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.     

A revision of the genus Labidochromis (Teleostei: Cichlidae) from Lake Malawi

The diagnosis of the genus Labidochromis is revised to include species with bicuspid outer teeth. Sixteen species, of which 13 are new, are described and illustrated and the validity of five others is discussed. The interrelationships between members of the genus and the relationship of the genus to other genera are discussed. No clearly defined sister group to the genus has been found either within or outside the Lake Malawi cichlid species flock. A key based on the coloration of known species is included.     

A sympatric sibling species complex of Petrotilapia Trewavas from Lake Malawi analysed by enzyme electrophoresis (Pisces: Cichlidae)

The protein electromorphs of three taxa of Petrotilapia , referred to in earlier works as sibling species, were examined by starch-gel electrophoresis. The results indicated that, for the three taxa, no alternative fixed alleles occurred for any of the 25 loci (enzymes) examined. However, heterogeneous gene frequencies were indicated at seven polymorphic loci. This suggests that either the taxa are isolated 'sibling' species which recently diverged, or that they are 'incipient' species with minimal gene flow between morphs. The genetic and field data lend support to the hypothesis that sympatric splitting of morphs could be important in the explosive radiation of the cichlidae.     

Description of a commercially important pelagic species of the genus Diplotaxodon (Pisces: Cichlidae) from Lake Malawi, Africa

A new species of pelagic haplochromine cichlid of the genus Diplotaxodon is described from Lake Malawi, Africa. The species, which attains a size of 14cm s. L., feeds mainly on zooplankton, shows no sexual dimorphism in size or body proportions, breeds throughout the year and lives at depths of 20 to at least 120 m. Approximately 600 tonnes of the species were caught by trawlers in 1990–1991, in the south–eastern arm of the lake alone. It appears that this species is a major component of the pelagic community, and occupies an ecological role similar to that of the clupeid Limnothrissa miodon in Lake Tanganyika.     

A comparison of the abilities of three sympatric species of Petrotilapia (Cichlidae,Lake Malawi) to penetrate deep water

Synopsis The ability of three sympatric species ofPetrotolapia, from Monkey Bay, Lake Malawi, to penetrate deep water was tested using pressure chambers. The three species were found to differ in terms of the maximum pressure (= depth) to which each was able to equilibrate. Mean maximum equilibration pressures were for the form designated MOL 39.0 m, for MBB 35.5 m and for the form MOC 30.4 m. The rate of compensation to pressure changes was slower in MBB (2.59 m day^–1) than in other two forms (MOL = 3.97 m day^–1, MOC = 3.98 day^–1). Natural depth distribution of all three forms, in particular MBB and MOC, is not determined by limits imposed by swin bladder physiology.     

Factors in habitat partitioning in Pseudotropheus aurora (Pisces: Cichlidae), an introduced species to a species-rich community of Lake Malawi

The territorial behaviour of a small rock-dwelling cichlid, Pseudotropheus aurora , was studied at Thumbi Island West at the southern end of Lake Malawi. P. aurora is one of several species that have been introduced to an already diverse fish community. Many of the endemic and introduced species are highly territorial. The coexistence of an introduced territorial species within a speciesrich community of other territorial species is examined. Territories from which P. aurora individuals were removed were invariably re-occupied by conspecifics. Similarly, when all territories of a certain area were emptied of all territory owners of any existing species, the former P. aurora territories were re-occupied by conspecifics.
The possible roles of site specificity, competitive exclusion and behaviourally transmitted traditions regarding territories are discussed.     

Evolution of the mitochondrial DNA control region in the mbuna (Cichlidae) species flock of lake Malawi, East Africa

Considerable controversy has surrounded the application of mitochondrial DNA data to reconstruction of evolutionary relationships among the endemic cichlids of Lake Malawi. Central to this debate has been the issue of whether lineage sorting is complete, and thus whether these data actually reflect species phylogeny, or simply gene genealogy. Review of all mtDNA control region sequences available for members of one monophyletic subset of this species flock, the Malawi rockfishes, or mbuna, strongly indicates that lineage sorting is incomplete: Character-based analyses of these sequences reconstruct gene, not species, interrelationships. Analysis of the pattern of nucleotide substitutions differentiating these mtDNA alleles suggests that pyrimidine residues undergo transition substitutions more often than do purines. Estimation of the magnitude of derived sequence differentiation in light of the reconstructed gene genealogy suggests that the mbuna may be of considerably more recent vintage than previous molecular characterizations have indicated. Received: 6 April 1996 / Accepted: 3 March 1997     

Identification of the Lake Malawi Oreochromis (Nyasalapia) spp. using multivariate morphometric techniques

With the aim of reliably distinguishing these commercially important species on the basis of external characteristics alone, morphometric techniques were employed on a sample of the four species of Oreochromis ( Nyasalapia ) described from Lake Malawi: O. karongae, O. lidole, O. saka and O. squamipinnis . Univariate analysis of variance on the ratios of 23 variables to standard length indicated many differences among all species, but there was considerable individual variation, and consequent overlap. Residuals from a regression of each variable on length were employed for multivariate analysis. Cluster analysis on the means of the residuals was used to construct a phenogram which formed the basis for denning a series of dichotomous discriminant analyses. In each discriminant analysis, variables were successively eliminated in the reverse order of the magnitude of their correlation with the discriminant function. The combination of variables producing 95% accuracy of classification was selected, and the discriminant function equations for each step calculated. Some further variables were eliminated by checking for redundancy through analysis of correlations. The resulting equations enable O. karongae to be separated using eight measurements, O. lidole using 10, and O. saka and O. squamipinnis to be distinguished by a combination of 13 measurements.     

Nuclear gene variation and molecular dating of the cichlid species flock of Lake Malawi

The cichlid fishes of Lake Malawi are famously diverse. However, evolutionary studies have been difficult because of their recent and uncertain phylogenetic history. Portions of 12 nuclear loci were sequenced in nine rock-dwelling species (mbuna) and three representatives of pelagic nonmbuna species. In contrast to the pattern of variation at mitochondrial genes, which do provide phylogenetic resolution at the level of mbuna versus nonmbuna, and among some genera, the nuclear loci were virtually devoid of phylogenetic signal. Only a small minority of variable positions were phylogenetically informative, and no phylogenetic branches are supported by more than one site. From the nuclear gene perspective the Malawian radiation appears to be a star phylogeny, as if the founding of the lake was accompanied by a partial bottleneck. The pattern is different from that found in Lake Victoria, in which nuclear loci share large amounts of ancestral variation. In the case of nuclear genes of Lake Malawi, the absence of phylogenetically informative variation suggests a relative absence of ancestral variation. Nuclear genes also differed from the mitochondria in having nearly twice the amount of divergence from Oreochromis (tilapia). An approximate splitting time between mbuna and nonmbuna lineages was estimated as 0.7 Myr. Oreochromis is estimated to have diverged from the cichlids in Lake Malawi and Lake Tanganyika about 18 MYA.     

A new species and geographical variation in the Telmatochromis temporalis complex (Teleostei, Cichlidae) from Lake Tanganyika

Telmatochromis brachygnathus sp.n. is described from the southern and central parts of Lake Tanganyika. It can be distinguished from the similar T. temporalis mainly by its smaller mouth. Morphological distinct populations were found in both species.     

Elevated mtDNA diversity in introduced populations of Cynotilapia afra (Günther 1894) in Lake Malawi National Park is evidence for multiple source populations and hybridization

Genetic variation in many invasive species shows little or no signs of a founder event, suggesting that high genetic diversity may facilitate establishment success. The rocky‐shore, plankton‐feeding cichlid fish Cynotilapia afra is endemic to Lake Malawi, but naturally absent from many suitable sites. In the 1960s, this species was introduced to the southern areas of the lake, presumably as a result of the aquarium fish trade. It has now become established on a number of rocky areas within the Lake Malawi National Park. Here, we analysed DNA sequence variation in the mitochondrial control region of six native and four introduced populations of C. afra, and three populations of the closely‐related and hybridizing Pseudotropheus zebra. In contrast to previous studies of Lake Malawi rock dwelling cichlids, network analyses suggested that native populations of C. afra showed high levels of lineage sorting in mtDNA. Introduced populations showed higher sequence and haplotype diversity than their native counterparts. Our analyses suggested that the elevated gene diversity was largely attributed to the fact that the introduced C. afra populations were derived from several genetically distinct and geographically separate populations, and to a lesser extent because of introgressive hybridization with native P. zebra. The establishment and spread of C. afra may be partly because of its ability to occupy a vacant ecological niche, but it may also have been facilitated by its enhanced genetic diversity.     

No evidence for parallel sympatric speciation in cichlid species of the genus Pseudotropheus from north-western Lake Malawi

To test the hypothesis of parallel speciation by sexual selection, we examined length variation at six microsatellite loci of samples from four sites of four to six putative species belonging to two subgenera of rocky shore mbuna cichlids from Lake Malawi. Almost all fixation indices were significantly different from zero, suggesting that there is presently little or no gene flow among allopatric populations or sympatric species. Analysis of variance indicated that genetic distances among allopatric populations of putative conspecifics were significantly lower than among sympatric populations of heterospecifics. The topology of trees based on distance matrices was also largely consistent with the hypothesis that the putative species are monophyletic and have thus not evolved in parallel in their present locations. If parallel speciation does occur in Malawi cichlids, it may be on a larger spatial scale than investigated in our study.     

Extreme microallopatric divergence in a cichlid species from Lake Malawi

We demonstrate significant population structuring on an extremely small spatial scale between adjacent demes of a Lake Malawi haplochromine cichlid species of the mbuna group, Pseudotropheus callainos, separated by only 35 m of habitat discontinuity. This substantiates further the notion that intralacustrine allopatric divergence may help to explain the high level of species richness of the mbuna in comparison to other Malawian cichlids, as well as of the Malawian haplochromines as a whole.     

Human rights,environment, and development: The dispossession of fishing communities on Lake Malawi

In a growing number of cases throughout Africa, communities' resource bases are being undermined or appropriated by outsiders, a process which seriously threatens the continuation of local cultures and livelihoods. In this article, we use a political ecology framework to examine how the linked processes of economic development, political power, and environmental change are transgressing the rights of fishing communities on the shores of Lake Malawi. In the cases described, these communities, or community members within them, find themselves powerless to prevent the expropriation of the resources over which they previously had either legal or customary control. Thus, it is not the economic processes of dispossession alone which lead to human rights violations but rather dispossession combined with an authoritarian political context.     

Oxygen concentration profiles in sediments of two ancient lakes: Lake Baikal (Siberia,Russia) and Lake Malawi (East Africa)

Oxygen concentration profiles have been measured, by means of with microelectrodes in sediments of Lake Baikal and Lake Malawi, along transects allowing to give a survey of two major ancient Rift lakes: Lake Baikal (Eastern Siberia) and Lake Malawi (East Africa), along depth transects in the constitutive basins of the lakes and/or of relevant depths with regard to oxygen (including including the deepest point, 1680 m, in Lake Baikal). Sediment oxygen penetration depths (SOPs) display very different patterns, depending on the lake in the two lakes. In Lake Baikal, SOPs are variable, show no significant relationship with bathymetric depth and are surprisingly deep on Akademichesky ridge (> 50.0 mm), emphasizing the distinctive feature of this region in the lake. While the Selenga river is an important source of eutrophication, the similarity of SOP-values in the Selenga shallow with those of most other sites suggests either a dilution of organic material by allochthonous matter, or a strong south-to-north transport of particles. In Lake Malawi, available oxygen is restricted to a maximum of three millimetres of the sediment, and there is a negative relationship with bathymetric depth, as a result of a steady decline of oxygen concentration with depth through the water column. Amongst the few parameters known to affect SOPs, the oxygen consumption by the sediment seems the most significant in both lakes. SOP-values furthermore confirm differences in the trophic status of Baikal and Malawi, respectively. The importance of oxygen as a factor likely to create ecological segregation for benthic organisms is discussed. Lake Malawi offers possibilities of bathymetric segregation but no vertical segregation in the sediment. In contrast, no bathymetric segregation related to oxygen is possible in Lake Baikal, but vertical segregation in the sediment is very likely. This revised version was published online in July 2006 with corrections to the Cover Date.     

Possible species specific courtship sounds by two sympatric cichlid fishes in Lake Malawi, Africa

Sounds were produced by the males of two species of cichlid fishes while courting females. Each courtship sound consisted of a series of distinct pulses occurring in rapid succession. Courtship sounds produced by Tramitichromis cf. intermedius and Copadichromis conophorus were significantly different in pulse rate and individual pulse durations. For C. conophorus calls (n=127) the mean ± sd number of pulses per call was 10 ± 3 and call duration was 181 ± 59 ms. There was a significant positive linear relationship between call duration and the number of pulses (r²=0.912, p < 0.001). The dominant frequency of the pulses in calls was 471 ± 50 (range 372–594) Hz (n=40 calls). T. cf. intermedius also produced a pulsed courtship call; data (mean ± sd) from two male T. cf. intermedius: 9 ± 2 pulses per call and duration 199 ± 44 ms (n=20 calls). The linear regression between call duration and number of pulses was positive (r²=0.463, p=0.001). Pulse rate within calls of T. cf. intermedius compared to C. conophorus were significantly different (p=0.018). Individual pulse durations were also significantly different (p=0.043) between species. However, interpulse intervals were not significantly different (p=0.177). These cichlids produced courtship sounds that were distinct by individual pulse durations and by pulse repition rate in a call.     

Evolutionary relationships of the Lake Malawi Oreochromis species: evidence from allozymes

Three tilapiine species belonging to the endemic Lake Malawi species flock known as' chambo' Oreochromis ( Nyasalapia ) karongae , O. (N.) lidole and O. (N.) squamipinnis and the species Oreochromis (Oreochromis) shiranus were collected from the wild. These four species were analysed at 43 enzyme loci using starch gel electrophoresis. No significant deviations from Hardy-Weinberg equilibrium were observed after correcting for multiple simultaneous testing. The expected heterozygosity ( H _e) was lowest in O. (O.) shiranus ( H _e=0.082). The three chambo species had higher levels: H _e=0.103 to H _e=0.116. The three chambo species as a whole were polymorphic (99%) at 16 different loci sharing 10 of these in common. No fixed differences between the three chambo species were observed but highly significant allele frequency differences existed between all the chambo species. The F _ST calculated for the three chambo species was 0051, closer to intra-specific than to inter-specific levels. O. (O.) shiranus could be clearly separated from all chambo species at five fixed loci. Comparison of the allozyme data from these species with five species from the same sub genera support the hypothesis that the chambo form a monophyletic group and that O. (N.) macrochir or a related species represents the sister taxon.     

Isotopic change throughout the life history of a Lake Malawi cichlid fish

Clear changes in body size-isotope (carbon and nitrogen) trajectories of Pseudotropheus callainos , a cichlid belonging to the endemic haplochromine species radiation in Lake Malawi, were found that corresponded with an ontogenetic dietary shift from predominantly planktonic to benthic food sources. The results indicated that dietary switching was a proximate cause of isotopic change over the life history of this species and confirmed the value of stable isotope signatures for inferring diet. The data also illustrated that possible variability of signatures over the life history of a species should be considered when using stable isotope ratios to investigate fine-scale ecological differentiation among taxa.     

Signal and preference divergence among populations of the non‐endemic basal Lake Malawi cichlid fish Astatotilapia calliptera (Perciformes: Cichlidae)

In Lake Malawi and Lake Victoria, cichlid fishes have diversified into hundreds of species, many reproductively isolated by mate choice. Territorial males tend to be more aggressive to similar‐coloured males, facilitating coexistence of divergent colour morphs or species. Behavioural mate choice and aggression biases of species and allopatric populations of specialized rocky shore cichlids are influenced by divergent signals such as male colour. Believed to be basal to the Lake Malawi haplochromine radiation, and inhabiting shallow weedy areas of the lake and neighbouring water bodies, Astatotilapia calliptera also shows allopatric variation in colour. Here, it is demonstrated that such signal divergence is associated with tendencies of females to mate with males of their own population and also for males to preferentially attack males of their own population, indicating that preference divergence related to signal divergence in allopatry may have operated throughout the adaptive radiation of the Malawian cichlids. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 180–188.