Recent research in the African Great Lakes: fisheries, biodiversity and cichlid evolution

The East African Great Lakes are now well known for (1) their fisheries, of vital importance for their rapidly rising riparian human populations, and (2) as biodiversity hotspots with spectacular endemic faunas, of which the flocks of cichlid fishes unique to each of the three largest lakes, Tanganyika, Malawi and Victoria, offer unique opportunities to investigate how new species evolve and coexist. Since the early 1990s research involving over a hundred scientists, financed by many international bodies, has produced numerous reports and publications in widely scattered journals. This article summarizes their main discoveries and examines the status of, and prospects for, the fisheries, as well as current ideas on how their rich endemic fish faunas have evolved. It first considers fisheries projects in each of the three lakes: the deep rift valley lakes Tanganyika and Malawi and the huge Victoria, all of which share their waters between several East African countries. Secondly it considers the biodiversity surveys of each lake, based on underwater (SCUBA) observations of fish ecology and behaviour which have revealed threats to their fish faunas, and considers what conservation measures are needed. Thirdly, using the lakes as laboratories, what have the international investigations (including DNA techniques and follow-up aquarium experiments) now revealed about the origins and relationships of their cichlid species flocks and mechanisms of evolution?

Coral reef resources of East Africa, Kenya, Tanzania and the Seychelles

Coral reefs are widespread along the east African coast and Seychelle islands. Their roles in island building and coastal protection are often underestimated, they are also important fishery habitats and major tourist attractions. The east African marine fishery production, estimated at 1.4-4.9 tonnes per km super(2), is principally a result of artisanal fishing. Siltation, trampling, and destructive fishing methods are the main cause of coral reef degradation along the east African coast and associated islands. Legislation has been implemented to protect coral reefs by establishing marine parks and reserves. However, poaching and anchor damage are widespread on these protected reefs. Legislative provision to increase the benefit to fishing communities may reduce poaching. The establishment of exclusive nature reserves may be one way to ensure preservation of some coral reefs in the region.

A survey of research and scientific services in East Africa, 1947-56

This "Survey of Research and Scientific Services in East Arica 1947-1956" has been prepared by Dr. E. B. Worthington, who held the post of Scientific Secretary in the Office of the Chief Secretary to the East African Governor's Conference and subsequently in the Administrator's Office of the East Mrica High Commission during the period January, 1947-May, 1951. Dr. Worthington is now Secretary General of the Scientific Council for, Arica South of the Sahara

A review of the fisheries of the Uganda waters of Lake Albert East Africa, 1928-1965/66, with catch data mainly from 1953

An historical account is given of the development of the Lake Albert fisheries since Worthington's survey in 1928. It is noted that the development of the fisheries was related to, and dependent upon, improvements in the type of gear and canoes, an incFease in the number of canoes and outboard engines in use, improved marketing facilities and better road communications. Summarized data, collected mainly since 1954, has been analysed and tabulated to show annual exports to the Congo, total annual catches 'and annual catches of individual species. A change in the relative abundance of the various species in the annual catches is described. It is noted that this change was caused by a change-over from large to small mesh size gill-nets, and that it was associated with an increased demand within Uganda for the smaller species of fish, such as Aleste's baremose and Hydrocynus forskahlii. A brief description of fish processing and marketing in the Lake Albert region is given, which emphasizes the suitability of salt-cured fish to the social and physical environment of the area. Finally, a summary of a recent survey of the potential fish resources of the lake is given in the discussion, and estimates of the 1965 catch at the north and south ends of the lake are compared with the findings of the survey. This showed that there is little danger of overfishing the Alestes baremose stocks of the Wanseko area at the 1965 rate of exploitation of the species, and that the total catch for 1965 at the south end of the lake was well below the estimated annual sustainable yield from the area.

Improved facilities for marine biology in East Africa

In 1967 the then University College of Dar es Salaam built a small laboratory on the shore at Kunduchi, 16 km from the main campus and 24 km north of Dar es Salaam. This was used for undergraduate field courses, and as a base for staff from the University to carry out research. It soon became apparent that the urgent need for studies of the marine environment in the East African area, and the lack of existing facilities, necessitated the development of the Kunduchi Marine Biology station into a research establishment with its own staff of full time scientists. This operation began in 1970: necessary structural modifications have been made to the building, staff have been recruited, and the station has been equipped with an adequate range of field and laboratory apparatus. A varied programme of research is now actively under way.

International Partnering strategy for e-learning provider in emerging economies: Evidence from East Africa learning institutions

International partnership has received growing interest in the literature during the past decades due to globalization, increased technological approaches and rapid changes in competitive environments. The study specifically determines the support provided by international partners on promotion of e-learning in East Africa, assess the motives of partner selection criteria, the determinants of selecting partners, partner models and partner competence of e-learning provider. The study also evaluates obstacles of e-learning partnering strategy in East Africa learning institutions. The research adopts a descriptive survey design. Target population involved East Africa learning institutions with a list of potential institutions generated from the Ministry of Higher Education database. Through a targeted reduction of the initial database, consisting of all learning institutions, both public and private, the study created a target sample base of 200 learning institutions. Structured questionnaires scheduled were used to collect primary data. Study findings showed the approach way East African communities in selecting their e-learning partners depend on international reputation of partners, partner with ability to negotiate with foreign governments, partner with international and local experiences, nationality of foreign partner and partners with local market knowledge.

The competitiveness of domestic rice production in East Africa: A domestic resource cost approach in Uganda

The rapid increase of rice imports in sub-Saharan Africa under the unstable situation in the world rice market during the 2000s has made it an important policy target for the countries in the region to increase self-sufficiency in rice in order to enhance food security. Whether domestic rice production can be competitive with imported rice is a serious question in East African countries that lie close, just across the Arabian Sea, to major rice exporting countries in South Asia. This study investigates the international competitiveness of domestic rice production in Uganda in terms of the domestic resource cost ratio. The results show that rainfed rice cultivation, which accounts for 95% of domestic rice production, does not have a comparative advantage with respect to rice imported from Pakistan, the largest supplier of imported rice to Uganda. However, the degree of non-competitiveness is not serious, and a high possibility exists for Uganda’s rainfed rice cultivation to become internationally competitive by improving yield levels by applying more modern inputs and enhancing labour productivity. Irrigated rice cultivation, though very limited in area, is competitive even under the present input-output structure when the cost of irrigation infrastructure is treated as a sunk cost. If the cost of installing irrigation infrastructure and its operation and maintenance is taken into account, the types of irrigation development that are economically feasible are not large-scale irrigation projects, but are small- and microscale projects for lowland rice cultivation and rain-water harvesting for upland rice cultivation.

The Accumulation of Reproductive Incompatibilities in African Cichlid Fish

The rate at which different components of reproductive isolation accumulate with divergence time between species has only been studied in a limited, but growing, number of species. We measured premating isolation and hybrid inviability at four different ontogenetic stages from zygotes to adults in interspecific hybrids of 26 pairs of African cichlid species, spanning the entire East African haplochromine radiation. We then used multiple relaxed molecular clock calibrations to translate genetic distances into absolute ages to compare evolutionary rates of different components of reproductive isolation. We find that premating isolation accumulates fast initially but then changes little with increasing genetic distance between species. In contrast, postmating isolation between closely related species is negligible but then accumulates rapidly, resulting in complete hybrid inviability after 4.4/8.5/18.4 million years (my). Thus, the rate at which complete intrinsic incompatibilities arise in this system is orders of magnitude lower than rates of speciation within individual lake radiations. Together these results suggest divergent ecological adaptations may prevent populations from interbreeding and help maintain cichlid species diversity, which may be vulnerable to environmental degradation. By quantifying the capacity to produce viable hybrids between allopatric, distantly related lineages our results also provide an upper divergence time limit for the "hybrid swarm origin" model of adaptive radiation.

Origins of Shared Genetic Variation in African Cichlids

Cichlid fishes have evolved tremendous morphological and behavioral diversity in the waters of East Africa. Within each of the Great Lakes Tanganyika, Malawi, and Victoria, the phenomena of hybridization and retention of ancestral polymorphism explain allele sharing across species. Here, we explore the sharing of single nucleotide polymorphisms (SNPs) between the major East African cichlid assemblages. A set of approximately 200 genic and nongenic SNPs was ascertained in five Lake Malawi species and genotyped in a diverse collection of 160 species from across Africa. We observed segregating polymorphism outside of the Malawi lineage for more than 50% of these loci; this holds similarly for genic versus nongenic SNPs, as well as for SNPs at putative CpG versus non-CpG sites. Bayesian and principal component analyses of genetic structure in the data demonstrate that the Lake Malawi endemic flock is not monophyletic and that river species have likely contributed significantly to Malawi genomes. Coalescent simulations support the hypothesis that river cichlids have transported polymorphism between lake assemblages. We observed strong genetic differentiation between Malawi lineages for approximately 8% of loci, with contributions from both genic and nongenic SNPs. Notably, more than half of these outlier loci between Malawi groups are polymorphic outside of the lake. Cichlid fishes have evolved diversity in Lake Malawi as new mutations combined with standing genetic variation shared across East Africa.

How does the taxonomic status of allopatric populations influence species richness within African cichlid fish assemblages?

Aim  Current estimates of species richness within rapidly evolving species flocks are often highly dependent on the species status of allopatric populations that differ in phenotypic traits. These traits may be unreliable indicators of biological species status and systematists may have inconsistently assigned species among lineages or locations on the basis of these traits, thus hampering comparative studies of regional species richness and speciation rates. Our aim was to develop a method of generating standardized estimates of regional species richness suitable for comparative analysis, and to use these estimates to examine the extent and consistency of species assignment of allopatric populations within rapidly evolving cichlid fish flocks present in three east African lakes. Location  Lakes Malawi, Victoria and Tanganyika. Methods  Using published taxon co-occurrence data, a novel approach was employed to calculate standardized ‘minimum’ estimates of regional species richness for hard substrate associated complexes of cichlids within each of the lakes. Minimum estimates were based on an explicit assumption that if taxa present on equivalent habitats have disjunct distributions, then they are allopatric forms of the same species. These estimates were compared with current observed ‘high-end’ regional species richness estimates for those complexes to determine the consistency of species assignment of allopatric populations between lineages within a lake. A ‘sympatry’ index was developed to enable comparisons of levels of species assignment of allopatric populations between-lakes to be made. Results  Within each lake, the minimum and high-end estimates for species richness were significantly correlated across complexes, indicating that the complexes that contain more recognized species contain the most genuine biological species. However, comparisons of complexes among lakes revealed considerable differences. For equivalent geographical areas, substantially higher proportions of recognized species were totally allopatric within the studied Lake Malawi and Lake Victoria complexes, than those of Lake Tanganyika. Main Conclusions  Among African lakes, levels of assignment to species status of allopatric populations were found to be distinctly different. It is unclear whether the discrepancies are a consequence of differences between the lake faunas in degrees of phenotypic divergence among allopatric populations, or are simply the result of inconsistent taxonomic practices. In either case, these results have considerable wider relevance for they emphasize that quantitative measures of regional and beta diversity are critically dependent on the species status of allopatric populations, an issue usually neglected in comparative studies of species richness. The technique introduced here can be used to standardize measures of regional diversity of lineages for comparative analyses, potentially enabling more accurate identification of processes influencing rates of speciation.

Contrasting long-term records of biomass burning in wet and dry savannas of equatorial East Africa

Rainfall controls fire in tropical savanna ecosystems through impacting both the amount and flammability of plant biomass, and consequently, predicted changes in tropical precipitation over the next century are likely to have contrasting effects on the fire regimes of wet and dry savannas. We reconstructed the long-term dynamics of biomass burning in equatorial East Africa, using fossil charcoal particles from two well-dated lake-sediment records in western Uganda and central Kenya. We compared these high-resolution (5 years/sample) time series of biomass burning, spanning the last 3800 and 1200 years, with independent data on past hydroclimatic variability and vegetation dynamics. In western Uganda, a rapid (<100 years) and permanent increase in burning occurred around 2170 years ago, when climatic drying replaced semideciduous forest by wooded grassland. At the century time scale, biomass burning was inversely related to moisture balance for much of the next two millennia until ca. 1750 ad, when burning increased strongly despite regional climate becoming wetter. A sustained decrease in burning since the mid20th century reflects the intensified modern-day landscape conversion into cropland and plantations. In contrast, in semiarid central Kenya, biomass burning peaked at intermediate moisture-balance levels, whereas it was lower both during the wettest and driest multidecadal periods of the last 1200 years. Here, burning steadily increased since the mid20th century, presumably due to more frequent deliberate ignitions for bush clearing and cattle ranching. Both the observed historical trends and regional contrasts in biomass burning are consistent with spatial variability in fire regimes across the African savanna biome today. They demonstrate the strong dependence of East African fire regimes on both climatic moisture balance and vegetation, and the extent to which this dependence is now being overridden by anthropogenic activity.

The genomic substrate for adaptive radiation in African cichlid fish

Cichlid fishes are famous for large, diverse and replicated adaptive radiations in the Great Lakes of East Africa. To understand the molecular mechanisms underlying cichlid phenotypic diversity, we sequenced the genomes and transcriptomes of five lineages of African cichlids: the Nile tilapia (Oreochromis niloticus), an ancestral lineage with low diversity; and four members of the East African lineage: Neolamprologus brichardi/pulcher (older radiation, Lake Tanganyika), Metriaclima zebra (recent radiation, Lake Malawi), Pundamilia nyererei (very recent radiation, Lake Victoria), and Astatotilapia burtoni (riverine species around Lake Tanganyika). We found an excess of gene duplications in the East African lineage compared to tilapia and other teleosts, an abundance of non-coding element divergence, accelerated coding sequence evolution, expression divergence associated with transposable element insertions, and regulation by novel microRNAs. In addition, we analysed sequence data from sixty individuals representing six closely related species from Lake Victoria, and show genome-wide diversifying selection on coding and regulatory variants, some of which were recruited from ancient polymorphisms. We conclude that a number of molecular mechanisms shaped East African cichlid genomes, and that amassing of standing variation during periods of relaxed purifying selection may have been important in facilitating subsequent evolutionary diversification.

Model simulation output data (CCSM3) related to uplift of African topography and the intensification of upwelling in the Benguela region

The Benguela Current, located off the west coast of southern Africa, is tied to a highly productive upwelling system**1. Over the past 12 million years, the current has cooled, and upwelling has intensified**2, 3, 4. These changes have been variously linked to atmospheric and oceanic changes associated with the glaciation of Antarctica and global cooling**5, the closure of the Central American Seaway**1, 6 or the further restriction of the Indonesian Seaway**3. The upwelling intensification also occurred during a period of substantial uplift of the African continent**7, 8. Here we use a coupled ocean-atmosphere general circulation model to test the effect of African uplift on Benguela upwelling. In our simulations, uplift in the East African Rift system and in southern and southwestern Africa induces an intensification of coastal low-level winds, which leads to increased oceanic upwelling of cool subsurface waters. We compare the effect of African uplift with the simulated impact of the Central American Seaway closure9, Indonesian Throughflow restriction10 and Antarctic glaciation**11, and find that African uplift has at least an equally strong influence as each of the three other factors. We therefore conclude that African uplift was an important factor in driving the cooling and strengthening of the Benguela Current and coastal upwelling during the late Miocene and Pliocene epochs.

(Table 1) Magnetic susceptibility readings in the western part of the Sør Rondane Montains

The Sør Rondane Mountains (SRM) in eastern Dronning Maud Land (DML) are located in an area, where two apparent Pan-African (650-520 Ma) orogenic mobile belts appear to intersect, the East African-Antarctic Orogen and the Kuunga Orogen. Hence, a better understanding of the tectonic structure of the Sør Rondane region is an important key for unravelling the complex geodynamic evolution of the eastern DML and adjacent regions of East Antarctica during the Late Neoproterozoic/Early Palaeozoic amalgamation of Gondwana. The SRM were recently (2011-2012) aerogeophysically investigated with a 5 km flight line spacing, covering a total area of ~140,000 km². The aeromagnetic data are correlated with ground-based magnetic susceptibility measurements and geological field data and allow to project tectonic terranes and individual structures into ice-covered areas. Magnetic anomalies and basement foliation trends are collinear in areas dominated by simple shear deformation, whereas an area of large-scale refolding correlates with a subdued small-scale broken magnetic anomaly pattern. The latter area can be regarded as a distinct tectonic domain, the central Sør Rondane corridor. It magnetically separates the SRM into an eastern, a central, and a western portion. This subdivision is presumably related to late Pan-African extensional tectonics and suggests that such a tectonic regime may play a larger role than previously assumed. Voluminous late Pan-African granitoids, which are mainly undeformed, correlate with positive magnetic anomalies between +30 and +80 nT, while a strong magnetic high (+680 nT) near the granitic intrusion at Dufekfjellet is caused by a highly magnetised enigmatic body. The recently discovered prominent magnetic anomaly province of southeastern DML continues into the southern part of the Sør Rondane region, where only a few outcrops are exposed. Findings at these westernmost nunataks of the SRM indicate that the subdued magnetic anomaly pattern of this southeastern DML province is most likely caused by the predominance of metasedimentary rocks of yet unknown age.

Geological map of Heimefrontfjella, Dronning Maud Land, Antarctica (Scale 1:250,000)

The geological overview map was compiled from 15 geological maps (1 : 25,000) and is based on Jacobs et al. 1996. The topographic basemaps were adapted from unpublished 1:250,000 provisional topographic maps, Institut f. Angewandte Geodäsie, Frankfurt, 1983. Part of the contour lines are from Radarsat (Liu et al. 2001).