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.     

WATER TEMPERATURE AND THE 1987 FISH KILL AT LAKE ST LUCIA ON THE SOUTH EASTERN COAST OF AFRICA

Summary

An unusually cold spell during the winter of 1987 caused a drop in water temperature at Lake St Lucia. This coupled with typical estuarine salinities in the lake resulted in a fish kill. The kill was one of the largest recorded in a South African estuarine environment and involved an estimated 250,000 fish comprising at least 21 species. Most fish that died belonged to small species. Densities of dead fish recorded along the lake's shoreline ranged from 0.1 to 16.6 fish per metre. It is concluded that the combination of low water temperature and near marine salinities was the major cause of the 1987 fish kill at Lake St Lucia.     

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.     

Epiphytes of the St. Lucia Estuary and their response to water level and salinity changes during a severe drought

St. Lucia is the largest estuary in South Africa with extensive areas of submerged macrophytes that fluctuate rapidly in response to water level and salinity changes. Epiphytes associated with submerged macrophytes were sampled during a severe drought between November 2004 and October 2005 when very low water level and high, variable salinity characterised the estuary. Potamogeton pectinatus and Ruppia cirrhosa were the dominant submerged macrophytes throughout the estuary, with P. pectinatus occurring at relatively low salinity (∼10 ppt) and R. cirrhosa at higher salinity (9–33 ppt). Zostera capensis, normally the other dominant submerged macrophyte, was conspicuously absent under the prevailing conditions. Epiphytic biomass, estimated as chlorophyll a, varied greatly between sites and over the 12 month sampling period, ranging from 10.9 to 71.7 mg Chl a m⁻² leaf area for P. pectinatus and 16.9–165.0 mg Chl a m⁻² leaf area for R. cirrhosa. Epiphytic biomass was twice as high in the Southern Lake where R. cirrhosa occurred, probably because the dominant epiphytes were macroalgae. An assessment of the diatom species composition of the epiphytic community indicated the dominance of only six species throughout the estuary. Neither epiphytic biomass nor diatom species composition showed strong statistical relationships with the environmental variables measured and it is believed that biological factors may be more important than the physico-chemical environment in determining epiphyte biomass distribution. Because epiphyte biomass is dependent on the presence of host surface area it will only contribute substantially to overall system biomass and productivity when submerged macrophyte area cover is high. In the St. Lucia Estuary this will occur when the water level is high and the upper level of the salinity gradient does not increase above that of seawater.     

Neogosseidae (Gastrotricha,Chaetonotida) from the iSimangaliso Wetland Park,KwaZulu-Natal,South Africa

Among the mostly benthic gastrotrichs, the Neogosseidae (Gastrotricha, Chaetonotida) are particularly interesting from an evolutionary point of view in virtue of their planktonic lifestyle; yet, they are poorly known and uncertainties concerning morphological traits hamper accurate in-group systematics. During a recent survey of meiofauna in the iSimangaliso Wetland Park, South Africa, two species of Neogosseidae were found in a freshwater pond near Charter’s Creek on the Western Shores of Lake St Lucia. Based on morphological traits, one species has been identified as Neogossea acanthocolla, originally described from Brazil, while the other, affiliated to the genus Kijanebalola, is proposed as new to science. Using a combination of differential interference contrast and scanning electron microscopy, fine anatomical details were observed and are here discussed in a larger taxonomic framework, especially regarding Kijanebalola devestiva sp. n. Results have also provided reasons for a revision of the diagnostic traits of Kijanebalola, Neogossea and the whole Family Neogosseidae. Besides expanding awareness about the biodiversity hosted by South Africa’s first UNESCO World Heritage Site, our study will be beneficial to future phylogenetic studies of the Gastrotricha based on morphology, by allowing the selection and/or a more precise character coding of traits of phylogenetic relevance.     

Population structure of an invasive parthenogenetic gastropod in coastal lakes and estuaries of northern KwaZulu-Natal, South Africa

Background

Estuaries and coastal lakes receive little attention despite being heavily invaded by non-indigenous invasive species (NIS). In these situations, studies of population dynamics in invaded habitats can provide valuable insights into how NIS interact with new environments. Tarebia granifera is a prosobranch gastropod from south-east Asia which has invaded other sub-tropical parts of the world. This study addresses whether a small number of key environmental factors influences gastropod communities, and specifically how the population density and size structure of T. granifera were influenced by environmental change in estuaries and coastal lakes in southern Africa.

Methodology/Principal Findings

T. granifera''s density, number of brooded juveniles and size structure were measured at the St. Lucia Estuary, Mgobozeleni Estuary, Lake Sibaya and Lake Nhlange. Size structure was classified according to shell height (SH). All dissected individuals were found to be female and free from trematode infection. Salinity, water depth, temperature, and pH were the main factors correlated with population density of gastropod communities. T. granifera often reached densities well over 1000 ind. m⁻², displacing indigenous gastropods and becoming a dominant component of the benthic community. T. granifera successfully invaded estuaries despite frequent exposure to high salinity and desiccation, which could together eliminate >97% of the population. The persistence of T. granifera was ensured due to its high fecundity and the environmental tolerance of large adults (20–30 mm SH) which carried an average of 158±12.8 SD brooded juveniles. Repeat introductions were not essential for the success of this parthenogenetic NIS.

Conclusion/Significance

There is a need for a broader study on the reproductive biology of T. granifera (including the previously overlooked “brood pouch ecology”), which affects population dynamics and may be relevant to other parthenogenetic NIS, such as Melanoides tuberculata and Potamopyrgus antipodarum.     

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.     

In situ feeding rates and grazing impact of Mesopodopsis africana O. Tattersall in the St Lucia Estuary, South Africa

Mesopodopsis africana is an important mysid in southern African coastal zooplankton and a key species in the St Lucia estuarine lake, which is currently undergoing severe desiccation owing to freshwater deprivation. M. africana populations through much of the system are consequently under severe environmental stress. This study investigates the grazing dynamics of this mysid species, in relation to autotrophic food availability and other environmental constraints in two contrasting areas of the St Lucia Estuary, Charters Creek, heavily affected by the desiccation process and the Mouth, virtually under unchanged conditions. Gut evacuation experiments were conducted once each during the day and the night. Evacuation rates were consistently higher during the night, ranging from 0.27 to 0.33 h^-1 at Charters Creek and from 1.13 to 1.24 h^-1 at the Mouth. Ingestion rates were, therefore, higher at the Mouth resulting in population grazing impacts of 2.5% of the total microalgal biomass, while the grazing impact at Charters Creek was only 0.5%. The spatial variation in ingestion rates could be attributed to seasonal differences in gut evacuation rates, differences in the mean size of mysids used, or the physicochemical conditions present at the two stations. It is suggested that mysid populations at Charters Creek are predominantly driven by bottom-up forces, initiated by the harsh environmental conditions. Despite the lower ingestion rates exhibited at Charters Creek, results indicate that these mysids are capable of meeting all their energetic requirements from a microalgal diet alone, although they may also utilise a heterotrophic diet.     

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.     

A new genus for a rare African vespertilionid bat: insights from South Sudan

A new genus is proposed for the strikingly patterned African vespertilionid “Glauconycteris” superba Hayman, 1939 on the basis of cranial and external morphological comparisons. A review of the attributes of a newly collected specimen from South Sudan (a new country record) and other museum specimens of “Glauconycteris” superba suggests that “Glauconycteris” superba is markedly distinct ecomorphologically from other species classified in Glauconycteris and is likely the sister taxon to Glauconycteris sensu stricto. The recent capture of this rarely collected but widespread bat highlights the need for continued research in tropical sub-Saharan Africa and in particular, for more work in western South Sudan, which has received very little scientific attention. New country records for Glauconycteris cf. poensis (South Sudan) and Glauconycteris curryae (Gabon) are also reported.     

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.     

Fluctuations in numbers of great white pelicans at Lake St Lucia in response to changing water-levels

In January 2004, Lake St Lucia, a major part of the Greater St Lucia Wetland Park, was reduced to a fraction of its normal capacity as a result of a severe drought in this region of KwaZulu-Natal, South Africa. After rains in the area, the lake level rose and then fluctuated considerably over the next 42 months. During the first 38 months the mouth of the estuary into the sea was closed. The area entered a second severe period of prolonged drought from mid 2005 through to the spring of 2006. Great white pelican ( Pelecanus onocrotalus Linnaeus, 1758 ) numbers and lake levels were monitored during these 42 months. Pelican numbers were highly variable ranging from 0 to 6000. When lake levels were very low or too high, no pelicans were present. Pelican numbers appeared more indicative of food availability. Implications of these trends to the management of the lake and the conservation of the avifauna are discussed. It is suggested that the great white pelican could be used as an indicator species for the fish dynamics of the lake at medium to low lake levels.     

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.     

Impact of a flood event on the zooplankton of an estuarine lake

Shallow coastal lakes are prone to large fluctuations in physico-chemical variables such as salinity and turbidity. This is now escalating in response to global change. A flood event in March 2014 resulted in a silt plume spreading through part of Lake St Lucia (South Africa). To determine the impact of this event on zooplankton, the Narrows region of St Lucia was sampled on a monthly basis from March to September 2014. For comparative purposes, data from samples collected prior to the flood event were included in the analyses. Analysis of similarity (ANOSIM) revealed dissimilarities in zooplankton community structure among the sampling occasions. The March 2014–May 2014 period was characterized by the highest abundance of freshwater species. Conversely, the abundance of the resident St Lucia copepods Acartiella natalensis and Oithona brevicornis was lowest during this time, and highest in September 2014. The other dominant copepod Pseudodiaptomus stuhlmanni prevailed in March 2014, but declined markedly in April. As of September 2014, P. stuhlmanni had yet to regain its pre-flood densities. The BIOENV procedure, which relates biological and environmental data, revealed that turbidity, salinity and dissolved oxygen were responsible for the observed changes in zooplankton community structure during the study period. Careful management of turbidity and salinity is stressed, as both factors are major drivers of the biota of St Lucia and similar systems worldwide.     

Afroprinus cavicola gen. et sp. n. from the Afrotropical region with notes on cave-dwelling Saprininae (Coleoptera,Histeridae)

A new genus and species from Kenya, Afroprinus cavicola is herein described and illustrated and its systematic position is discussed. By the prosternal pre-apical foveae connected by marginal prosternal stria it resembles most of the Afrotropical species of the genus Chalcionellus Reichardt, 1932, or some species of the genus Pholioxenus Reichardt, 1932 from South Africa and Namibia. Afroprinus can be distinguished from Chalcionellus chiefly by the lack of pronotal depressions and a coarsely sculptured, non-metallic dorsum; from Afrotropical species of Pholioxenus it can be most easily distinguished by the asetose pronotal hypomeron. The new taxon was discovered in a cave, but lacks obvious troglophilic adaptations. Notes on other Saprininae taxa found in caves are given. An identification key to the genera of Afrotropical Saprininae is provided.     

Temperature and salinity tolerance of Mesopodopsis africana O. Tattersall in the freshwater-deprived St. Lucia Estuary, South Africa

Mesopodopsis africana is a key species in the St. Lucia Estuary, Africa's largest estuarine lake. This system is currently undergoing an unprecedented crisis due to freshwater deprivation. A reversed salinity gradient has persisted with hypersaline conditions (> 300) occurring in the upper regions of the estuarine lake. In the context of climate change, rising temperatures will not only push the thermal tolerance limits of estuarine organisms, but increased evaporation from this lake's large surface area will lead to further salinity increases. The present study aims to determine the temperature and salinity tolerance of M. africana, both through in situ studies and the use of laboratory experiments. Results indicate that M. africana is a broad euryhaline species. Mysids were recorded at salinity levels ranging from 2.55 to 64.5 in situ. While experiments revealed a narrower salinity tolerance, acclimation resulted in a significant increase in the tolerance range of this species. It is probable, however, that slower acclimation times may increase survival rates even further, particularly in the higher salinity treatments. M. africana was especially tolerant of the lower salinity levels. In the 20 °C acclimation experiment, LS₅₀ at 1 and 2.5 was only reached after 8 and > 168 h, respectively. Survival at 10 and 40 °C was negligible at all salinity levels. This concurs with field results which documented mysids at temperatures ranging from 16.2 to 30.9 °C. Salinity and temperature increases associated with global climate change may, therefore, have significant implications for these mysid populations, with cascading effects on the higher trophic levels which they support.     

Testing two contrasting evolutionary patterns in ancient lakes: species flock versus species scatter in valvatid gastropods of Lake Ohrid

Ancient lakes have long been recognized as “hot spots of evolution” and “evolutionary theatres” and they have significantly contributed to a better understanding of speciation and radiation processes in space and time. Yet, phylogenetic relationships of many ancient lake taxa, particularly invertebrate groups, are still unresolved. Also, the lack of robust morphological, anatomical, and phylogeographical data has largely prevented a rigorous testing of evolutionary hypotheses. For the freshwater gastropod genus Valvata—a group with a high degree of endemism in several ancient lakes—different evolutionary scenarios are suggested for different ancient lakes. Lake Baikal, for example, is inhabited by several endemic Valvata taxa that presumably do not form a monophyletic group. For such an evolutionary pattern, the term ancient lake species scatter is introduced here. In contrast, for the Balkan Lake Ohrid, workers previously suggested the presence of a monophyletic group of endemic Valvata species, that is, an ancient lake species flock. Sequence data of the mitochondrial cytochrome oxidase c subunit I gene (COI) from worldwide taxa, with a strong emphasis on Balkan species, are here used to test whether the putative Ohrid Valvata endemics represent an ancient lake species flock and to study patterns of speciation both on the Ohrid and the Balkan scale. The study reveals three distinct clades of endemic Valvata in Lake Ohrid. Monophyly of these taxa, however, is rejected, and they therefore do not represent an ancient lake species flock, but rather an ancient lake species scatter. Also, in contrast to many other gastropod groups in Lake Ohrid, the valvatids apparently did not radiate. Many Valvata taxa in ancient lakes are characterized by enhanced levels of shell complexity. However, it remains unclear whether these patterns are associated with ancient lake environments per se. It is here suggested that similarities in shell structure between North American and Balkan taxa might simply be due to convergent evolution.