Feeding schools among Lake Malawi cichlid fishes

Synopsis Seven species of rock-frequenting cichlids, endemic to Lake Malawi, formed feeding-schools. Petrotilapia genalutea was the nuclear species comprising 92% of the school members (n = 2690 individuals from 50 schools). Fifty-eight per cent of all schools encountered consisted only of P. genalutea. School size ranged from 10 to 300 members (x = 81 ± 74 SD, median = 62). Schooling behaviour was facultative. Ninety-seven per cent of the feeding sites used by the schools occurred within the territories of highly aggressive, territorial species. These sites support considerably greater standing crops of algae, the food of the schooling species, than undefended sites. Individuals of schooling species were unable to gain access to the aggresively defended areas unless they joined schools.     

Translocations of rocky habitat cichlid fishes to Nkhata Bay, Lake Malawi

Over the last 10 years, 12 rocky habitat 'mbuna' taxa have been translocated to Nkhata Bay on the north-western shores of Lake Malawi from other regions of the lake. Given potential for competition and hybridization with indigenous taxa, further translocations should be avoided.     

Aggressive biases towards similarly coloured males in Lake Malawi cichlid fishes

In haplochromine cichlids, female mate choice based on male nuptial coloration has played an important role in speciation. Recent studies suggest that male coloration strongly influences the distribution of these fishes based on male-male aggression; males direct more aggression towards similarly coloured opponents while tolerating differently coloured individuals. We explored the role of male nuptial colour in aggression among the mbuna of Lake Malawi, examining aggression by male Metriaclima mbenjii, the red top cobalt zebra, towards conspecific opponents, similarly coloured heterospecific opponents and differently coloured heterospecifics. In trials in which focal males were offered a single opponent, while the total number of aggressive behaviours did not vary among opponent species, the types of behaviours did; focal males directed more lateral displays towards conspecifics than towards the other opponent species. When focal males were offered two opponents simultaneously, M. mbenjii directed more aggressive behaviours and more lateral displays towards similarly coloured opponents, regardless of species. Furthermore, when offered a conspecific and a similarly coloured opponent simultaneously, there were no differences in behaviour towards either opponent. Thus, nuptial coloration is used by males to identify competitors, and it suggests that male-male aggression may have also been an important diversifying force in speciation in rock-dwelling Lake Malawi cichlids.     

Genetic relationships of cichlid fishes from Lake Malawi based on mitochondrial DNA sequences

Limnology - The purpose of this study was to estimate the genetic distance, diversity and relationship of two large groups of Malawian cichlid, ‘mbuna’ living in rocky area and feeding...     

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.     

A small number of genes underlie male pigmentation traits in Lake Malawi cichlid fishes

Pigmentation patterns are one of the most recognizable forms of phenotypic diversity and an important component of organismal fitness. While much progress has been made in understanding the genes controlling pigmentation in model systems, many questions remain about the genetic basis of pigment traits observed in nature. Lake Malawi cichlid fishes are known for their diversity of male pigmentation patterns, which have been shaped by sexual selection. To begin the process of identifying the genes underlying this diversity, we quantified the number of pigment cells on the body and fins of two species of the genus Metriaclima and their hybrids. We then used the Castle-Wright equation to estimate that differences in individual pigmentation traits between these species are controlled by one to four genes each. Different pigmentation traits are highly correlated in the F(2) , suggesting shared developmental pathways and genetic pleiotropy. Melanophore and xanthophore traits fall on opposite ends of the first principal component axis of the F(2) phenotypes, suggesting a tradeoff during the development of these two pigment cell types.     

A population genetic assessment of taxonomic species: The case of Lake Malawi cichlid fishes

Organisms sampled for population‐level research are typically assigned to species by morphological criteria. However, if those criteria are limited to one sex or life stage, or the organisms come from a complex of closely related forms, the species assignments may misdirect analyses. The impact of such sampling can be assessed from the correspondence of genetic clusters, identified only from patterns of genetic variation, to the species identified using only phenotypic criteria. We undertook this protocol with the rock‐dwelling mbuna cichlids of Lake Malawi, for which species within genera are usually identified using adult male coloration patterns. Given high local endemism of male colour patterns, and considerable allele sharing among species, there persists considerable taxonomic uncertainty in these fishes. Over 700 individuals from a single transect were photographed, genotyped and separately assigned: (a) to morphospecies using photographs; and (b) to genetic clusters using five widely used methods. Overall, the correspondence between clustering methods was strong for larger clusters, but methods varied widely in estimated number of clusters. The correspondence between morphospecies and genetic clusters was also strong for larger clusters, as well as some smaller clusters for some methods. These analyses generally affirm (a) adult male‐limited sampling and (b) the taxonomic status of Lake Malawi mbuna, as the species in our study largely appear to be well‐demarcated genetic entities. More generally, our analyses highlight the challenges for clustering methods when the number of populations is unknown, especially in cases of highly uneven sample sizes.     

Reproductive isolation among deep-water cichlid fishes of Lake Malawi differing in monochromatic male breeding dress

Male nuptial colour hues are important for the maintenance of reproductive isolation among cichlid fish species, and environmental changes that lead to narrower light spectra can lead to hybridization. However, cichlid species can naturally co-occur in narrow light spectrum habitats, such as turbid shallow lakes and the deep benthic zones of African rift lakes. Closely related species from narrow light spectrum habitats tend to differ little in the palette of male nuptial colours, thus for these taxa differences in colour patterns may be more important than differences in colour hue for species recognition. To investigate this hypothesis we examined morphometric and genetic differentiation among males of four sympatric putative species within the deep-water genus Diplotaxodon. These taxa live in a narrow-light spectrum environment where only blue light is present, and males differ primarily in 'monochromatic' black, white and silver patterning of the body and fins. Significant genetic differentiation was present among taxa in both microsatellite DNA and mitochondrial DNA, including one pair with no significant morphometric differentiation. Thus, these taxa represent reproductively isolated biological species, a result consistent with male nuptial patterning being important for species recognition and assortative mating. As such, we suggest that narrow-light spectra need not always represent barriers to effective visually mediated mate recognition.     

Beta diversity of rock-restricted cichlid fishes in Lake Malawi: importance of environmental and spatial factors

The rock-restricted cichlid fish assemblages of Lake Malawi exhibit high spatial diversity in their species composition and relative abundance. However the extent to which this is due to the effects of local environmental differences, dispersal limitation of constituent taxa, and the assignment of allopatric populations to species is uncertain. We examined the factors associated with diversity within an assemblage from the north-western shores, encompassing a spatial scale of 170 km. For both the whole assemblage, and all constituent species-complexes, spatial variance in community structure was significantly dependent upon both geographic distances between locations and local habitat variables. Pronounced effects of distance indicate limited dispersal, but our results also show that that the spatial variance explained by geographic distance alone was strongly linked to proportion of allopatric populations within a species-complex with species status. Thus, the taxonomic status of allopatric populations underlies, at least partially, the biogeographical structure of this assemblage. Substrate composition and habitat depth were also significant determinants of community structure, although spatial variance attributed to these variables was less than that associated with distance alone. Substantial unexplained variance may be a consequence of the effects of unmeasured habitat variables, high ecological similarity between co-occurring species, stochastic influences on population abundance, and the effects of local adaptation. Despite low spatial variance explained by the assessed environmental variables, significant environmental influence on cichlid assemblage structure across a wide spatial scale indicates that even slight future environmental changes may have the capacity to significantly alter species composition.     

Assortative mating among rock-dwelling cichlid fishes supports high estimates of species richness from Lake Malawi

It has been estimated that Lake Malawi, Africa, contains 500–650 endemic species of cichlid fishes, the largest number of vertebrate species endemic to any comparable sized area on the planet. As many of these putative species cannot be distinguished anatomically, these estimates of species richness depend to a great extent on the assumption that sympatrically occurring male colour morphs represent biological species. We have tested this assumption using a combination of behavioural observations of courtship and microsatellite DNA analysis for six putative species of the Pseudotropheus ( Tropheops ) complex and three of the Pseudotropheus ( Maylandia ) complex occurring sympatrically at Nkhata Bay. We were unable to demonstrate assortative courtship for the species pairs Pseudotropheus ( Maylandia ) zebra / P . 'gold zebra' or P. ( Tropheops ) 'band'/ P. ( T .) 'rust' because we were unable to distinguish between the females of these taxa. All other taxa showed clear assortative courtship, except for P. ( T .) 'deep', a deep-water species which was rarely observed. Fixation indices (θ_ST for the infinite allele model, and R _ST for the stepwise mutation model) calculated from six microsatellite DNA loci demonstrated significant deviations from panmixia in all pairwise comparisons of putative species, indicating little or no gene flow between populations. All taxa showed high levels of allelic diversity providing evidence that genetic bottlenecking may have been of limited importance in the speciation process. Assortative mating among taxa differing only in male colouration is supportive of theories that speciation in these fishes has been driven by sexual selection by female choice.     

Polygynandry inPseudotropheus zebra, a cichlid fish from Lake Malawi

Synopsis Parental care in the Malawian cichlid fishPseudotropheus zebra BB is extensive and exclusively maternal; males contribute only genetic material. The costs of searching for multiple mates (in this case risk of predation on orally incubated eggs) suggested that females should be monandrous; microsatellite genetypes of seven brooding females and their young, however, reveal extensive multiple paternity in this species, with a mean of 3.8 paternal individuals per brood. Polygynandry inP. zebra is probably not maintained by selection for genetically diverse offspring; potential explanations include avoidance of inbreeding, and bet-hedging on other male characteristics that females are unable to evaluate when selecting a mate. The observed degree of multiple paternity strongly suggests that females are free to choose mates as they will, a prerequisite of many theories positing sexual selection as a key element in Malawi chichlid evolution. It should also result in elevation of effective population sizes, and thus be antagonistic to runaway evolution of male secondary sexual characteristics, but not necessarily to other modes of sexual selection.     

Genetic interactions controlling sex and color establish the potential for sexual conflict in Lake Malawi cichlid fishes

Sex-determining systems may evolve rapidly and contribute to lineage diversification. In fact, recent work has suggested an integral role of sex chromosome evolution in models of speciation. We use quantitative trait loci analysis of restriction site-associated DNA -tag single nucleotide polymorphisms to identify multiple loci responsible for sex determination and reproductively adaptive color phenotypes in Lake Malawi cichlids. We detect a complex epistatic sex system consisting of a major female heterogametic ZW locus on chromosome 5, two separate male heterogametic XY loci on chromosome 7, and two additional interacting loci on chromosomes 3 and 20. Our data support the known chromosomal linkage between orange blotch color and ZW, as well as novel genetic associations between male blue nuptial color and two sex determining regions (an XY and ZW locus). These results provide further empirical evidence for a complex antagonistic sex–color system in this species flock and suggest a possible role for, and effect of, polygenic sex-determining systems in rapid evolutionary diversification.     

Evolutionary history of Lake Tanganyika's scale-eating cichlid fishes

Although Lake Tanganyika is not the most species-rich of the Great East African Lakes it comprises by far the greatest diversity of cichlid fishes in terms of morphology, ecology, and breeding styles. Our study focuses on the Tanganyikan cichlid tribe Perissodini, which exhibits one of the most peculiar feeding strategies found in cichlids-scale-eating. Their evolutionary history was reconstructed from 1416 bp DNA sequence of two mitochondrial genes (ND2 and partial control region) and from 612 AFLP markers. We confirm the inclusion of the zooplanktivorous genus Haplotaxodon in the tribe Perissodini, and species status of Haplotaxodon trifasciatus. Within the Perissodini, the major lineages emerged within a short period roughly 1.5-2 MYA, which makes their radiation slightly younger than that of other Tanganyikan cichlid tribes. Most scale-eaters evolved in deep-water habitat, perhaps associated with the previously documented radiations of other deep-water dwelling cichlid lineages, and colonized the shallow habitat only recently.     

The feeding habits of a group of herbivorous rock-dwelling cichlid fishes (Cichlidae: Perciformes) from Lake Malawi,Africa

Synopsis The feeding; habits of a group of tropical herbivorous rock-dwelling cichlid fishes from Lake Malawi, Africa, are investigated using stomach content analyses. The various species fed selectively on the periphyton of the rocky shores. Blue-green alga of the genus Calothrix was the most common item ingested by the group. Diatoms (Chrysophyta) also were abundant food items. Discriminant analysis showed that dietary items were good variables to identify species. Interspecific dietary differences showed a continuum from those species feeding primarily on Calothrix to those feeding primarily on diatoms. Algal resources exhibit distinct patterns of spatial variation. Diet was correlated with foraging behavior and trophic morphology. Interspecific differences in diet could possibly facilitate ecological coexistence among various species. Such coexistence would contribute to the maintenance of the high diversity fish faunas characteristic of the Great Rift Lakes of Africa.     

Unusually fine-scale genetic structuring found in rapidly speciating Malawi cichlid fishes

Mechanisms behind the explosive radiation of over 500 cichlid fish species from a single founding population in Lake Malawi during the last 700 000 years are poorly understood. Recent studies have suggested that the degree of population subdivision among the habitat patches within the lake may be responsible, but the evidence has been circumstantial: lack of a dispersal stage in haplochromine cichlids; genetic and colour variation among populations separated by large-scale geographical barriers; and fluctuating lake levels. One reason for the rapidity of speciation in these fishes may be that population subdivision is on a much finer scale than previously thought. Here we quantify the level of population subdivision and estimate migration at a scale of 700-1400 m, in order to investigate whether cichlid populations are sufficiently isolated from each other for allopatric divergence and perhaps speciation to take place. Using six microsatellite loci, we demonstrate the existence of highly significant genetic differentiation between subpopulations on adjacent headlands in each of four rock-dwelling haplochromine cichlid species. Our results suggest that these fish populations are divided into thousands of subunits among which genetic divergence is currently occurring, and that this may provide unprecedented opportunities for allopatric speciation. <br>     

Alternative life-history styles of some African cichlid fishes

Synopsis About 50% of African cichlid species have been described. Little is known about cichlid ecology, behaviour or about the evolution and interactions of communities. Nevertheless, trends which provide an insight into cichlid life histories, into evolutionary alternatives and into behavioural alternatives which may be followed during the life time of these fishes are emerging. Cichlids which spend their entire life history in a single habitat belong to species flocks that have spectated greatly. In contrast, those groups which live in a variety of habitats during their life history have spectated little. Despite the trophic specializations that have occurred among cichlids, many and perhaps all species, have the ability to feed upon alternative food resources. They do so by switching or by markedly modifying their behaviour and all are particularly opportunistic. The extensive adaptive radiation of cichlids with regard to those attributes of morphology and behaviour that are associated with survival and growth, is apparently not matched by similar radiations in reproductive behaviour. The perceived evolutionary conservatism with regard to reproductive behaviour is attributed to the effects of stabilizing selection, but this still needs to be tested. The constraints of stabilizing selection appear to be lifted with regard to coloration. If colour is an important component of the specific-mate recognition system then it should also be subjected to stabilizing selection so the readiness with which colour variation occurs within and between populations needs explanation. Parental care in cichlids has followed a variety of evolutionary alternatives, all of which are geared to improve the chances of survival of offspring in their specific micro-habitats.     

Age and spread of the haplochromine cichlid fishes in Africa

The Haplochromini are by far the most species-rich cichlid fish tribe that originated along with the so-called primary radiation of the Lake Tanganyika cichlid species flock, i.e. at the same time during which the majority of the endemic Lake Tanganyika cichlid tribes emerged. Unlike the other tribes, the haplochromines are not restricted to Lake Tanganyika but distributed throughout Africa, except for the northwestern part of the continent. Haplochromine cichlids seeded the adaptive radiation of cichlid fishes in Lakes Malawi, Kivu, Victoria, Turkana, as well as in the now extinct paleo-Lake Makgadikgadi. Here we present a comprehensive phylogenetic and phylogeographic analysis of haplochromine cichlids that is based upon DNA sequences of two mitochondrial gene segments of riverine taxa covering all major African biogeographic regions where haplochromines are found. Our analysis revealed that six lineages of haplochromines originated within a short period of time, about 5.3-4.4 MYA. These haplochromine lineages show a highly complex phylogeographic pattern, probably severely influenced by climate- and/or geology-induced changes of the environment, with river capture events most likely playing an important role for species dispersal.