Genetic and morphological studies of Nothobranchius (Cyprinodontiformes) from Malawi with description of Nothobranchius wattersi sp. nov.

Molecular and morphological data were used to explore evolutionary differentiation among populations of Nothobranchius in the Lake Malawi–upper Shire River and the Lakes Chilwa–Chiuta drainage systems in Malawi. The aim of the study was to test the hypothesis that Nothobranchius of the Malawi–Shire system constitute a separate evolutionary group from Nothobranchius kirki. Mitochondrial and nuclear sequence data show a strongly supported phylogenetic split into two monophyletic groups separating the Lake Malawi basin fish from N. kirki. Unlike N. kirki, Lake Malawi–Shire fish do not deviate from neutrality and express an excess of rare haplotypes and mutations in terminal branches, characteristic of recently expanded populations. Further, the two groups significantly differ in morphology. Two body characters (dorsal‐fin base length and pre‐pelvic–pre‐anal distance) are significantly different between the two species in both sexes. Several other characters are significantly different in either male or female comparisons with respect to both standard and head lengths, and robust morphological differentiation is detected by multivariate analysis. The two groups are readily distinguished on the basis of male colouration, especially in scale centres and the caudal fin. On the basis of this differentiation at the molecular and morphological levels, in addition to colouration, the Lake Malawi–Shire fish are hereby formally recognized as constituting a new species, Nothobranchius wattersi. This distinction is in agreement with the geomorphologic and recent climatic history in the region.     

Development of polymorphic microsatellite DNA loci for characterizing Oreochromis shiranus subspecies in Malawi

Six sets of microsatellite primers were developed from Oreochromis shiranus nuclear DNA. These cross‐primed in species of the genera Sarotherodon and Tilapia, and were polymorphic in most of the species. The total number of alleles ranged from 18 to 30 per locus; the mean heterozygosity per population ranged from 0.51 ± 0.12 to 0.82 ± 0.03, which is higher than reported in allozyme studies. Five sets of primers were used to characterize five putative populations of O. shiranus, an indigenous mouth brooding tilapia which has been widely distributed in fish farms and reservoirs in Malawi. Two postulated subspecies of O. shiranus (subspecies shiranus and subspecies chilwae) have been difficult to distinguish morphologically. Lake Chilwa, Lake Chiuta and Bunda Reservoir populations cluster together as O. sh. chilwae, while the Lake Malombe and Bishop Reservoir populations form a second cluster as O. sh. shiranus. The assignment of the Chiuta and Chilwa populations of O. sh. chilwae to a single subspecies is consistent with the fact that the two lakes were a single open lake until about 8000–9000 _bp. There is no connection between them and the Lake Malawi‐Malombe drainage system where the O. sh. shiranus subspecies is found.     

THE SCIENCE: LIMNOLOGY

SUMMARY

Lake Chilwa, which has a fishery of economic importance to Malawi, declined in water level from 1964–68 and dried up completely for a few months in 1968. It then regained its former depth of 2–3 m in one wet season of five months. In the pre-drying period, zoo-plankters produced different diapause forms; in the drying period, zooplankton appeared to die out; in the refilling period rotifers predominated in the first year and crustacean populations were low; in the post-filling period crustaceans predominated. The same species occurred both before and after dryness. The conditions that possibly determined these responses are considered. After a lag of one year numbers of zooplankters were high, but decreased annually when the three dominant, largely planktivorous species of fishes recolonised the lake in succession over three years.

A main detritus food web is postulated and evidence for it is discussed. A practical method for fish conservation in times of low lake level, using artificial ‘lagoons’ in the swamp, and boreholes, is suggested.     

The role of the littoral zone in the functioning of a shallow tropical lake ecosystem

Lake Chilwa (Malawi) is a shallow tropical lake surrounded by an extensive zone of littoral swamp mainly composed of Typha domingensis. The data produced show that the physical and chemical conditions are different and more varied in the littoral region when compared with the open lake. Primary production in the littoral is confined almost entirely to Typha domingensis, which also acts as a nutrient pump by absorbing nutrients from the soils and eventually releasing them into the lake waters. Long term and short term exchanges in organic and inorganic materials between the littoral swamps and the open lake are described. These, as well as available evidence on the distribution, reproductive habits and diet of certain faunal groups, indicate that the littoral region of Lake Chilwa has an important regulating influence on the functioning of the whole lake.     

Zooplankton in a quasi-stable phase in an endorheic lake (Lake Chilwa,Malawi)

Lake Chilwa, a fairly large, turbid, brackish and very shallow endorheic lake in Malawi, Central Africa, dried up completely in 1968 and filled up again in the following wet season. Compared with the zooplankton in the drying, filling and post-filling phases (1966–1971), differences in their composition, density, distribution and reproduction were found in 1975–1976, a year when the lake was 1–2 metres deeper. This situation is discussed in relation to environmental factors, fish predation and the supply of detritus from the surrounding Typha swamp, to illustrate the relative stability of the zooplankton populations in periods between lake recessions.University of Malawi     

Vegetation changes in a shallow African Lake: Response of the vegetation to a recent dry period

This paper shows how the bed of Lake Chilwa in Malawi was colonised by plants during a recent dry period. Aeschynomene pfundii and Diplachne fusca were the principal colonising species on the alkaline mud flats. The subsequent changes in the vegetation after the lake refilled are described. Aeschynomene pfundii survived the flooded conditions for four years and eventually died out as seed germination could not occur underwater. The salinity of the lake during the low water and drying period prevented the lake bed from being extensively colonised by Typha domingensis.     

Abundance and distribution of fish in the lower Mnembo River, Malawi-Mozambique

Endorheic Lake Chilwa is one of the most productive lakes for fisheries in Africa. High human population densities and agricultural practices in the Chilwa catchment have been linked to observed declines of the annual fish catch in the lake. The Mnembo River is a major inflow into Lake Chilwa which has received little scientific study to date. In 2003/2004, fish diversity, abundance and distribution were monitored monthly at three sites in the Mnembo River. Twelve fish species were collected, a subset of the fourteen fish species present in Lake Chilwa. Although Barbus spp. were the most abundant species sampled in the Mnembo, total biomass was higher for Labeo cylindricus (6709 and 10,434 g, respectively). Total biomass of Pareutropius longifilis (1741 g) and Brycinus imberi (1174 g) were also high in the river. Catches of Barbus and Labeo were highly correlated (r = 0.763). Compared with other inflows into Lake Chilwa (Likangala and Domasi), the Mnembo River appears to be less adversely influenced by human populations, with high abundance of Barbus and strong presence of the riverine species Labeo, Pareutropius and Brycinus.     

The effects of suspensoids on fish

High suspensoid loads are a common feature of many Southern Hemisphere inland waters. Case studies on a natural lake (Chilwa), a man-made lake (Le Roux) and S.W. Indian Ocean estuaries reveal that the effects of turbidity on fish in these systems differ widely. In Lake Le Roux, high suspensoid loads influence fish by causing a reduced growth rate, a decrease in size at first maturity and maximum size, and a movement inshore by large fishes to feed on phytobenthos. High turbidities in Lake Chilwa sharply reduce food availability in benthic offshore zones, and restrict fishes to pelagic and inshore food resources. The resuspension of sediments by wind action may cause fish mortalities through deoxygenation of the water column. Moderate turbidity levels appear to be beneficial to fish in estuaries by affording protection from predators in shallow, food-rich areas. Turbidity gradients may also provide a navigational aid to fish entering estuaries.While ecological studies on Southern Hemisphere inland waters are fairly advanced, experimental work is restricted. Such research in the Northern Hemisphere has revealed that high suspensoid loads may influence breeding success, egg and larval survival, population structure and size, as well as food availability and feeding efficiency. The effect of suspensoids on the breeding success and feeding efficiency of Southern Hemisphere fish needs further investigation.Moderate suspensoid loads are a natural feature of many inland waters, but sustained high levels reduce the photic zone, blanket the benthos and interfere with the feeding efficiency of fish. The curtailment of soil erosion should therefore be regarded as a priority in the management of inland fisheries, especially in Third World countries where poor land-use practises have resulted in a marked deterioration of water quality. The need to study and manage river catchments holistically is emphasized.     

Geographical ancestry of Lake Malawi's cichlid fish diversity

The Lake Malawi haplochromine cichlid flock is one of the largest vertebrate adaptive radiations. The geographical source of the radiation has been assumed to be rivers to the south and east of Lake Malawi, where extant representatives of the flock are now present. Here, we provide mitochondrial DNA evidence suggesting the sister taxon to the Lake Malawi radiation is within the Great Ruaha river in Tanzania, north of Lake Malawi. Estimates of the time of divergence between the Lake Malawi flock and this riverine sister taxon range from 2.13 to 6.76 Ma, prior to origins of the current radiation 1.20–4.06 Ma. These results are congruent with evaluations of 2–3.75 Ma fossil material that suggest past faunal connections between Lake Malawi and the Ruaha. We propose that ancestors of the Malawi radiation became isolated within the catchment during Pliocene rifting that formed both Lake Malawi and the Kipengere/Livingstone mountain range, before colonizing rivers to the south and east of the lake region and radiating within the lake basin. Identification of this sister taxon allows tests of whether standing genetic diversity has predisposed Lake Malawi cichlids to rapid speciation and adaptive radiation.     

Lake flies and the deep-water demersal fish community of Lake Malawi

Larvae and pupae of Chaoborus edulis may be the most important food resource of some deep-water demersal fish species in southern Lake Malawi. C. edulis feed on crustacean zooplankton, which in turn feed on phytoplankton. Hence, demersal fish production is supported directly by planktonic primary production, rather than through a detrital food chain.     

Prey preferences in captivity of the freshwater crab Potamonautes lirrangensis from Lake Malawi with special emphasis on molluscivory

Freshwater crabs play an important role for the diversification of shell morphologies in freshwater gastropods. For example, the radiation of the freshwater crab genus Platythelphusa in Lake Tanganyika is thought to have driven shell diversification of the lake’s snail fauna, promoting the evolution of thalassoid shells. No comparable thalassoid snails are known from Lake Malawi. Accordingly, it was hypothesized that the lake’s only freshwater crab, Potamonautes lirrangensis, is not a snail predator. We tested this hypothesis using feeding experiments with specimens caught in the southern part of Lake Malawi. Individual crabs were held in experimental containers offshore and were presented with various food items overnight, after which ingestion frequency was recorded. Potamonautes lirrangensis can be characterized as a scavenger that is opportunistically carnivorous. A preference for fish and snail flesh could be observed, indicating a bias toward carnivory. We observed occasional cracking of the shell in different snail species, with frequent ingestion of artificially crushed specimens, suggesting that crabs do attempt to feed on snails. However, the investigated Lake Malawi gastropods appear to be partly protected against crab predation through thick-walled and low-spired shells (especially Lanistes and Bellamya), obviating the evolution of thalassoid shells carrying rims, ridges, or spines.     

Foraging of rocky habitat cichlid fishes in Lake Malawi: coexistence through niche partitioning?

The haplochromine cichlid fish communities of the rocky habitats of Lake Malawi are highly diverse; however, many species live side by side with apparently very similar resource requirements. There is a long-standing debate concerning whether these species partition their resources on a finer scale than has been previously reported or if species that are truly ecologically indistinguishable can coexist. A field study of food resource use was conducted to determine whether coexisting species segregate their diet and foraging sites. Significant differences between species were found, yet considerable inter-specific resource use overlap was commonplace. The data indicate that these cichlid species coexist both with and without niche differentiation. We propose that alternatives to niche differentiation should be considered to explain how many species coexist in Lake Malawi cichlid communities. Received: 5 October 1998 / Accepted: 30 June 1999     

Schistosomiasis transmission in Lake Malawi

Schistosomiasis (bilharziasis) transmission due to Schistosoma haematobium has for many years been known to occur along protected shorelines in Lake Malawi, but the recent finding that transmission also can also occur along open shorelines with sandy sediment has had a detrimental effect on tourism to Lake Malawi. The present paper shows that transmission, as evidenced from presence of intermediate hosts, along open shorelines is occurring in the southern part of the Lake. Relatively simple precautions can be taken to ensure minimal risk of attracting schistosome infection, and the schistosomiasis risk should not deter people from visiting and diving in Lake Malawi.     

Feeding-site utilization in three sympatric species of Petrotilapia (Pisces, Cichlidae) from Lake Malawi

Three sympatric sibling species of Petrotilapia , with similar dietary and macrohabitat requirements, occur near Monkey Bay, Lake Malawi. Males and females of all three species fed selectively on the rocky shores. Males utilized less than 6% of the space they defended as a breeding territory for feeding and although interspecific overlap of territories was considerable, only 2.5% of the feeding space used by territorial males was shared. Males primarily utilized rich food patches which occurred within the territories of highly aggressive Pseudotropheus species, whereas females tended to feed in undefended areas. Although Petrotilapia species are not highly aggressive fishes, there was a well-defined social dominance hierarchy amongst them and it is suggested that feeding-site utilization is dependent on social rank. Inter-and intraspecific differences in feeding-site utilization probably facilitate species coexistence and thus contribute to the maintenance of the high species diversity which is a characteristic of the ichthyofauna of Lake Malawi.     

Evolution of a cichlid fish in a Lake Malawi satellite lake

Allopatric divergence in peripheral habitats may lead to rapid evolution of populations with novel phenotypes. In this study we provide the first evidence that isolation in peripheral habitats may have played a critical role in generation of Lake Malawi's cichlid fish diversity. We show that Lake Chilingali, a satellite lake 11.5 km from the shore of Lake Malawi, contains a breeding population of Rhamphochromis, a predatory genus previously thought to be restricted to Lake Malawi and permanently connected water bodies. The Lake Chilingali population is the smallest known Rhamphochromis, has a unique male nuptial colour pattern and has significant differentiation in mitochondrial DNA from Lake Malawi species. In laboratory mate choice trials with a candidate sister population from Lake Malawi, females showed a strong tendency to mate assortatively indicating that they are incipient biological species. These data support the hypothesis that isolation and reconnection of peripheral habitats due to lake level changes have contributed to the generation of cichlid diversity within African lakes. Such cycles of habitat isolation and reconnection may also have been important in evolutionary diversification of numerous other abundant and wide-ranging aquatic organisms, such as marine fishes and invertebrates.     

The birth of an endemic species flock: demographic history of the Bellamya group (Gastropoda,Viviparidae) in Lake Malawi

Changes in habitat stability may significantly shape evolutionary patterns and processes in ancient lakes. In the present study, we use a hierarchical combination of molecular phylogenetic and coalescent approaches to investigate the evolutionary history of the endemic species of the gastropod genus Bellamya in the African rift‐lake Malawi. By integrating our findings with reported palaeontological and palaeolimnological data, we demonstrate that all but one evolutionary lineage of the Pliocene Bellamya fauna in Lake Malawi became extinct. Coalescent analyses indicate that the modern radiation underwent both a sudden demographic and a spatial expansion after a genetic bottleneck. We argue that a reflooding of the lake after severe Pleistocene low stands offers a straightforward explanation for this pattern and may have triggered speciation processes in the modern endemic Bellamya radiation in Lake Malawi. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 130–143.     

Hybrid origin of a cichlid population in Lake Malawi: implications for genetic variation and species diversity

The importance of species recognition to taxonomic diversity among Lake Malawi cichlids has been frequently discussed. Hybridization - the apparent breakdown of species recognition - has been observed sporadically among cichlids and has been viewed as both a constructive and a destructive force with respect to species diversity. Here we provide genetic evidence of a natural hybrid cichlid population with a unique colour phenotype and elevated levels of genetic variation. We discuss the potential evolutionary consequences of interspecific hybridization in Lake Malawi cichlids and propose that the role of hybridization in generating both genetic variability and species diversity of Lake Malawi cichlids warrants further consideration.     

Zooplankton of Lake Kivu, East Africa, half a century after the Tanganyika sardine introduction

The introduction into Lake Kivu of the planktivorous fish Limnothrissamiodon at the end of the 1950s raised major concern about thefate of mesozooplankton, but few data were available to assessthe impact of predation. In this study, we followed zooplanktonvariations in Lake Kivu for 3.5 years. Present Lake Kivu mesozooplanktonis dominated by cyclopoid copepods (Thermocyclops consimilis,Mesocyclops aequatorialis and Tropocyclops confinis), but cladoceransand rotifers are also present. Each year, total crustacean abundancein the plankton increased to a distinct seasonal maximum followinga rise of phytoplankton production associated with a deep epilimneticmixing in the dry season (August–September). This dependenceon phytoplankton resource suggests that mesozooplankton dynamicsin Lake Kivu is mainly bottom-up controlled, contrary to expectationsfrom the food web structure. However, measurements of body sizeindicate that sardine predation affects the cladoceran Diaphanosomaexcisum, whereas the larger copepods may efficiently escapepredation by migration in the deep mixolimnion at daytime. Totalbiomass of mesozooplankton in Lake Kivu is lower than in lakesTanganyika and Malawi. This may be related to the disappearanceof a large grazer that existed before the sardine introduction,whereas the large lakes of the same region have a more complexpelagic food web, with piscivorous fish, and with calanoid copepods,which can more efficiently exploit phytoplankton production.     

Causes of mortality in the endemic Tilapia of Lake Chilwa (Malawi)

Summary During 1966 and 1967 extensive mortalities of Tilapia shirana chilwae took place in Lake Chilwa, Malawi. By collating field observations and physico-chemical data from the lake with experimental evidence from the laboratory, it has been possible to predict what the most likely causes of mortality of Tilapia might have been.High winds caused bottom erosion in October 1966, and the released mud deoxygenated the water. Total deoxygenation of the water results if the level of suspended matter is raised to 12,860 ppm. Tilapia are able to survive levels of oxygen as low as 0.6 mg/l, but become distressed below this figure.The median lethal alkalinity for T. s. chilwae in artificial lake water is 61.6 meq/l. This concentration was reached in the lake in December 1966, and probably caused further mortalities. T. s. chilwae is remarkably tolerant of high temperature, and this factor alone is unlikely to have caused natural mortalities. Elevated temperatures can cause decreased resistance to low oxygen and high ionic concentrations, however, and the dense blooms of Anaebena may have caused sufficient deoxygenation of the water at night to cause further mortalities. T. s. chilwae is more resistant to high alkalinity and high temperature than T. melanopleura. This undoubtedly reflects the long history of stress in the endemic Tilapia. No doubt the heightened resistance of T. s. chilwae has enabled it to successfully populate L. Chilwa, despite its characteristic instability.

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Zusammenfassung Betrachtliche Mortalitäten des Fisches Tilapia shirana chilwae erfolgte im Chilwa See (Malawi) in 1966 und 1967. Bei der Zusammenfügung von den Beobachtungen am See und den physischchemischen Daten des Sees mit experimentellen Daten aus dem Laboratorium, war es moglich die wahrscheinlichen Ursachen des Mortalitäts festzustellen.Starke Winde verursachten die Erosion des Bodens im Oktober 1966 und der aufgelöste Schlamm entfernte die Sauerstoffe des Wassers. Totale Entfernung des Sauerstoffes folgte als tas Niveau des aufgelösten Materials einen Wert von 12,800 t.p.m. erreichte. Obgleich Tilapia einen so niedrigen Sauerstoffniveau von 0.6 mg/l überleben können fühlen she sich unheimlich unter diesem Wert.Die mediane letale Alkalinitat für T. s. chilwae im künstlichen Seewasser ist 61.6 meq/l. Diese Konzentration wurde im Dezember 1966 im See erreicht und verursachte wahrscheinlich eine weitere Mortalität. T. s. chilwae ist auffällig tolerant hohen Temperaturen gegen-über und es ist unwahrscheinlich dasz these Tatsache allein natürliche Mortalitäten verursacht hat. Dennoch können erhdhte Temperaturen eine abnehmende Resistenz gegen niedrigen Sauerstoff und hohe Ionen-Konzentration verursachen und die dicke Bliite von Anaebena könnte eine genügende Desoxygenation des Wassers in der Nacht veranlasst haben um weitere Mortalitaten zu verursachen. T. s. chilwae zeigt stärkere Resistenz gegenüber hoher Alkalinität und hoher Temperaturen als T. melanopleura. Dieses reflektiert, ohne Zweifel die lange Geschichte von Stress in die endemische Tilapia. Die erhöhte Resistenz des T. s. chilwae, dasz es für these Fische möglich war den Chilwa See zu bevölkern, trotz ihrer charakteristischen Instabilität.

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University of Malawi     

A CONCEPTUAL MODEL OF A SILTATION SYSTEM IN SHALLOW LAKES WITH LITTORAL VEGETATION

SUMMARY

In shallow lakes a siltation system develops when wind driven water turbulence erodes the lake floor, silt is resuspended and transported by means of waterdrift into the littoral vegetation. Here, due to the resistance of the vegetation, the turbulence is reduced and the silt deposited. Through this mechanism, the prevailing winds have a major rôle in the erosion and siltation pattern, lake morphology, vegetation pattern and in the mixing of water and nutrients from the open water and the reedswamp and vice versa. The model explains the uneven distribution of vegetation at the lakes Neusiedlersee (Austria) and Chilwa (Malawi) satisfactorily, if one assumes a positive correlation between siltation and reedswamp growth and spread.