Fishes and salinities in the St Lucia estuarine system—a review

The recorded salinity ranges of freshwater, estuarine and marine fish species in Lake St Lucia, a Ramsar and World Heritage Site, are documented. The freshwater group is most diverse and abundant under oligohaline conditions, although the Mozambique tilapia (Oreochromis mossambicus) was common under all salinity regimes. Estuary resident species also favoured oligohaline conditions but, in contrast to the freshwater taxa, were well represented in salinities up to 40 ‰. The marine group was most diverse and abundant within the salinity range 10–40 ‰, but a large number of species could also be found in salinities up to 70 ‰. Very few fish species were able to tolerate salinities between 70 ‰ and 110 ‰, with only O. mossambicus surviving for extended periods in salinities above 110 ‰. All the aquatic macrophytes and most of the zoobenthos within the lake appear to die out within the salinity range of 50–60 ‰, thus creating additional stress to those fish present under such conditions. The food resources least affected by extreme hypersalinity are the microphytobenthos and detritus food chains, with detritivorous fishes being dominant when the lake is in this state. Mass mortalities of fishes in Lake St Lucia have been recorded under both low (<5 ‰) and high salinity (>70 ‰) conditions. The fish kills are often triggered by exceptionally low or high water temperatures which affect the osmoregulatory abilities of these species. Hypersaline conditions and fish mortalities under the most recent closed estuary mouth conditions (2002–2005) are reviewed. If the surface area of St Lucia (35,000 ha) is compared to the total surface area of all South African estuaries (approximately 70,000 ha), then the possibility exists that the loss of the Lake St Lucia nursery area for estuary-associated marine fish species over the past few years may cause significant short-term declines in the future abundance of these taxa on both a local and regional scale.     

SOME NOTES ON THE BIRDS OF THE KAOKOVELD

Whitfield, A. K. &; Blaber, S. J. M. 1978. Feeding ecology of piscivorous birds at Lake St Lucia, Part 1: Diving birds. Ostrich 49:185-198.

The diets of three species of diving piscivorous birds at Lake St Lucia, Natal, South Africa during 1975 and 1976 are described and related to availability and abundance of different prey species. Mugilidae and Clarias gariepinus were the chief prey of the Fish Eagle Haliaeetus vocifer, while Johnius belengcrii and Sarotherodon mossambicus were, respectively, the main food items of Caspian Terns Hydroprogne tschegruva, and Pied Kingfishers Ceryle rudis. The methods of prey capture used by the three species of birds are described.

Predation on particular species of fish was related primarily to their swimming depth, abundance, size and distribution in littoral areas. Fishing habits, densities and foraging periodicity of the birds are discussed with regard to physical parameters such as thermals, wind speeds and water turbidity. The timing of breeding seasons at Lake St Lucia is related to fish densities, lake levels, turbidity of the water and air temperatures.     

THE PROBLEM OF NOMENCLATURE

Whitfield, A. K. &; Blaber, S. J. M. 1978. Feeding ecology of piscivorous birds at Lake St Lucia, Part 3: Swimming birds. Ostrich 50:10-20. The diets, foraging periodicities and feeding behaviour of the Reed Cormorant Phalacrocorax africanus, Whitebreasted Cormorant Phalacrocorax carbo and White Pelican Pelecanus onocrotalus were studied at Lake St Lucia, Natal, South Africa, during 1975 and 1976. The Reed Cormorant fishes in shallow water within 100 m of the shore and mainly caught Sarotherodon mossambicus and Solea bleekeri, while the Whitebreasted Cormorant caught Mugilidae, Rhabdosargus sarba and Thryssa vitrirostris in deeper water. The diet of the White Pelican followed three distinct phases: a pre-incubation phase when the birds followed and preyed heavily on migrating Mugil cephalus shoals; an incubation and post-incubation phase when the adults flew a round trip of 200 km to the north to obtain freshwater fish, mainly cichlids, from the Pongolo pans where fishes were concentrated and densities high; and a post-fledgling phase when both adults and juveniles fed on a variety of marine species of fish in Lake St Lucia for about a month before dispersing to other areas. The feeding and breeding of Reed Cormorants and White-breasted Cormorants is discussed in relation to wind speeds, water turbidity and flooding of backwaters. The diet and long-range foraging behaviour of White Pelicans at St Lucia are compared with data from other African lakes. The breeding season of White Pelicans at St Lucia is related to availability of fish and inaccessibility of the breeding site to predators. The latter is determined by lake levels.     

The influence of turbidity on juvenile marine fishes in estuaries. part 1. field studies at Lake St. Lucia on the southeastern coast of Africa

Lake St. Lucia, the largest estuarine system in Africa (325 km²), was chosen as the field study area for a 3.5-yr ((1980)–83) investigation into relationships between water turbidity and estuarine fish distribution. The variety of habitats, from clear water, open sandy shores to shallow muddy substrata and turbid waters, together with high species diversity (108 species) rendered the area suitable for this study. The relationships between fish distribution and environmental factors were monitored by monthly seine netting of fishes at seven sites representative of the range of conditions in St. Lucia. Simultaneously, water turbidity, salinity, and temperature were recorded.The possible influences of substratum type and food availability were also investigated by using recently published data on invertebrate benthos and Zooplankton distributions. Published data were also used to determine the diet of the common fish species. The results showed that the distribution of juveniles of the 20 commonest fish species were statistically correlated only with water turbidity, water temperature, and food availability. The correlation with temperature was related to seasonal not spatial temperature patterns.Turbidity and food type influences were difficult to separate but exceptions were the anchovyThryssa vitrirostris (Gilchrist and Thompson) and the soleSolea bleekeri Boulenger which occurred only in turbid water despite the widespread occurrence of their prey, andGenes acinaces Bleeker,G. rappi (Barnard), andG. fllamentosus Cuvier, all of which occurred only in clear water although the greatest densities of their bivalve prey were in turbid waters. Similarly, the sparidsRhabdosargus holubi (Steindachner) andR. sarba (Forsskal) were distributed according to turbidity and not their preferred foods.Principal component analysis with a minimum spanning tree plot and a canonical correlation test showed that the fish fauna could be divided into five groups according to their occurrence in various turbidities. These were: clear water species (e.g. Gerreidae) in < 10 NTU, clear to partially turbid species (e.g.Liza dumerilii (Steindachner) andL. macrolepis (Smith)) in < 50 NTU, intermediate turbidity species (e.g.Valamugil cunnesius (Valenciennes) andLeiognathus equula (Forsskal)) in 10–80 NTU, turbid-water species (e.g.Elops machnata (Forsskal) andThryssa vitrirostris) in > 50 NTU, and species indifferent to turbidity (e.g.Acanthopagrus berda (Forsskal) andTeraponjarbua (Forsskal). It is, therefore, suggested that turbidity plays a significant roˆle, either singly, or in combination with other variables in determining the distribution of juvenile marine fishes in estuaries.     

Biological effects of salinity gradient reversals in a southeast African estuarine lake

St Lucia Lake on the north coast of Natal, South Africa, has an area of 325 km² and is the largest estuarine complex in Africa. It consists of a 20 km tidal channel, averagingca. 400 m in width, linking the sea with the non-tidal lake which is H-shaped with a maximum length ofca. 40 km and width ofca. 20 km. Except during flood periods the depth of the lake does not exceed 2 m. The salinity gradient depends on evaporation, the configuration of the mouth and on the input of fresh water from four rivers which discharge into the northern and western areas of the lake. If fresh water input is high, the lake and much of the channel may be fresh. An intermediate stage features a normal salinity gradient while a third stage shows a reversed salinity gradient with salinities in excess of 100‰ in the upper reaches of the system. Changing salinities have marked effects on the biota. Aquatic macrophytes show cycles of appearance and disappearance depending on salinity tolerance and the presence of dormant stages. The resident benthic faunal species go through cycles of range expansion and contraction depending on prevailing salinities and recolonisation by dispersal phases. To date salinities in the southern part of the lake have approached, but not exceeded, lethal levels and this has therefore acted as a reservoir area. Catchment degradation and water abstraction are anticipated to exacerbate future salinity extremes. This has resulted in concern for the long term viability of this Ramsar site which has major southern African populations of hippopotamus and crocodile, provides breeding sites for South African Red Data water bird species and plays an important nursery role for marine fish and penaeid prawns.     

Prawn community structure in the subtropical Mfolozi–Msunduzi estuarine system,KwaZulu-Natal,South Africa

The life cycles of many marine and freshwater prawn species are closely associated with estuarine habitat. Here, the prawn community of the Mfolozi–Msunduzi estuarine system is described and the system's potential as an alternative nursery for prawns during prolonged closure of the adjacent St Lucia estuarine system is examined. Sampling was conducted at five localities in summer and winter in 2007–2012. The Mfolozi–Msunduzi Estuary is a river-dominated system with mouth condition, sediment grain size and turbidity being the most important factors affecting the structure of its prawn community. Twelve prawn species were recorded, including five marine penaeid and five freshwater palaemonid species. The prawn community was numerically dominated by freshwater Macrobrachium equidens and two penaeids, Fenneropenaeus indicus and Metapenaeus monoceros. Significant differences were recorded between summer and winter samples, with freshwater species dominating the catch in summer, while penaeid species were dominant in winter. The marked seasonal change in the prawn community was related to strong river flows during summer creating low-salinity conditions, unsuitable for penaeid postlarval development, throughout most of the system. Compared to St Lucia, the Mfolozi–Msunduzi Estuary can be regarded as a poor alternative nursery area for penaeids during the summer peak postlarval recruitment period.     

2. NOTES ON THE STATUS OF SOME PALAEARC-TIC MIGRANTS IN NORTHERN RHODESIA

Whitfield, A. K. &; Cyrus, D. P. 1978. Feeding succession and zonation of aquatic birds at False Bay, Lake St Lucia. Ostrich 49:8-15.

Seasonal fluctuations in water levels at Lake St Lucia resulted in the inundation of depressions adjacent to the lake. Backwaters formed and were immediately colonized by aquatic invertebrates and fishes. Piscivorous birds dominated the early stages of the backwater feeding aggregations but with the decrease in water levels due to evaporation, crustacean and mollusc feeding birds became abundant. Both piscivorous and invertebrate-feeding wading birds formed a series of overlapping zones parallel to the shoreline.     

Genetics and shell morphometrics of assimineids (Mollusca,Caenogastropoda, Truncatelloidea) in the St Lucia Estuary,South Africa

The Assimineidae are a family of amphibious microgastropods that can be mostly found in estuaries and mangroves in South Africa. These snails often occur in great numbers and are ecologically important to the St Lucia Estuary, which forms a crucial part of the iSimangaliso Wetland Park, a UNESCO World Heritage Site. Genetic and shell morphometric analyses were conducted on individuals collected from nine localities distributed from the northern lake regions to the southern lake and the mouth of the St Lucia estuarine lake. Mitochondrial (COI) and nuclear (28S) DNA was used to construct Bayesian Inference, Neighbour-joining, Maximum Parsimony and Maximum Likelihood trees. Principal Component Analysis and Cluster Analysis were performed on standard shell parameter data. Results indicate that two different taxa are present in St Lucia. The taxon comprising individuals from the South Lake and St Lucia Estuary Mouth is identified as Assiminea cf. capensis Bartsch, in accordance with the latest taxonomic consensus. The taxon comprising assimineid individuals from False Bay, North Lake and South Lake, is here tentatively named “Assiminea” aff. capensis (Sowerby). These two taxa exhibit patterns of spatial overlap that appear to vary depending on environmental parameters, particularly salinity. The need to resolve the complex taxonomy of assimineids is highlighted.     

Persistent phytoplankton bloom in Lake St. Lucia (iSimangaliso Wetland Park, South Africa) caused by a cyanobacterium closely associated with the genus Cyanothece (Synechococcaceae, Chroococcales)

Lake St. Lucia, iSimangaliso Wetland Park, South Africa, is the largest estuarine lake in Africa. Extensive use and manipulation of the rivers flowing into it have reduced freshwater inflow, and the lake has also been subject to a drought of 10 years. For much of this time, the estuary has been closed to the Indian Ocean, and salinities have progressively risen throughout the system, impacting the biotic components of the ecosystem, reducing zooplankton and macrobenthic biomass and diversity in particular. In June 2009, a bloom of a red/orange planktonic microorganism was noted throughout the upper reaches of Lake St. Lucia. The bloom persisted for at least 18 months, making it the longest such bloom on record. The causative organism was characterized by light and electron microscopy and by 16S rRNA sequencing and was shown to be a large, unicellular cyanobacterium most strongly associated with the genus Cyanothece. The extent and persistence of the bloom appears to be unique to Lake St. Lucia, and it is suggested that the organism's resistance to high temperatures, to intense insolation, and to hypersalinity as well as the absence of grazing pressure by salinity-sensitive zooplankton all contributed to its persistence as a bloom organism until a freshwater influx, due to exceptionally heavy summer rains in 2011, reduced the salinity for a sufficient length of time to produce a crash in the cyanobacterium population as a complex, low-salinity biota redeveloped.     

THE LAKE ST. LUCIA SYSTEM—A RESEARCH ASSESSMENT

SUMMARY

This paper assesses the research undertaken at Lake St. Lucia over the past 25 years based on over 300 documents from that period. Trends related to both time and subject matter are evident, and these are considered in relation to the gaps in our current knowledge concerning the system.

A feature evident throughout the period under consideration is that the major portion of documented material available relates to reports and contributions to workshops (77%) with only 23% from scientific publications. Contributions by these two sources to the subject group being considered for St. Lucia are markedly different with research publications dominating the biological field and reports dominating in physical aspects, catchment characteristics, man's activities, management, dredging and hydrological modelling. However, some 55% of all unpublished data related to reviews or assessments of the state of research on St. Lucia.

The importance of the scientific publications group as an indicator of the state of research into the system is considered in the light of an apparent decline in the number of completed projects being published. It is also considered in the light of the recent establishment of a co-ordinated Lake St. Lucia Research Programme, which may provide the impetus for a more concentrated and directed research effort on the Lake System.     

Biodiversity census of Lake St Lucia,iSimangaliso Wetland Park (South Africa): Gastropod molluscs

The recent dry phase experienced by the St Lucia estuarine system has led to unprecedented desiccation and hypersaline conditions through most of its surface area. This has changed only recently, at the end of 2011, with the onset of a new wet phase that has already caused a major shift to oligo- and mesohaline conditions. The estuary mouth, however, remains closed to the ocean, making the weak connection recently established between the St Lucia and the Mfolozi estuaries the only conveyance for marine recruitment. As a result, only 10 indigenous and two alien aquatic gastropod species are currently found living in the St Lucia estuarine lake. This is out of a total of 37 species recorded within the system since the earliest survey undertaken in 1924, half of which have not been reported in the literature before. The tick shell, Nassarius kraussianus, which was consistently found in large abundance prior to the recent dry phase, appears to have temporarily disappeared from the system, probably as a result of the extinction of Zostera marine grasses inside the lake. Population explosions of the bubble shell Haminoea natalensis, with its distinct egg masses, were recorded seasonally until 2009, but the species has subsequently not been observed again. A molecular DNA analysis of the various populations previously reported as belonging to the same assimineid species, variably referred to as Assiminea capensis, A. ovata, or A. bifasciata, has revealed that the St Lucia assemblage actually comprises two very distinct taxa, A. cf. capensis and a species provisionally referred to here as “A.” aff. capensis or simply Assimineidae sp. In the mangroves, the climbing whelk Cerithidea decollata is still found in numbers, while ellobiids such as Cassidula labrella, Melampus semiaratus and M. parvulus are present in low abundances and all previously recorded littorinids have disappeared. A number of alien freshwater species have colonized areas of the system that have remained under low salinity. These include the invasive thiarid Tarebia granifera, which can be found in concentrations exceeding 5000 ind.m^-2, the lymnaeid Pseudosuccinea columella and the physid Aplexa marmorata.     

DISTRIBUTION,POPULATION SIZE AND CONSERVATION OF THE CASPIAN TERN STERNA CASPIA IN SOUTHERN AFRICA

Cooper, J., Brooke, R.K., Cyrus, D.P., Martin, A.P., Taylor, R.H. & Williams, A.J. 1992. Distribution, population size and conservation of the Caspian Tern Sterna caspia in southern Africa. Ostrich 63: 58–67.

The Caspian Tern Sterna caspia occurs along the whole southern African coastline and on large river systems and water bodies away from the coast. A total of 28 definite breeding localities has been recorded in southern Africa. Breeding has occurred recently at at least 14 coastal localities between Swakopmund, Namibia, and Lake St Lucia, Natal, South Africa. Inland breeding has been recorded in recent years at Sua Pan, Botswana and Kalkfonteindam, Orange Free State, South Africa. Based on censuses conducted between 1980 and 1991, the southern African breeding population is estimated to be of the order of 500 pairs, 91% of which breed coastally and 89% breed on islands. Up to 290 pairs (58%) bred at Lake St Lucia. Seventyone per cent of the 1980–1991 breeding population falls within eight nature reserves. Conservation of the Caspian Tern in southern Africa requires protection at breeding localities, including commercial salt and soda ash extraction works, against the effects of changing water levels, human disturbance and predation. Pesticide levels of addled and abandoned eggs should be measured at selected breeding localities.     

PRELIMINARY OBSERVATIONS ON THE MACROCRUSTACEA OF COASTAL LAKES IN THE VICINITY OF RICHARDS BAY,ZULULAND, SOUTH AFRICA

SUMMARY

A limited amount of research has been carried out on the four coastal lakes in the Richards Bay area. Due to the impending threat of water pollution from industrial, mining and residential areas within the catchments of these lakes there is an urgent need for the collection of baseline data before it is too late.

This paper reports on preliminary observations on Macrocrustacea of the lakes. A total of 27 species were recorded, of which 8 are new distribution records for the species in fresh water and one, an isopod, is a species new to science from a family previously unrecorded on the African continent. Sixteen of the Macrocrustacea recorded were found to be relict estuarine species.

The relationship between the Macrocrustacea are discussed in terms of habitat and dietary associations. The former category was found to be variable in the shallower Richards Bay lakes as opposed to Lake Sibaya whose depth (40 m) plays an important role in species distribution. Information is provided on habitat, substrata preference, associations with plant communities, and known depth distribution. Dietary data on the Macrocrustacea present in the four coastal lakes showed that seven major feeding groups are present: benthic filter feeders, nektonic filter feeders, periphyton grazers, shredders of dead macrophytes, benthic scavengers, partial carnivores and fish ectoparasites.     

The influence of zooplankton food resources on the morphology of the estuarine clupeid Gilchristella aestuarius (Gilchrist, 1914)

Synopsis The body shape of the estuarine clupeidGilchristella aestuarius from the St. Lucia system is different from that of the same species in other estuaries. The morphology ofG. aestuarius is discussed in relation to long term food availability in the St. Lucia system.     

Ecological studies on sporobolus virginicus (L.) kunth with particular reference to salinity and inundation

Sporobolus virginicus is shown to occupy low-lying areas on the shores of Lake St. Lucia, where it is subjected to periodic inundation and widely varying salinities. Whilst seeds are not permanently affected by storage under waterlogged saline conditions, young plants are markedly influenced by both inundation and salinity. Older plants are more tolerant of these conditions.     

Penaeid and carid community changes in the St Lucia estuarine lake system,South Africa,under low water level,extended closed periods and marine reconnection conditions

Penaeid and carid communities were assessed in St Lucia over a wide range of saline conditions, water level conditions and mouth states, including a six-month marine connection and various connections to the Mfolozi Estuary. Samples were collected biannually in spring and autumn from November 2004 to May 2012 using seine nets. Seven penaeid and seven carid species were recorded, of which Palaemon pacificus, P. peringueyi, Penaeus indicus and Metapenaeus monoceros were the most abundant. The decline from 2004 to 2007 in both species recorded and densities of freshwater, estuarine and marine species during the initial closed period was linked to increased salinity, lack of recruitment and reduced estuarine surface area. Increased wave action and high seas following Cyclone Gamede opened the mouth in March 2007 after a 57-month closed period, resulting in recruitment of marine species, predominantly P. indicus, followed by a decline in densities after reclosure in August 2007. Increases in marine and freshwater species were evident after Mfolozi flood connections during 2008, 2009 and 2010. Highest densities were recorded after breaching in 2007 and after flood connections, highlighting the importance of a marine link to maintain the recruitment of penaeid species.     

Argulus izintwala n. sp. (Crustacea: Branchiura) from Lake St Lucia,South Africa

During surveys of the fish parasites of Lake St Lucia on the coast of northern KwaZulu-Natal, specimens of an hitherto undescribed argulid were found on the skin of the kelee shad Hilsa kelee (Cuvier). This, only the fourth argulid species to be found in South African coastal waters, is here described as Argulus izintwala n. sp.     

BREEDING NOTES ON THE BLACK-NECKED GREBE PODICEPS NIGRICOLLIS BREHM

Summary

Tarboton, W. R. &; Fry, C. H. 1986. Breeding and other behaviour of the Lesser Jacana. Ostrich 57: 233–243.

Breeding Lesser Jacanas were studied briefly at Lake St Lucia (Zululand), Hwange (Zimbabwe) and the Okavango Delta (Botswana). The species is monogamous and breeding birds are dispersed as territorial pairs. Male and female share incubation nearly equally, alternating at the nest in shifts averaging 39min; the eggs are attended (incubated or shaded), on average, for 82% of the daylight hours. Eggs are incubated by holding them against the breast with the underside of the wings; at least one chick was seen carried under a parent's wing. The pullus, foraging behaviour, courtship and vocalisations are described. It is suggested that the Lesser Jacana's small egg necessitates a high rate of nest attendance which could account for the sociosexual differences between this species and other jacanas.     

ON THE RACE OF CARDINAL WOODPECKER Dendropicos fuscescens (Vieillot) OCCURRING IN THE EASTERN TRANSVAAL

Summary

Hustler, K. &; Marshall, B.E. 1996. The abundance and food consumption of piscivorous birds on Lake Kariba, Zimbabwe-Zambia. Ostrich 67:23-32.

About 40 species of piscivorous birds have been recorded on Lake Kariba (Zimbabwe-Zambia) but only 26 were observed during this study, carried out in 1986–87. The Reed Cormorant Phalacrocorax africanus was the most numerous and made up 51.4% of all the birds seen, followed by the Whitewinged Black Tern Chlidonias leucoptera (26.7%), and the Darter Anhinga melanogaster (10.8%). Several species roosted or bred in the drowned trees which project above the lake surface but gently sloping shores were the most important feeding grounds. Relatively few birds were seen on steeply sloping shores or in open water. There was a distinct pattern of seasonal abundance, with birds being most numerous in the rainy season (November-April). The relative abundance of some fish species also was greatest during this period. There was a positive relationship between the numbers of some bird species and the relative abundance of these fish. The mean density of piscivores (0.05 ha^?1) was low compared to some other African waters. This is attributed to the fact that Lake Kariba is relatively deep and its shoreline is mostly steep. The mean population of 25 000 birds consumed less than 1000 t of fish per annum, compared to almost 40000 t taken by the commercial fisheries. A considerable portion of the birds' diet was fish that were not taken commercially.     

Tidal behaviour of post-larval penaeid prawns (Crustacea: Decapoda: Penaeidae) in a southeast African estuary

Although the post-larvae of many species of inshore penaeid prawns migrate from the sea into estuarine habitats the mechanisms by which this process occurs have not been adequately explained. Collections of penaeid post-larvae in the St. Lucia estuary in Natal, South Africa during flood and ebb tides, day and night, were dominated by Penaeus japonicus Bate and P. indicus Milne Edwards. P. indicus was most abundant over flood tides, day and night but P. japonicus was markedly more nocturnal and only abundant over night floods. Vertical distribution differed in the two species. P. japonicus was more abundant in bottom samples but this was much less apparent in P. indicus. It is suggested that movement into the water column is triggered by pressure changes and this is modified by light, salinity, and the nature of the substratum. These responses are discussed in relation to the invasion of estuaries by penaeid post-larvae and the apparent survival of P. indicus but not P. japonicus in the St. Lucia system.