that reflects ontogeny, habitat and likely prey availability. Smaller, more pelagic individuals included a substantial number of pelagic organisms in their diet and the proportion of benthic organisms increased with the size of the goby. Given the importance of this species in recycling benthic carbon through the foodweb of the northern Benguela ecosystem, there is a need to describe and quantify the benthic food environment.     

African penguins as predators and prey – coping (or not) with change

African penguins Spheniscus demersus live in the Benguela and western Agulhas ecosystems off southern Africa. Their numbers decreased throughout the 20^th century from at least 1.5 million to about 0.18 million adults, although different regional trends were apparent. They feed to a large extent on shoaling epipelagic fish, notably anchovy Engraulis capensis and sardine Sardinops sagax, and regional trends in the abundance of penguins are associated with trends in the abundance and distribution of these prey fish. Many first-time breeders emigrate from colonies where feeding or other conditions at the time are unfavourable to more favourable breeding localities. This has led to both the extinction and formation of colonies. Food now may limit colonies at relatively small sizes, a fact attributable to industrial fisheries reducing the densities of forage fish. African penguins share their habitat with several other predators, with which they compete for food and breeding space. One of these, the Cape fur seal Arctocephalus p. pusillus, increased through the 20^th century to 1.5 – 2 million animals at its close. Reported observations of predation by fur seals on seabirds have increased in recent decades and threaten the continued existence of small colonies of penguins. Stochastic modelling suggests that colonies of 10 000 pairs have a 9% probability of extinction in 100 years, so smaller populations should be regarded as "Vulnerable". However, in a period of prolonged food scarcity off southern Namibia, the regional population decreased from more than 40 000 pairs in 1956 to about 1 000 pairs in 2000, and many colonies numbering less than 1 000 pairs became extinct. The minimum viable population for African penguins is currently considered to be >40 000 pairs, likely of the order of 50 000 pairs, a figure equivalent to its level in 2000. The chance of survival of the species through the 21^st century is tenuous.     

Spatial variability in branchial basket meristics and morphology of southern African sardine Sardinops sagax

We examined spatial variability in meristic and morphological characteristics of the branchial basket of sardine Sardinops sagax collected from four geographical regions around the southern African coast, namely Namibia and the South African west, south and east coasts. Our analysis tested the hypothesis of three putative sardine stocks off South Africa, one in each of the three geographical regions. We therefore collected fish data from Namibia to compare with South Africa, because sardine from the two countries are considered to be separate stocks. Morphometric measurements (gill arch length and gill raker spacing) and meristic data (number of gill rakers) were collected from the left side of the first gill arch from a total of 377 sardine, approximately equally divided between the regions. A multivariate general linear model with caudal length as covariate was used to assess differences among fish from the four regions and significant differences were observed, although not always consistently across all fish size classes. Small South Coast sardine had shorter gill arches than small West Coast sardine, but adults had gill arches of similar length, longer than those from Namibia and the East Coast. Small sardine from the South Coast had fewer gill rakers than small sardine from the West Coast, but larger fish had similar numbers of gill rakers, significantly more than sardine from Namibia and the East Coast. Sardine from the West and South coasts had similar gill raker spacings, which were smaller than those of fish from the East Coast and Namibia. Despite spatially and particularly temporally unbalanced sampling, we consider that these differences provide evidence of spatial variation in Benguela sardine phenotype and that it would support the hypothesis of discrete sardine stocks off Namibia and South Africa. The results are consistent with the hypothesis of three sardine stocks within the southern Benguela.     

Groupers of the World: A Field and Market Guide

The diet of Chrysaora hysoscella was investigated from 55 specimens caught near the surface throughout 24h in Walvis Bay Lagoon, Namibia, during September 2003. The diet was diverse, ranging from dinoflagellates to carideans, and included abundant benthic species (adults and larvae). There was no clear overall relationship between medusa size and either the number or type of prey ingested, although aspects of both measures were significantly related to size when analyses were confined to the nocturnally collected data. Whereas there was a significant difference in the diet of medusae collected by day and night, principally because of an increase in the abundance of benthic prey items, it was not possible to partition this between the vertical migration of predator (downwards) or prey (upwards) owing to a lack of information on the ambient prey environment. Regardless, the abundance of benthic prey in the diet could help explain the predominance of medusae off Namibia, where there has been an increased flow of surface production to the benthos since the collapse of the pelagic fisheries.     

Spatial variation in meristic and morphometric characteristics of sardine Sardinops sagax around the coast of southern Africa

Spatial variability in phenotypic characteristics within a fish population may be used to infer the existence of multiple stocks, and knowledge of the population structure of exploited species is important for their sustainable management. In this study we investigated geographic variability in meristic (vertebral count) and morphometric (body shape) characteristics of sardine Sardinops sagax from three southern African regions: Namibia, the South African west coast, and the South African south coast. There were significant regional differences in vertebral counts, with Namibian fish tending to have more vertebrae than South African fish, but with no difference between sardine off the South African west and south coasts. Body shape was characterised using morphometric landmarks, and shape changes were investigated using geometric morphometrics, with influences on fish shape assessed using multivariate regressions. To remove the effects of allometry, the 22 shape variables (represented by Procrustes coordinates) were regressed against fish size (represented by centroid size). There was a significant allometric effect, which was removed by using the residuals of these regressions in further analyses. A multivariate multiple regression was applied to the 22 size-corrected shape variables and to three covariates: condition factor, stomach fullness and fat stage. The significant influence of these covariates was removed by using the residuals from this regression in a canonical variate analysis, where corrected shape variables were classified in relation to region, year and sex. The Mahalanobis distances from the canonical variate analysis differed significantly for most combinations of classifier variables, with clear separations among the three regions. Sardinops sagax off Namibia had thicker bodies and more contracted tail regions than South African fish, whereas S. sagax off the South African south coast had thinner bodies and smaller heads than those off the west coast. These results support the hypothesis of multiple sardine stocks around southern Africa.     

Distribution,population size and conservation of the jackass penguin Spheniscus demersus

The jackass penguin, endemic to southern Africa, is the only member of the Sphenisciformes now breeding in Africa. Its breeding distribution extends from Sylvia Hill, South West Africa (Namibia), to Bird Island, Algoa Bay, South Africa. The non-breeding range extends from Sette Cama, Gabon, to Inhaca Island, Moçambique. Jackass penguins do not usually occur far off shore. The minimum total breeding population in 1978/79 was estimated to be approximately 134 000 birds. Numbers halved between 1956 and 1978 alone as a result of collapses in the South and South West African pilchard populations. Numerical decreases at Possession and Dassen Islands were particularly severe. Prior to 1956 numbers had also decreased, primarily because of large collections of eggs. Sealing activities and guano harvests may also have influenced trends. Only colonies at the periphery of the breeding distribution have increased numerically in recent times, and such increases appear to be related to the availability of suitable prey. Although the species breeds in captivity and is unlikely to become extinct, its continued survival in the wild probably relies on the continued good state of the pelagic fish resources.     

Diet of the South African fur seal (1974–1985) and an assessment of competition with fisheries in southern Africa

Almost 1 000 South African fur seals Arctocephalus pusillus pusillus were collected at sea (inshore and offshore) for examination of stomach contents and other biological parameters. The main prey of seals above the age of one year was teleost fish, of which at least 28 species were identified. Overall, 74,6 per cent of the diet was teleost fish, 16,7 per cent cephalopods, 4,8 per cent crustaceans such as shrimps, prawns, amphipods and isopods and 3,2 percent rock lobster. Within this overall dietary pattern there were marked regional differences. The importance of individual fish species in the diet was investigated from the numbers and the sizes of otoliths occurring in the stomachs, from which the identity and size classes of the prey species could be calculated. Again there were regional differences in the importance of different prey species. On the west coast of South Africa the most important species were anchovy and Cape hakes (each 23% of total diet), whereas on the south coast the important species were anchovy (17%), horse mackerel (14%), pilchard (12%) and Cape hake (14%). Off South West Africa the two dominant species were pelagic goby (52%) and horse mackerel (23%), anchovy and Cape hake constituting only 4 and 5 per cent respectively of total diet. The total annual consumption of anchovy and Cape hake by seals in South African waters was calculated at 125 000 and 120 000 tons respectively.     

Trophic relationships in the demersal fish community off Namibia

The benthic fish community off Namibia (between the Cunene River and Walvis Bay) in 50–650 m of water was studied during three bottom trawl surveys. The community was investigated on the basis of distribution, abundance and diet of 51 species, constituting 95 per cent of the demersal fish biomass. Dietary studies revealed the existence of three major trophic groups, one containing species that prey on pelagic and nektonic organisms, a second dominated by predators that feed on benthic polychaetes and copepods and a third group containing predators of benthic crabs, demersal shrimps and fish. Well differentiated from these groups are a few species that prey mainly on jellyfish and ophiuroids. Because for most of the species, their trophic level changes with growth, they were divided into size classes which are analysed independently. To describe interactions between the size classes, a similarity index combining diet affinity with spatial coincidence was applied. Cluster analysis showed that, for Merluccius capensis, Raja straeleni, R. clavata and Trachurus trachurus, there is a large difference in both geographic distribution and dietary preference between the size classes. For species such as Hoplostethus atlanticus, Austroglossus microlepis and Coelorinchus flabellispinis, the different size classes seem to share the same ecological niche. Variation in the number of species and the parameters mean abundance and niche width are described to enhance knowledge of the general structure of the community.     

Diet of the Cape fur seal Arctocephalus pusillus pusillus at the Robberg Peninsula,Plettenberg Bay,and implications for local fisheries

Cape fur seals Arctocephalus pusillus pusillus were harvested to extirpation on the Robberg Peninsula, Plettenberg Bay, on the south-east coast of South Africa, between the 17th and early 20th centuries. Seals returned to Robberg in small numbers during the early 1990s and their numbers subsequently increased. We studied the diet of this increasing population using faecal (scat) sampling to determine: the species composition and size of prey in the diet of Cape fur seals at Robberg; to explore temporal variation in the diet; and to investigate the potential for competition between seals and the fisheries around Plettenberg Bay. Of the 445 scats collected, 90% contained hard prey remains and 15 teleost prey species were represented in the 3 127 otoliths that could be identified. The seals’ most important prey species in terms of numerical abundance, frequency of occurrence and mass in the diet, were anchovy Engraulis encrasicolus, sardine Sardinops sagax, horse mackerel Trachurus capensis, sand tongue-fish Cynoglossus capensis and shallow-water hake Merluccius capensis (in decreasing order of importance for numerical abundance). The proportion of anchovy in the diet increased during the study period (2003–2008), whereas the proportion of sardine decreased. The estimated average annual consumption of sardine by seals was higher than the average annual catch made by purse-seine fisheries in this area, suggesting resource competition between seals and purse-seiners, especially in the light of continuing growth of seal numbers in the area. However, direct competition between seals and linefisheries appeared to be minimal. Scat sampling of Cape fur seals holds potential to serve a useful and cost effective indicator of temporal changes in sardine abundance.     

The food and feeding of seven fish species from the Campbell Plateau,New Zealand

The food and feeding habits of hoki (Macruronus novaezelandiae), southern blue whiting (Micromesistius australis), javelin fish (Lepi‐dorhynchus denticulatus), ling (Genypterus blacodes), smooth rattail (Coelorinchus aspercephalus), silverside (Argentina elongata), and small‐scaled notothenid (Notothenia microlepidota) sampled from the Campbell Plateau in 1979 were examined. The importance of prey items in the diet has been assessed by an ‘index of relative importance’, which combines measurements of frequency of occurrence, number, and weight of prey. Hoki, southern blue whiting, and javelin fish are pelagic feeders. Hoki preyed largely on natant decapod crustaceans, amphipods, and myctophid and photichthyid fishes. The main prey of southern blue whiting were amphipods, natant decapods, and euphausiids. Javelin fish fed on natant decapods, amphipods, and small squid. Seasonal and regional differences in feeding, and dietary changes with length of fish were evident. Ling, smooth rattail, silverside, and small‐scaled notothenid are predominantly benthic feeders. Ling preyed on natant decapods, macrourid fishes, and small hoki. The diet of rattail comprised natant decapod crustaceans, opal fishes (Hemerocoetes spp.), and poly‐chaetes. Silverside fed almost solely on salps. Salps, amphipods, brachyuran crustaceans, and opal fishes were the main prey of small‐scaled notothenid.     

A hard-knock life: the foraging ecology of Cape cormorants amidst shifting prey resources and industrial fishing pressure

Once one of the most numerous seabirds of the Benguela upwelling system, the population of Cape cormorants Phalacrocorax capensis has decreased by 60% in the past three decades and the species is listed as Near Threatened. Declines in prey availability and/or abundance brought about by recent changes in the distribution of pelagic fish stocks and industrial purse-seine fishing are hypothesised to be a key driver of seabird population decreases in the southern Benguela. We investigated the foraging behaviour of breeding Cape cormorants by deploying GPS and temperature–depth recorders on 24 breeding adults from three islands off the coast of South Africa, two of them to the north of Cape Point and a third farther south on the western Agulhas Bank. This provided the first measures of foraging dispersal by a cormorant in the Benguela system, and enabled a comparison of foraging behaviour between birds from these islands. Foraging trips of Cape cormorants lasted between 17 min and >7 h, at a maximum distance of between 2 and 58 km away from their colony. Foraging effort was significantly greater for birds from farther north off the West Coast in terms of trip duration, distance travelled, number of dives and time spent flying compared to those from the southernmost island (Dyer), which is probably a response to low prey availability in the north. Coastal reserves that exclude pelagic fishing from inshore feeding grounds around Cape cormorant breeding colonies may result in increased local prey availability, which would benefit Cape cormorant populations.     

Assessing trophic adaptability is critical for understanding the response of predatory fishes to climate change: a case study of Pomatomus saltatrix in a global hotspot

There is a growing need to incorporate biotic interactions, particularly those between predators and their prey, when predicting climate-driven shifts in marine fishes. Predators dependent on a narrow range of prey species should respond rapidly to shifts in the distribution of their prey, whereas those with broad trophic adaptability may respond to shifts in their prey by altering their diet. Small pelagic fishes are an extremely important component of the diet of many marine predators. However, their populations are expected to shift in distribution and fluctuate in abundance as the climate changes. We conducted a comparative study of the seasonal diet of adult Pomatomus saltatrix over two periods (June–December 2006 and 2012) and examined the available data on small pelagic fishes biomass in a global hotspot (the coastal region of southern Angola, southern Africa) to gain an understanding of the tropic adaptability of the species. Despite a drop (630 000 t to 353 000 t) in the abundance of their dominant prey (Sardinella aurita) in the region, it remained the most important prey item during both study periods (Period 1 = 99.3% RI, Period 2 = 85.3% RI, where %RI is a ranking index of relative importance). However, the diet during Period 2 was supplemented with prey typically associated with the nearshore zone. The seasonal data showed that P. saltatrix were capable not only of switching their diet from S. aurita to other prey items, but also of switching their trophic habitat from the pelagic to the nearshore zone. These findings suggest that P. saltatrix will not necessarily co-migrate if there is a climate-driven shift in the distribution of small pelagic fishes (their dominant prey). Accordingly, understanding the trophic adaptability of predators is critical for understanding their response to the impacts of climate change.     

Top predators in the Benguela ecosystem — implications of their trophic position

Many top predators in the Benguela ecosystem feed on prey species targeted by commercial fisheries. Their roles as indicators of the state of exploited prey resources, as competitors with commercial fisheries for resources, and as susceptible to impact from commercial fishing on those resources are briefly considered. Trends in the occurrence of anchovy Engraulis capensis and pilchard Sardinops ocellatus in the diet of Cape gannets Morus capensis off South Africa's west coast are related significantly to survey estimates of the abundance of these fish species, and they provide useful confirmation of those estimates. In the 1980s, anchovy decreased in the diet of Cape gannets, but pilchard increased. In both the northern and southern portions of the Benguela system, groundfish were thought to eat most (66–73%) of the total quantity of cephalopods and vertebrates consumed by predators and man in the 1980s. South African fur seals Arctocephalus pusillus pusillus, predatory pelagic fish and man removed roughly equal amounts, with squids, seabirds and cetaceans having a smaller impact. In the 1980s, man and seals removed about two million tons live mass more than in 1930. Indices of the rate of natural mortality of anchovy and pilchard attributable to Cape gannets are not related to biomass of the prey species. That for anchovy was high in 1989 when a poor anchovy year-class was formed. Decreased abundance of anchovy led to poor breeding by Cape cormorants Phalacrocorax capensis in 1989 and 1990. A model linking the Cape cormorant population with anchovy is used to explore the impact of possible exploitation strategies for anchovy on Cape cormorants.     

An overview of the living marine resources of Namibia

This paper gives an overview of the main living marine resources of Namibia. It focuses on the scientific research conducted during the past decade as input to the management of these resources. The distribution and habitats of the most important harvested species and the main seabird populations are briefly described and discussed. The life histories of the major exploited species are summarized, with emphasis on spatial and temporal spawning patterns, dispersal of early life stages, migration patterns of recruits and adults, and diet, the latter particularly as it relates to potential competition between species. A number of commercially important species, such as the hake Merluccius capensis and M. paradoxus, deep-sea red crab Chaceon maritae, West Coast rock lobster Jasus lalandii, skipjack tuna Katsuwonus pelamis, southern albacore Thunnus alalunga and to a lesser extent Cape horse mackerel Trachurus capensis, southern African sardine Sardinops sagax and Cape anchovy Engraulis capensis, are distributed across national boundaries, requiring regional cooperation in research and management. The history and current status of the major fisheries is discussed. Over the past 30–40 years total annual catches have declined from a peak of around 2 million tons in the late 1960s to less than a million tons in the 1990s. This decline has been due, mainly, to a collapse in the sardine stock in the late 1960s and 1970s, and a reduction in the catches of hake and horse mackerel under a conservative management strategy in the past decade. Changes in the abundance and distribution of commercially important species, as determined by acoustic and trawl surveys and catch-based analytical methods, are presented. The effect of major environmental anomalies on the distribution and abundance of the resources in recent years is discussed. The most dramatic anomaly in recent years was the wide-scale advection of low-oxygen water into the northern Benguela from the Angola Dome in 1994, and the subsequent Benguela Niño of 1995, which appear to have severely impacted the Namibian sardine population and many other resources. The present socio-economic value of the Namibian fishing industry is given together with the broad policy, legislation and formal structures for managing the living marine resources.     

Breeding success of African penguins Spheniscus demersus at Dassen Island,especially after oiling following the Apollo Sea spill

The reproductive success of African penguins Spheniscus demersus at Dassen Island from 1994 to 2000 was variable, but much higher than previously reported figures for the species. Breeding success was positively related to the abundance of anchovy Engraulis encrasicolus and sardine Sardinops sagax, and the high reproductive output during the study was attributed to the large biomass and high availability of these two species for much of the study period. De-oiled penguins from the Apollo Sea spill had a slightly lower hatching success but a significantly lower overall breeding success than that of un-oiled birds, driven mainly by the reduced number of chicks which fledged. Nests with two de-oiled Apollo Sea parents were less successful than nests with one de-oiled bird. There was increased mortality of chicks 40 days and older in nests with de-oiled birds. Chicks from nests with one de-oiled Apollo Sea parent grew at a similar rate to chicks from nests with no de-oiled parents. However, chicks from nests that comprised two de-oiled Apollo Sea birds had significantly slower growth rates than these other two groups. Breeding success and chick growth at nests with de-oiled birds were more negatively impacted when feeding conditions were less favourable. These results suggest that one of the main reasons for lower breeding success in de-oiled birds was their reduced ability to provision chicks, especially during the period in which the energy demands of the chicks is greatest. The rate of mate fidelity was lower in de-oiled birds than un-oiled birds, and there was a positive association between mate fidelity and breeding success.     

Use of particle-size data to predict potential pelagic-fish yields of some southern African areas

A particle-size spectrum model is used to estimate standing stocks of some pelagic fish from measurements of phytoplankton chlorophyll in two exploited regions in southern Africa (southern Benguela and off South West Africa/Namibia) and in two unexploited regions (the Agulhas Bank and off the eastern Cape Province). The model is based on the assumption that equal biomasses occur in logarithmically equal size classes in the pelagic marine environment. Phytoplankton, with an equivalent spherical diameter ranging from 1 to 128 μm, occupy 21 size classes on the logarithmic scale. Two different size ranges are assumed for some commercial, pelagic fish species, equivalent to exploitation with two different purse-seine mesh sizes. A mesh of 12,7 mm would catch 8 size classes of pilchard, horse mackerel and anchovy whereas a mesh of 32 mm would catch only 3,3 size classes of pilchard and horse mackerel. From the model, the potential biomass of these commercial pelagic fish is estimated, after allowing for the presence of other commercial and non-commercial fish and other taxa in the exploited size range. Total pelagic fish production is estimated by assuming constant turnover rates of 1·y^?1 and 1,5·y^?1 when exploited with 32 and 12,7 mm mesh nets respectively. Consideration of the maximum and mean reported catches in the exploited areas indicates that only some 25 per cent of pelagic fish production is exploitable by man. On this basis, the unexploited Agulhas Bank region may yield some 400 000 metric tons (wet) of pelagic fish of the species considered, and the East Coast region some 90 000 tons. Exploitation in these regions cannot be recommended, because the Agulhas Bank is an important spawning ground for many pelagic species, and the fish in both regions probably act as a reserve buffer for the heavily exploited pelagic resource of the Western Cape.     

Diving behaviour of African penguins: do they differ from other Spheniscus penguins?

African penguins Spheniscus demersus closely resemble Magellanic S. magellanicus and Humboldt S. humboldti penguins and have similar breeding and feeding ecologies. Adults feed on pelagic schooling fish in continental shelf waters, but African penguins have been reported to have shallower dive angles and remain submerged longer for dives to a given depth than their congeners. The few data for African penguins were gathered using relatively large time-depth recorders. We measured diving behaviour of 36 African penguins provisioning small chicks at three colonies near Cape Town, South Africa. Maximum and mean dive depths were 69m and 14m respectively. Diving took place mainly during the day. Although dive depths differed between colonies, there were no significant differences in dive duration or maximum, median or mean depth. Total dive duration, descent time, bottom time, ascent time and dive angle all were strongly correlated with the maximum depth attained. The diving behaviour of African penguins is similar to that of its congeners. Diving performance probably was compromised by the data-logger used in the previous study. Comparative data from Humboldt penguins also indicate potential biases in an earlier study of this species. Care is needed when comparing the diving performance of penguins measured using different loggers.     

Diet of sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The diet of sardine Sardinops sagax in the southern Benguela was investigated by microscopic examination of stomach contents. The relative dietary importance of prey size and prey type was assessed by calculating the carbon content of prey items. Sardine is an omnivorous clupeoid, ingesting both phytoplankton and zooplankton, with the relative importance of these two food types varying both spatially and temporally. Stomach contents were numerically dominated by small prey items, principally dinoflagellates, followed by crustacean eggs, cyclopoid copepods, calanoid copepods and diatoms. Virtually all prey items ingested by sardine were <1.2 mm maximum dimension, the particle size below which sardine only filter-feed. Despite the numerical dominance by phytoplankton, zooplankton contributed the major portion to sardine dietary carbon, small calanoid and cyclopoid copepods, anchovy eggs and crustacean eggs being the primary prey types. These results indicate that, like anchovy Engraulis capensis, sardine in the southern Benguela are primarily zoophagous, and contrast with earlier dietary studies on sardine in the region. However, the two species appear to partition their prey on the basis of size; sardine consume small zooplankton, whereas anchovy consume large zooplankton. This difference has been observed in other upwelling ecosystems where the two genera co-exist and is likely to contribute to the regime shifts observed between sardine and anchovy.     

Mesozooplankton dynamics in the Benguela ecosystem,with emphasis on the herbivorous copepods

Recent research developments on the ecology, dynamics and trophic position of copepods in the Benguela ecosystem are synthesized. Attention is focused on herbivorous species of the southern Benguela and how they cope with the physical and biological variability characteristic of this upwelling region. Copepods constitute on average approximately half of the total zooplankton carbon and. are most abundant during the upwelling season. They are able to maintain large population densities within local coastal upwelling areas by combining ontogenetically based vertical migration behaviour with features of the current system. Some species have developed finely tuned strategies to overcome periods of starvation between upwelling bouts by storing lipid reserves or by entering temporary developmental arrest. In situ measurements of production rates of local species are among the highest recorded for copepods. Despite an apparent excess of food, copepods exert only limited impact on the phytoplankton, removing on average <25 per cent of that available daily. Indirect estimates of carbon flux indicate that 11–25 per cent of copepod daily ration is used for egestion of faecal pellets. Copepods are the preferred prey of a wide variety of invertebrate and vertebrate predators. Large copepods in particular are important in the diet of commercially exploited pelagic fish. Localized areas of low abundance of copepods have been found in association with high densities of anchovy during peak spawning and recruitment periods. Copepods may therefore constitute a central limiting factor for pelagic fish production in the southern Benguela.     

Diel feeding of the transparent goby Aphia minuta (Pisces,Gobiidae) in the Northwestern Adriatic Sea in spring time

The present study provides the first estimate of the daily ration for a goby species in the Mediterranean Sea, using a new approach to determine the mass of fish stomach contents through the sum of individual prey dry mass derived indirectly from prey size. Diel feeding activity and daily ration of the pelagic goby Aphia minuta were studied under natural conditions in the coastal waters off Comacchio (Northwestern Adriatic Sea) during the spring season. A total of 318 individuals of A. minuta, collected by 14 trawls carried out on 4 consecutive days, was examined for gut contents. The diel pattern of the vacuity index and the stomach fullness index indicated that A. minuta has nocturnal feeding behaviour. The daily ration, computed over a period of 16 h, was equivalent to 2.23% wet body mass.