Movement and habitat use by the marbled lungfish Protopterus aethiopicus Heckel 1851 in Lake Baringo, Kenya

The marbled lungfish, Protopterus aethiopicus, a recent introduction into Lake Baringo, Kenya is now an important commercial species there. Because little is known about its behaviour, we used ultrasonic telemetry to investigate its movements and use of habitat as part of a broader biological study. Twelve marbled lungfish were implanted with ultrasonic tags and tracked for variable periods between September 2001 and 2002. Two individuals were tracked for most of the study period. Daily movement ranged from little or none to 5.2 km. Mean hourly rates of movement for three fish located twice a day (morning and late afternoon) over several days suggested that individuals were active throughout the diel period. Maximum lake depth was about 3 m and fish utilized all depths greater than 1 m. Six home ranges described for four lungfish varied in size from 5.8 to 19.8 km ² and were occupied for between 2 and 4.5 months. Use of habitat and the movement of marbled lungfish in Lake Baringo appeared to be influenced more by biotic than abiotic factors.     

Marbled lungfish growth rates in Lake Baringo, Kenya, estimated by mark‐recapture

Mark‐recapture was used to quantify the relationship between body mass ( M _T) and individual growth rates of sub‐adult marbled lungfish Protopterus aethiopicus in Lake Baringo, Kenya. Specific growth rate (in mass) was found to be a well defined and decreasing function of M _T over the size range of recaptured fish. Growth trajectories based on this function indicated that Lake Baringo African lungfish reached maturity at an age of c . 3 years and had a low reproductive effort.     

Biology of the marbled lungfish, Protopterus aethiopicus Heckel, in Lake Baringo, Kenya

In this paper we provide life history characteristics for an introduced marbled lungfish (Protopterus aethiopicus) population in Lake Baringo, Kenya, including their length–weight relationship, body condition, fecundity, sex ratio, reproduction and food. These data are compared with those of other populations. Samples were obtained from the commercial fishery between January and October 2001. The lungfish exhibited positive allometric growth, with the length–weight relationship described by the equation: logW = ?6.41 + 3.52 logTL. The mean relative condition factor was close to unity, did not vary much between months and was not significantly different between sexes. The overall sex ratio was skewed in favour of females. The size of fish at first maturity was 70–76 cm total length (TL) for females and 82–88 cm TL for males. Fish in maturity stages III–IV occurred in all monthly samples suggesting year round spawning. The marbled lungfish in Lake Baringo is largely piscivorous and its impact on other fishes certainly warrants further research.     

Translocation of marbled African lungfish,Protopterus aethiopicus (Telostei: Protopteridae), and its fishery in Lake Baringo,Kenya

The translocation of the marbled African lungfish Protopterus aethiopicus into Lake Baringo created a new fishery for the local community, who capture them primarily in a bottom-set long line fishery. Its introduction, development and current fishery status in the lake are documented. Annual catch data were obtained from the District Fisheries Office, while catch and effort data of the long line fishery were recorded at one active fish-landing site between February and October 2001. Lungfish comprise a significant component of commercial landings, sometimes exceeding catches of the Baringo tilapia Oreochromis niloticus baringoensis as the most landed species by weight. Daily catch rates and effort varied considerably, ranging from 5.51–15.18kg day^?1 and from 50–590 baited hooks day^?1, respectively. Daily catch per fisher ranged from 0–35.72kg, while the overall mean catch per unit effort (CPUE) was only 0.003kg per hook-hour. This study provides the only baseline CPUE data for a lungfish long line fishery that the authors are aware of, for comparison with future studies in this or other lakes where lungfish are similarly exploited. Recommendations for improved management and the maintenance of a viable lungfish long line fishery in Lake Baringo are presented.     

Population-structure and genetic diversity in a haplochromine cichlid fish [corrected] of a satellite lake of Lake Victoria

The approximately 500 species of the cichlid fish species flock of Lake Victoria, East Africa, have evolved in a record-setting 100,000 years and represent one of the largest adaptive radiations. We examined the population structure of the endangered cichlid species Xystichromis phytophagus from Lake Kanyaboli, a satellite lake to Lake Victoria in the Kenyan Yala wetlands. Two sets of molecular markers were analysed--sequences of the mitochondrial control region as well as six microsatellite loci--and revealed surprisingly high levels of genetic variability in this species. Mitochondrial DNA sequences failed to detect population structuring among the three sample populations. A model-based population assignment test based on microsatellite data revealed that the three populations most probably aggregate into a larger panmictic population. However, values of population pairwise FST indicated moderate levels of genetic differentiation for one population. Eleven distinct mitochondrial haplotypes were found among 205 specimens of X. phytophagus, a relatively high number compared to the total number of 54 haplotypes that were recovered from hundreds of specimens of the entire cichlid species flock of Lake Victoria. Most of the X. phytophagus mitochondrial DNA haplotypes were absent from the main Lake Victoria, corroborating the putative importance of satellite lakes as refugia for haplochromine cichlids that went extinct from the main lake in the last decades and possibly during the Late Pleistocene desiccation of Lake Victoria.     

Patterns in fish radiation are compatible with Pleistocene desiccation of Lake Victoria and 14,600 year history for its cichlid species flock

Geophysical data are currently being interpreted as evidence for a late Pleistocene desiccation of Lake Victoria and its refilling 14,600 years ago. This implies that between 500 and 1000 endemic cichlid fish species must have evolved in 14,600 years, the fastest large-scale species radiation known. A recent review concludes that biological evidence clearly rejects the postulated Pleistocene desiccation of the lake: a 14,600 year history would imply exceptionally high speciation rates across a range of unrelated fish taxa. To test this suggestion, I calculated speciation rates for all 41 phylogenetic lineages of fish in the lake. Except for one cichlid lineage, accepting a 14 600 year history does not require any speciation rates that fall outside the range observed in fishes in other young lakes around the world. The exceptional taxon is a lineage of haplochromine cichlids that is also known for its rapid speciation elsewhere. Moreover, since it is unknown how many founding species it has, it is not certain that its speciation rates are really outside the range observed in fishes in other young lakes. Fish speciation rates are generally faster in younger than in older lakes, and those in Lake Victoria, by far the largest of the young lakes of the world, are no exception. From the speciation rates and from biogeographical observations that Lake Victoria endemics, which lack close relatives within the lake basin, have such relatives in adjacent drainage systems that may have had Holocene connections to Lake Victoria, I conclude that the composition of the fish assemblage does not provide biological evidence against Pleistocene desiccation. It supports a hypothesis of recent colonization from outside the lake basin rather than survival of a diverse assemblage within the basin.     

High prevalence of non-synonymous substitutions in mtDNA of cichlid fishes from Lake Victoria

When a population size is reduced, genetic drift may fix slightly deleterious mutations, and an increase in nonsynonymous substitution is expected. It has been suggested that past aridity has seriously affected and decreased the populations of cichlid fishes in Lake Victoria, while geographical studies have shown that the water levels in Lake Tanganyika and Lake Malawi have remained fairly constant. The comparably stable environments in the latter two lakes might have kept the populations of cichlid fishes large enough to remove slightly deleterious mutations. The difference in the stability of cichlid fish population sizes between Lake Victoria and the Lakes Tanganyika and Malawi is expected to have caused differences in the nonsynonymous/synonymous ratio, ω (= d_N/d_S), of the evolutionary rate. Here, we estimated ω and compared it between the cichlids of the three lakes for 13 mitochondrial protein-coding genes using maximum likelihood methods. We found that the lineages of the cichlids in Lake Victoria had a significantly higher ω for several mitochondrial loci. Moreover, positive selection was indicated for several codons in the mtDNA of the Lake Victoria cichlid lineage. Our results indicate that both adaptive and slightly deleterious molecular evolution has taken place in the Lake Victoria cichlids' mtDNA genes, whose nonsynonymous sites are generally conserved.     

Genetic affinities of an introduced predator: Nile perch in Lake Victoria, East Africa

Although the introduction of Nile perch, Lates niloticus , to Lake Victoria has received intense global attention, especially in relation to its impact on endemic cichlid species and on fishery yields, fundamental information on its taxonomy and population genetics is lacking. Most importantly, the introduced fish originated from two lakes (Lakes Albert and Turkana) containing three Lates species, and it has never been entirely clear which of these became established in Lake Victoria, or indeed whether the Lake Victoria population is derived from hybridization between Lates species. In addition, genetic drift caused by the relatively small founder population (≈ 400), the initially slow population increase followed by a period of explosive population growth, and selection pressures in the new environment may have resulted in substantial genetic changes. Allozyme data indicated that the introduced Nile perch of Lake Victoria were mainly L. niloticus from Lake Albert, although maximum likelihood estimates of stock contributions (GSI) suggested the presence of L. macrophthalmus. In contrast, introduced Nile perch in adjacent smaller lakes (Lakes Kyoga and Nabugabo) appeared to be entirely L. niloticus . The effect of the introductions on allozyme diversity varied among lakes and appeared to be uncorrelated to the number of fish introduced.     

Decline of the African lungfish (Protopterus aethiopicus) in Lake Victoria (East Africa)

Catch and effort data for the period 1973–1990 demonstrate a dramatic decline of lungfish in the Tanzanian waters of Lake Victoria. Bottom trawl catches in the Mwanza Gulf showed a decline in catch rates from 67.5 kg h⁻¹ in 1973 to 5.5 kg h⁻¹ in 1986. Trawling of commercial vessels in the Speke Gulf revealed a decline in lungfish catches from 1.3 kg h⁻¹ in 1986 to 0.07 kg h⁻¹ in 1990. The development of anoxia in the deeper waters of Lake Victoria, the algal blooms, and the decline of water transparency, all associated with eutrophication, are not likely to have contributed to the decreased catch rate. However, the lungfish decline may reflect the interaction of overexploitation by the fishery and a low level of Nile perch predation that restricts lungfish to wetland refugia. We suggest that this may have been reinforced over the past few decades by large-scale conversion of wetlands to agricultural land and harvesting of nest-guarding male lungfish leading to decreased recruitment of young.     

Reconciling geography and genealogy: phylogeography of giant freshwater prawns from the Lake Carpentaria region

There is convincing geological evidence for the historical existence of an ancient lake on the Australian-New Guinea continental shelf during the late Pleistocene. Lake Carpentaria was a vast fresh- to brackishwater lake that would presumably have provided habitat for, and facilitated gene flow among, aquatic taxa that tolerate low to moderate salinities in this region. Moreover, it has been argued that the outflow of Papua New Guinea's Fly River was diverted westward into Lake Carpentaria during this period, although this hypothesis is controversial. We predicted that these events, if a true history, would have promoted gene flow and population growth via range-expansion events in the giant freshwater prawn (Macrobrachium rosenbergii) and restricted gene flow subsequently by way of a vicariant event as sea levels rose during the late Pleistocene, and a marine environment replaced Lake Carpentaria. We tested these hypotheses using phylogeographical and phylogenetic analyses of mitochondrial DNA variation in M. rosenbergii populations sampled from the Lake Carpentaria region. Our results support the hypothesis that Lake Carpentaria facilitated gene flow among populations of M. rosenbergii that are today isolated, but contest claims of a westward diversion of the Fly River. We inferred the timing of initial expansion in the 'Lake Carpentaria lineage' and found the timing of this event to be broadly concordant with geological dating of the formation of Lake Carpentaria. Reconciling geological and molecular data, as presented here, provides a powerful framework for investigating the influence of historical earth history events on the distribution of biological (i.e. molecular) diversity.     

Impact of rifting and hydrography on the genetic structure of Clarias gariepinus in eastern Africa

In a study of the genetic structure of Clarias gariepinus at the ND5 and ND6 loci of mitochondrial DNA using RFLP–PCR in 16 populations (267 fish), a total of 18 haplotypes was detected. The most common haplotype was EA1 and represented 46% of all genotypes. Three phylogenetic groups characterized the region, with the East African clade occurring between the western rift in the west, Lake Baringo in the north, the Indian Ocean in the east and the Rufiji River in the south. The Lower Tana River (unlike the Upper Tana River) and Lake Kamnarok grouped with the North clade, while the Ruaha River (tributary of the Rufiji River) represented a contact zone between the East and the South-central African clade. Genetic structure was distinct ( F _ST=0·66), with the Lower Tana River and Lake Mtera forming specific units. Within the East clade, the Oloibortoto River, Upper Tana River and Lake Victoria each formed a distinct population. Lake Baringo grouped either with the eastern rift or the western rift depending on the analysis. Lake Jipe, the Pangani River, Rusizi River, Lake Edward, Nyabugogo River and the Luiche River clustered together. Among group variance explained 66·8% of the total variance; the impact of vicariance (rifting and uplifting) on the phylogeography was considerable. A distribution model of the catfish is proposed which combines the separation by rifting several million years ago with a high gene flow in the western rift due to historical connectivity among lakes and rivers.     

The bounty of minor lakes: the role of small satellite water bodies in evolution and conservation of fishes in the Lake Victoria Region,East Africa

The Lake Victoria Region (LVR) encompasses the large lakes Victoria, Kyoga, Edward, George and Kivu, as well as scores of small satellite lakes within the parent catchments. Taken as a whole, the LVR originally harbored a unique fish fauna that included in excess of 600 endemic species of cichlid fishes. As a result of human influence, including a commercial fishery and the introduction of several exotic species, nearly 200 cichlid species and several endemic genera have become extinct from lakes Victoria and Kyoga. Recently, we have discovered that some of the apparently extinct taxa survive as extant representatives in the satellite lakes. Here, we summarize the findings of our ecological survey of the fish species of the satellite lakes. We also discuss the results of some preliminary genetic analyses, and highlight major genetic and ecological changes in the fish fauna that have taken place in the regional fishery. Minor lakes now play a crucial role in conserving the endangered species of the entire region, and also as living museums of East African ecological history. Our findings allude to the historical importance of minor satellite lakes as natural refugia for the fishes of the Lake Victoria Region, a region characterized by a history of geological and climatic instability.     

Testing biogeographical hypotheses in space and time: faunal relationships of the putative ancient Lake Egˇirdir in Asia Minor

Aim The aims of this study are to establish a multi‐locus phylogeny‐based hypothesis for the biogeographical relationship of gastropods from the putative ancient Lake Egˇirdir, to test the respective null hypothesis, to estimate the timing of biogeographical events based on independent molecular clock approaches, and to interpret the data with respect to the putative ancient character of Lake Egˇirdir. Location Lake Egˇirdir, western Taurus Lake District, Turkey. Methods DNA sequences from the putatively only extant endemic taxon of Lake Egˇirdir, Falsipyrgula pfeiferi, as well as representatives of other pyrgulinid genera from Europe and western Asia are used for phylogenetic analyses based on Bayesian inference. The respective null hypothesis is tested utilizing parametric bootstrapping. The timing of evolutionary events is estimated based on two independent molecular clock approaches, which involve the modelling of judicious errors associated with branch‐length calculations and calibration points. Results Bayesian inference indicates a very close relationship between the Lake Egˇirdir and Ponto‐Caspian taxa. Parametric bootstrapping rejects the null hypothesis that these taxa are not monophyletic (P ≤ 0.01). The alternative hypothesis, namely monophyly of the Ponto‐Caspian and Egˇirdir taxa, can therefore be accepted. The two independent molecular clock approaches show diversion times for the Ponto‐Caspian/Egˇirdir taxa of 0.42 ± 0.18 and 0.43 ± 0.63 Ma. Main conclusions The present study shows that there is no close biogeographical affiliation between the probably only remaining endemic taxon of Lake Egˇirdir and taxa from central Europe or the Balkan region. Instead, there is a very close and relatively young (i.e. late Pleistocene) biogeographical relationship with the Ponto‐Caspian pyrgulinids. However, fossil and comparative data from other invertebrates indicate that biogeographical connections between Lake Egˇirdir and the Ponto‐Caspian region existed during various time periods, i.e. the Miocene/Pliocene, early Pleistocene, and late Pleistocene. Acknowledging the still‐restricted knowledge of the evolutionary history of the lake, the data presented here do not reject the putative ancient status of Lake Egˇirdir. Future studies utilizing endemic taxa of other lakes in the region need to show whether the western Taurus Lake District represents a melting pot of Pleistocene refuge biodiversity from different regions, and whether the admixture of divergent lineages has created a genetically distinct set of taxa that would warrant the designation of the area as a unique biogeographical subregion.     

Characterization of Vibrio cholerae Bacteriophages Isolated from the Environmental Waters of the Lake Victoria Region of Kenya

Over the last decade, cholera outbreaks have become common in some parts of Kenya. The most recent cholera outbreak occurred in Coastal and Lake Victoria region during January 2009 and May 2010, where a total of 11,769 cases and 274 deaths were reported by the Ministry of Public Health and Sanitation. The objective of this study is to isolate Vibrio cholerae bacteriophages from the environmental waters of the Lake Victoria region of Kenya with potential for use as a biocontrol for cholera outbreaks. Water samples from wells, ponds, sewage effluent, boreholes, rivers, and lakes of the Lake Victoria region of Kenya were enriched for 48 h at 37 °C in broth containing a an environmental strain of V. cholerae. Bacteriophages were isolated from 5 out of the 42 environmental water samples taken. Isolated phages produced tiny, round, and clear plaques suggesting that these phages were lytic to V. cholerae. Transmission electron microscope examination revealed that all the nine phages belonged to the family Myoviridae, with typical icosahedral heads, long contractile tails, and fibers. Head had an average diameter of 88.3 nm and tail of length and width 84.9 and 16.1 nm, respectively. Vibriophages isolated from the Lake Victoria region of Kenya have been characterized and the isolated phages may have a potential to be used as antibacterial agents to control pathogenic V. cholerae bacteria in water reservoirs.     

Nuclear markers reveal unexpected genetic variation and a Congolese-Nilotic origin of the Lake Victoria cichlid species flock

Phylogenetic analyses based on mitochondrial (mt) DNA have indicated that the cichlid species flock of the Lake Victoria region is derived from a single ancestral species found in East African rivers, closely related to the ancestor of the Lake Malawi cichlid species flock. The Lake Victoria flock contains ten times less mtDNA variation than the Lake Malawi radiation, consistent with current estimates of the ages of the lakes. We present results of a phylogenetic investigation using nuclear (amplified fragment length polymorphism) markers and a wider coverage of riverine haplochromines. We demonstrate that the Lake Victoria-Edward flock is derived from the morphologically and ecologically diverse cichlid genus Thoracochromis from the Congo and Nile, rather than from the phenotypically conservative East African Astatotilapia. This implies that the ability to express much of the morphological diversity found in the species flock may by far pre-date the origin of the flock. Our data indicate that the nuclear diversity of the Lake Victoria-Edward species flock is similar to that of the Lake Malawi flock, indicating that the genetic diversity is considerably older than the 15 000 years that have passed since the lake began to refill. Most of this variation is manifested in trans-species polymorphisms, indicating very recent cladogenesis from a genetically very diverse founder stock. Our data do not confirm strict monophyly of either of the species flocks, but raise the possibility that these flocks have arisen from hybrid swarms.     

Is speciation of demersal fishes in Lake Tanganyika restrained by physical limnological conditions?

Both Lake Malawi and Lake Victoria have many species of offshore demersal cichlids, most of which are stenotopic. In Lake Tanganyika there are fewer, more eurytopic, demersal species. These differences may be the effect of the physical regimes of the lakes. In Lake Victoria there is no permanent anoxic layer. In both the other lakes, water below 250 m depth is permanently anoxic and the seasonal cycle is dominated by upwelling. In Lake Malawi this is mainly derived from intermediate water containing free oxygen. In Lake Tanganyika upwelling involves hypolimnetic water, which may rise to within 80 m of the surface, causing great short-term changes in oxygen concentration over the depth range 50–250 m. This has inhibited the development of deep water species restricted to narrow depth ranges.     

Evolution of the ribosomal RNA internal transcribed spacer one (ITS-1) in cichlid fishes of the Lake Victoria region

The nucleotide sequences of the first internal transcribed spacer (ITS-1) of the ribosomal RNA gene cluster have been determined for 11 species of closely related endemic cichlid fishes of the Lake Victoria region (LVR) and 6 related East African cichlids. The ITS-1 sequences confirmed independently derived basal phylogenies, but provide limited insight within this species flock. The line leading to Pseudocrenilabrus multicolor arose early, close to the divergence event that separated the tilapiine and haplochromine tribes of the "African Group" of the family Cichlidae. In this phylogeny, Astatoreochromis alluaudi and the riverine Astatotilapia burtoni are sister taxa, which together are a sister group to a monophyletic assemblage including both Lake Victoria and Lake Edward taxa. The ITS-1 data support the monophyly of haplochromine genera across lakes. Since Lake Victoria is believed to have been dry between 14, 500 and 12,400 BPE, the modern assemblage must have been derived from reinvasion by the products of earlier cladogenesis events. Thus, although the regional superflock is monophyletic, the haplochromines of Lake Victoria itself did not evolve in situ from a single ancestor.     

Lake level fluctuations synchronize genetic divergences of cichlid fishes in African lakes

Water level fluctuations are important modulators of speciation processes in tropical lakes, in that they temporarily form or break down barriers to gene flow among adjacent populations and/or incipient species. Time estimates of the most recent major lowstands of the three African Great Lakes are thus crucial to infer the relative timescales of explosive speciation events in cichlid species flocks. Our approach combines geological evidence with genetic divergence data of cichlid fishes from the three Great East African Lakes derived from the fastest-evolving mtDNA segment. Thereby, we show for each of the three lakes that individuals sampled from several populations which are currently isolated by long geographic distances and/or deep water form clusters of equally closely related haplotypes. The distribution of identical or equally closely related haplotypes in a lake basin allows delineation of the extent of lake level fluctuations. Our data suggest that the same climatic phenomenon synchronized the onset of genetic divergence of lineages in all three species flocks, such that their most recent evolutionary history seems to be linked to the same external modulators of adaptive radiation. A calibration of the molecular clock of the control region was elaborated by gauging the age of the Lake Malawi species flock through the divergence among the utaka-cichlid and the mbuna-cichlid lineages to minimally 570,000 years and maximally 1 Myr. This suggests that the low-lake-level period which established the observed patterns of genetic relatedness dates back less than 57,000 years, probably even to 17,000-12,400 years ago, when Lake Victoria dried up and Lakes Malawi and Tanganyika were also low. A rapid rise of all three lakes about 11,000 years ago established the large-scale population subdivisions observed today. Over that period of time, a multitude of species originated in Lakes Malawi and Victoria with an impressive degree of morphological and ecological differentiation, whereas the Tanganyikan taxa that were exposed to the same habitat changes hardly diverged ecologically and morphologically. Our findings also show that patterns of genetic divergences of stenotopic organisms provide valuable feedback on geological and sedimentological time estimates for lake level changes.     

Sediment formation in Lake Baikal

The following stages in the formation of sediments in Lake Baikal have been distinguished. In the Middle Miocene lacustrine deposits accumulated in the southern depression. In Early and Middle Pliocene, as well as in Early-Middle Pleistocene the bottom of the southern and middle Baikal depressions was occupied by large lakes, but the northern basin was mainly dry with a system of small lakes, streams and rivers. This accounts for the great facies variety and the genetic heterogeneity of sediments in the northern depression. At the end of Middle and in the Late Pleistocene an abrupt warping of the depression took place, and a shore-line similar to the present one was formed. Sands in the upper part of the sediments are considered to be the product of intensive erosion which probably occurred in the pluvial epoch in the Late Pleistocene period.