Estimates of standing stock,production and consumption of meso- and macrozooplankton in the Benguela ecosystem

Attempts are made to consolidate current information on estimates of standing stock, production and consumption of meso- and macrozooplankton from the shelf region of the west and south coasts of southern Africa for inclusion in a network analysis of carbon flow in the Benguela system. The meso- and macrozooplankton communities consist chiefly of copepods and euphausiids respectively. Although geographic and seasonal differences in standing stock are reasonably well described for the inner shelf (<200 m), knowledge of production and rate processes affecting standing stock is limited, and extrapolation to the whole shelf area (<500 m) provides only a crude appraisal of the real situation. Despite the uncertainties, direct measurements have improved on estimates previously inferred. It seems likely that grazing by meso- and macrozooplankton plays a minor role in phytoplankton losses in the Benguela system. However, it would appear that predation by macrozooplankton, particularly under swarming conditions, has an appreciable impact on mesozooplankton biomass.     

Forecasting in South African pelagic fisheries management: the use of expert and decision support systems

Understanding long-term variability of pelagic fish populations is important in developing forecasting strategies for fisheries management and planning. However, many current fisheries models have only short-term datasets available, whereas those of suitable duration often lack reliability. As resources are placed under increasing pressure, all available information should be used to assist management. Two simple rule-based deterministic modelling approaches are described, which use semi-quantitative and qualitative rules to relate recruitment success of South African anchovy Engraulis capensis to physical and biological indices. The first model relates recruitment success to indices of wind and sea surface temperature by way of a rule-based decision support system. In the second model, significant environmental and biological factors were identified and related to anchovy recruitment by way of an expert system approach. These two approaches are evaluated and compared. It is suggested that these types of models, when satisfactorily validated, have great potential in supporting the future management of the South African anchovy fishery in the dynamic environment of the Benguela Current.     

Population Numbers of Fur Seals at Prince Edward Island,Southern Ocean

During the period 17–22 December 2001, the onshore distribution and the abundance of Antarctic fur seals Arctocephalus gazella and Subantarctic fur seals A. tropicalis were determined for Prince Edward Island. Two breeding colonies of Antarctic fur seals were located on the south-east coast of the island; the first a mixed (with Subantarctic fur seals) breeding colony with an estimated 24 pups on a vegetated promontory on the northern section of Boggel Beach, and the second, a presumably pure Antarctic fur seal breeding colony with an estimated 380 pups, at Penguin Beach. At a mean intrinsic rate of natural increase of 16.2% per year, Antarctic fur seals appear to be in the rapid recolonization phase of population growth. Breeding colonies of Subantarctic fur seals, largely found on the entire east coast, produced an estimated 15 000 pups, and the population had maintained a mean intrinsic rate of natural increase of some 9.5% per year since 1987/88.     

Anchovy spawning in relation to the biomass and the replenishment rate of their copepod prey on the western Agulhas Bank

Anchovy biomass and copepod standing stocks and growth rates on the Agulhas Bank were compared during the peak spawning period (November) in 1988 and 1989. In 1988, copepod biomass over the western Agulhas Bank was low (1,0 g dry mass·m^?2) relative to anchovy biomass there (14,7 g dry mass·m^?2). In November 1989 in the same area, fish biomass was much lower (5,7 g dry mass·m^?2), following a recruitment failure, and copepod biomass was higher (2,4 g dry mass·m^?2), possibly as a result of lesser predation by anchovy. By contrast, the eastern Agulhas Bank had a larger biomass of copepods (4–6 g dry mass·^?2) and a lower biomass of anchovy during both years. Knowing, from laboratory studies, that a prey biomass of 0,78 g·m^?2 is required for fish to obtain their daily maintenance ration, it is suggested that spawning on the western Agulhas Bank was food-limited in 1988. Copepods on the western Bank may be replaced by local growth or transport from the eastern Bank. Growth rates of copepods on the western Bank were 10–50 per cent of maximum in 1988, but total production (c. 100 mg dry mass·m^?2·day^?1) was low, primarily because biomass was low and less than the rate of consumption by anchovy (243 mg copepod dry mass·m^?2·day^?1). On the eastern Bank, copepod production exceeded anchovy consumption and it is concluded that the flux of copepod biomass onto the western Bank may be as important as local growth in replenishing copepod stocks there. Feeding conditions for anchovy on the western Agulhas Bank are often marginal compared to the situation on the eastern Bank, and it is suggested that the selection of the western Bank as the major spawning area is related more to the success of transport and survival of eggs and larvae on the West Coast recruiting grounds than to feeding conditions per se.     

A preliminary investigation on how meteorological changes may affect beach-seine catches of Lougo vulgaris in the Thracian Sea (Eastern Mediterranean)

Daily catches of Loligo vulgaris and the fishing effort of commercial beach-seiners operating along the coasts of the Thracian Sea were recorded between October 1994 and May 1995. Taking into account the spatial distribution of the ports, the squid fishing grounds and the daily activity of the beach-seiners, five coastal zones were determined for the monitoring of fishery data. Multivariate analysis failed to show significant differences between the monthly catch rate (cpue) in the zones. However, Loligo vulgaris cpue was clearly seasonal, with a distinct peak in November and an evident decline from winter to spring. Temperature, rainfall and local wind records were used in making a preliminary investigation into meteorological factors that may affect beach-seine catches of squid. Multiple linear regression analysis revealed that temperature is a highly significant explanatory variable for cpue variation, whereas the intensity of the wind plays an important role, but mostly during winter. Rainfall was significantly related to variation in cpue in one zone. Cpue was generally negatively correlated with air temperature and rainfall. Most significant correlations of cpue with wind were also negative.     

Counts of Surface-nesting Seabirds Breeding at Prince Edward Island,Summer 2001/02

The first midsummer survey of surface-nesting seabirds at Prince Edward Island was conducted during December 2001. It was also the first census for most species since the early 1980s. Despite concerns about the impacts of longline fishing mortality on various albatrosses and giant petrels, there was no evidence that populations of these species had decreased. Indeed, the estimate of wandering albatrosses Diomedea exulans (Vulnerable) has increased significantly, making Prince Edward Island equal with Marion Island as supporting the largest single-island populations of this species. Species that underwent significant decreases were macaroni penguins Eudyptes chrysolophus (Vulnerable), Crozet shags Phalacrocorax [atriceps] melanogenis and Kerguelen terns Sterna virgata (Near-Threatened). The reasons for these decreases are unclear, but for macaroni penguins may be partly a consequence of competition for space with the burgeoning population of fur seals Arctocephalus spp. The 2001 survey increased the population estimates for Subantarctic skuas Catharacta antarctica, lightmantled sooty albatrosses Phoebetria palpebrata (Vulnerable) and southern giant petrels Macronectes giganteus (Vulnerable), mostly as a result of greater coverage than for previous counts. The 2001 survey confirms that Prince Edward Island remains a globally important breeding site for seabirds.     

Populations of Surface-nesting Seabirds at Marion Island, 1994/95–2002/03

During the 1990s and early 2000s, populations of surface-nesting seabirds at Marion Island showed different trends, but for the majority of species numbers decreased. Reduced numbers of gentoo penguins Pygoscelis papua, eastern rockhopper penguins Eudyptes chrysocome filholi, Crozet shags Phalacrocorax [atriceps] melanogenis and probably macaroni penguins E. chrysolophus are most plausibly attributed to an altered availability of food. Decreases in numbers of dark-mantled sooty albatrosses Phoebetria fusca, light-mantled sooty albatrosses P. palpebrata, southern giant petrels Macronectes giganteus and possibly northern giant petrels M. halli may have resulted from mortality of birds in longline fisheries. However, populations of wandering Diomedea exulans and grey-headed Thalassarche chrysostoma albatrosses fluctuated around a stable level. Numbers of Subantarctic skuas Catharacta antarctica and kelp gulls Larus dominicanus breeding at Marion Island also decreased. Kerguelen Sterna virgata and Antarctic S. vittata terns remain scarce at the island. Trends for king penguins Aptenodytes patagonicus were not reliably gauged, but numbers probably remained stable or increased. There were large fluctuations in numbers of king penguin chicks surviving to the end of winter.     

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.     

Population dynamics of southern elephant seals: a synthesis of three decades of demographic research at Marion Island

Southern elephant seal Mirounga leonina numbers declined precipitously throughout most of their circumpolar distribution since the 1950s. A long-term intensive demographic programme was initiated in 1983 on the relatively small population of southern elephant seals at sub-Antarctic Marion Island in an attempt to identify causative mechanisms associated with this decline. Weaned pups have been tagged annually since 1983, and this has produced a large number of individuals of known identity. A regular resighting programme yielded a mark-recapture dataset that has been subjected to numerous survival-based models. This ongoing programme produced a substantial body of scientific literature on population growth patterns, vital rates (survival and fecundity) and population regulation in southern elephant seals, which are reviewed in this synthesis. We briefly describe the analytical framework common to much of the demographic research, highlight important conclusions concerning population regulation of elephant seals at Marion Island, and discuss priorities for future research.     

Estimates of phytoplankton and bacterial biomass and production in the northern and southern Benguela ecosystems

Available data on phytoplankton and bacterial abundance and production off the coasts of southern Africa (to the 500 m depth contour) have been assembled and analysed for a network analysis of carbon flow in the Benguela ecosystem. Phytoplankton carbon biomass (from measurements of chlorophyll a) in the northern Benguela (2 558 300 tons) was considerably higher than in the southern Benguela (671 420 and 516 400 tons for the West and South coasts respectively). However, overall annual production (from C¹⁴-uptake measurements) was similar, 77 416 608, 76 399 973 and 78 988 020 tons C·year^?1 respectively. Phytoplankton respiration and sedimentation losses were calculated as functions of primary production and therefore followed similar trends. From the most conservative estimates (mean bacterial biomass of 10 mg C·m^?3 and average P:B of 0,2·day^?1) bacterial biomass is 2–7 per cent of phytoplankton biomass in the northern and southern Benguela, and bacterial production is 3–5 per cent of primary production. Assuming a net growth yield of 30 per cent, bacteria would need to consume 9–15 per cent of the total primary production in order to meet their requirements for carbon consumption. Calculations based on a mean bacterial biomass of 40 mg C·m^?3 and a mean growth rate of 0,5·day^?1 in the upper 30 m of the water column show bacterial biomass to be 8–27 per cent of phytoplankton biomass and bacterial production to be 26–44 per cent of phytoplankton production. Bacterial carbon consumption requirements at these rates amount to 86–147 per cent of total primary production.     

The Distribution and Spread of Alien Vascular Plants on Prince Edward Island

Surveys of alien plants at subantarctic Prince Edward Island in 2001 show that the ranges of all three introduced species have increased since the last survey in 1998. Poa annua, the longest-established species, increased its range substantially after 1987, prior to which it was confined to a single site for more than 20 years. It remains largely restricted to sites characterized by intense disturbance by seals and seabirds. Sagina procumbens, the most recently discovered alien plant, has spread even more rapidly (up to 800 m year^?1) and has colonized the west coast of the island (a leap of at least 3 km). Unlike Poa annua, it is not restricted to animal-disturbed areas, although its seeds probably are dispersed by both birds and fur seals. Cerastium fontanum also continues to spread at the island, but remains confined to the western coastal plain, where it occurs mostly on dry feldmark slopes or erosion scars. All three species have expanded their ranges faster than at Marion Island, possibly because of a warmer recent climate and higher densities of seabirds and seals at Prince Edward Island. The ranges of alien plants are likely to continue to expand, with S. procumbens causing considerable changes in the island's terrestrial ecology.     

Recent population trends of sooty and light-mantled albatrosses breeding on Marion Island

Sub-Antarctic Marion Island is one of the few islands where both species of Phoebetria albatrosses breed sympatrically. The last published assessment of their population trends, which reported counts up to 2008, concluded that the numbers of breeding pairs of sooty albatross P. fusca (Endangered) were decreasing, whereas numbers of light-mantled albatross P. palpebrata (Near Threatened) were increasing. Extending the counts to 2014 reversed these trends, with numbers of sooty albatrosses increasing from 2006 to 2014, and numbers of light-mantled albatrosses decreasing from 2007 to 2014. Confidence in island-wide counts is low due to the cryptic nature of the albatrosses on their largely inaccessible cliff-side nest sites, as well as counts for sooty albatrosses taking place late in the incubation period when 10?20% of nests have already failed. Given the greater conservation concern for the sooty albatross, we recommend that dedicated annual counts be conducted during the early incubation period, and be repeated shortly after the chicks hatch (late December), mid-way through the nestling period (late February) and prior to fledging (late April), to give a better idea of breeding success. Count zones also should be revised to facilitate more accurate counts, ensuring more reliable estimates of sooty albatross population trends at Marion Island.     

Effect of human disturbance on the behavioural responses and offspring survival of grey-headed albatrosses Thalassarche chrysostoma at Subantarctic Marion Island

Nest monitoring is commonly used to gather important seabird demographic data, but monitoring itself can affect seabird behaviour and offspring survival. The effect of monitoring on grey-headed albatrosses Thalassarche chrysostoma at Subantarctic Marion Island was investigated. The behaviour of chicks close to fledging and of brooding adults was recorded in research and non-research sections of the colony. This was done prior to, during and after researcher/s entered the research sections of the colony. During disturbance, both chicks and adults in research sections were more likely to be standing and to be more responsive than during the pre-disturbance phase. However, adults appeared to be more sensitised to disturbance and this is likely because of being previously handled. Offspring survival in research sections was compared with survival in (a) sections less-frequently entered (2004/2005) and (b) sections monitored through fixed-point photography (2005/2006 and 2006/2007). In February of each of the three years of study, offspring survival was 62.5%, 55.8% and 71.8% respectively. Disturbance level, size of section and the interaction of these two factors did not explain significant amounts of the among-section survival variance, but rainfall explained almost all of the temporal variation in offspring survival in 2006/2007. Although research activities influenced the short-term behavioural responses of chicks and adults in the research sections of the colony, it did not seem to influence the survival of offspring. Nevertheless, fixed-point photography is recommended as a non-intrusive monitoring method.     

Impact of predation by Cape fur seals Arctocephalus pusillus pusillus on Cape gannets Morus capensis at Malgas Island,Western Cape,South Africa

Cape fur seals Arctocephalus pusillus pusillus were estimated to kill some 6 000 Cape gannet Morus capensis fledglings around Malgas Island in the 2000/01 breeding season, 11 000 in 2003/04 and 10 000 in 2005/06. This amounted to about 29%, 83% and 57% of the overall production of fledglings at the island in these breeding seasons respectively. Preliminary modelling suggests this predation is not sustainable. There was a 25% reduction in the size of the colony, the second largest of only six extant Cape gannet colonies, between 2001/02 and 2005/06. There has been a large increase in predation by Cape fur seals on seabirds around southern African islands since the mid-1980s, coincidental with both an increase in the seal population, altered management of the islands and an altered distribution of prey for gannets and seals. At Malgas Island, most gannet fledglings were killed between 10:00 and 18:00, the period when most are in the water around the island, from mid-January to mid-March, the main fledging period. The Cape gannet is classified as Vulnerable.     

Congruent,decreasing trends of gentoo penguins and Crozet shags at sub-Antarctic Marion Island suggest food limitation through common environmental forcing

Numbers of gentoo penguins Pygoscelis papua and Crozet shags Phalacrocorax [atriceps] melanogenis breeding annually at Marion Island, one of South Africa's Prince Edward Islands in the South-West Indian Ocean, were strongly correlated over 19 split-years from 1994/1995 to 2012/2013. Both species decreased between the mid-1990s and the mid-2000s, exhibited a partial recovery in the late-2000s and then decreased to their lowest recorded levels in 2012/2013. In both instances, the partial recoveries in the late-2000s were associated with improved breeding success. At a colony of gentoo penguins, breeding success was negatively correlated with the date of arrival of adults to breed. Gentoo penguins and Crozet shags are demersal feeders in inshore waters around Marion Island and there is considerable overlap in the composition of their diets. Therefore, trends in their populations may be driven by food availability, which is likely to be influenced by benthic production around the island. We propose that, in South Africa, and based on the current assessment, the Crozet shag, which elsewhere breeds only at the Crozet Islands, is now Critically Endangered, and the more widely ranging gentoo penguin is Endangered.     

Age at first breeding and change in plumage of kelp gulls Larus dominicanus in South Africa

In South Africa, kelp gulls Larus dominicanus first breed when 4 years old. At least 50% of 4-year-olds breed; virtually all older birds breed annually. Most chicks leave natal colonies in the austral autumn and winter, when about 6 months old, and 60% of them may not return until aged 2 or 3 years. One 3-year-old showed signs of defending territory, but birds only attain full adult plumage when aged 4. The plumage of 3-year-olds is similar to that of adults, but the white spots on the outer primaries are less well developed and the secondaries and inner primaries have no white trailing edge. Chicks are dark, whereas birds aged 1 and 2 years gain increasing amounts of pale or white plumage on the face, neck, underparts and tail.     

Nest Densities of the Wandering Albatross Diomedea Exulans at the Prince Edward Islands,Estimated Using Gps

Hand-held Global Positioning System (GPS) receivers provide opportunities for detailed and rapid mapping of features, including biological ones, further enhanced by the removal during 2000 of "selective availability". GPS was used to map, describe and compare nest densities within wandering albatross Diomedea exulans colonies at subantarctic Marion and Prince Edward islands. On Prince Edward Island, the coordinates of 1 061 wandering albatross nests were determined and, on Marion Island, 1 779 nests. For describing nest densities of wandering albatrosses, a 50-m grid is recommended, at which scale, the densest area of Prince Edward Island was in Albatross Valley, where the area of the colony was 46 ha and nest density was 22.3 nests ha^?1. For Marion Island, the total area of the wandering albatross colonies was 306 ha and the nest density was 5.8 nests ha^?1. In the three study colonies there (Macaroni Bay, 28 nests; Sealer's Beach, 117 nests; Goney Plain, 140 nests), the density statistics did not differ greatly from the overall densities on the island, with overall mean densities of 4.9, 5.7 and 8.0 birds ha^?1 respectively. Although comparisons with nest densities at other breeding colonies are uncertain because of differing methods of computing them, the nest densities in Albatross Valley lie within the reported ranges for other colonies of great albatrosses.     

An analysis of yield variability from three harvesting strategies in the South African anchovy fishery,under conditions of randomly fluctuating recruitment success

An age-structured population model was used to examine the variability of yield created by randomly fluctuating recruitment success in the South African anchovy Engraulis capensis stock. Three different harvesting strategies were examined: (1) constant age-specific pattern of fishing mortality, (2) constant quota with effort limitation and (3) annual quota adjustments by means of an F _msy procedure. Variable recruitment was generated by a stochastic stock-recruit relationship, and mean yields, mean spawner biomasses and probability distributions of yield were calculated at each exploitation level after 150 projections from the population structure of anchovy in 1981. Under conditions of constant fishing mortality, the variability of yield increased continuously as the exploitation rate was increased. The maximum average yield (MAY) is the stochastic equivalent of the maximum sustainable yield (MSY) of deterministic models, but an attempt to sustain this MAY by a constant quota resulted in a very high risk of stock collapse. An Fmsy policy based on pre-season adult biomass resulted in more variable yields than were obtained at the equivalent constant fishing mortality at age. Stability of yield therefore demanded the acceptance of lower average yields than could be attained in the long term by quota adjustment. It was considered that the South African purse-seine industry could not cope with the wide fluctuations of yield necessary to attain MAY. Specific management policies sufficiently robust to withstand both fluctuating recruitment success and the uncertainties in the parameters of working population models would be required.     

Southward extension of the reproductive range of Macroramphosus scolopax in the upwelling area off North-West Africa

Ichthyological literature gives the distribution of M. scolopax as along the whole coast of West Africa. However, analysis of this literature and the results of trawling surveys in the North-East Atlantic suggest only exceptional occurrences in the tropics. Plankton surveys off N.W. Africa, up to and including 1977, rarely yielded M. scolopax larvae off the coast of the southern Sahara and not at all off Mauritania, though a few specimens were collected after having been swept southwards in the open ocean by the Canary Current. Larvae did occur off northern Mauritania in February 1983. Details of distribution, abundance and length composition of early stages caught during the period January–April 1983 are given. It is concluded that the reproductive range of M. scolopax (as well as that of some other temperate species) extends south of that documented and that its spawning season there may be shorter than in the centre of this species' distribution. The findings are discussed in relation to interannual variation of upwelling intensity and the current system off N.W. Africa.     

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.