Lake Tanganyika harbours the oldest and ecologically, morphologically and behaviourally most diverse species flock of cichlid fishes. Its species are excellent subjects for the study of explosive speciation and adaptive radiation. Many species are subdivided into numerous genetically and phenotypically distinct populations, often classified as distinct geographical races or colour morphs, which mostly differ in colour and much less in terms of morphology. This study for the first time quantifies morphological differences among such morphs by studying three populations of Tropheus moorii. We compared Ôtraditional morphometricsÕ (TM) and Ôgeometric morphometricsÕ (GM) to explore their potential for discriminating populations. So far species description and population discrimination are almost solely based on TM in the form of standardized measurements, although specialists are aware of their lack of diagnostic power for discrimination of closely related entities. Moreover, comprehensive TM measurements are time consuming and can best be done on dead specimens which have to be preserved in the case re-measuring is necessary. In contrast, GM can also be based on photographs and computer scans of anaesthetized fish, so that the same individual can be repeatedly analysed during its ontogeny. Here, we show that GM is more flexible in data acquisition and more powerful in the discrimination of species and closely related populations. While TM is restricted to distances and ratios of distances, GM not only includes these measurements indirectly, but also allows for body shape analysis using a semi-landmark approach. It can be equally standardized as TM by defining diagnostic landmarks. Data description by canonical variate analysis was most informative using GM data including semi-landmarks, whereas differences between populations were significant (p < 0.05) based on both morphological approaches.
The abundance of two main pelagic fish species in Lake Tanganyika (Stolothrissa tanganicae and Lates stappersii) has always been observed to fluctuate considerably at different time scales. The inverse correlation between the abundance of these species has often been interpreted as the consequence of predator-prey relations (avoidance behaviour by the prey). However, currently the two species often appear spatially segregated in the lake, S. tanganicae dominating in the north while L. stappersii is generally abundant in the south where it feeds mostly on shrimps. A fluctuating abundance of the species is nevertheless observed. As these fish species have a major importance for the fisheries, we investigated the limnological variability in relation to the shortterm variability of fish catches. The abundance of S. tanganicae was positively correlated to plankton biomass (r = 0.65), while water transparency (r = 0.56), depth of mixed layer (r = -0.70) and oxygenated water appeared important drivers for the abundance of L. stappersii. Alternating ''mixing'' and ''stable'' states of the epilimnion related to seasonal and internal waves variability are probably determinant for the short-term variability in abundance of Handling editor: M. Power
Haplochromine cichlids form the most species-rich lineage of cichlid fishes that both colonized almost all river systems in Africa and radiated to species flocks in several East African lakes. The enormous diversity of lakes is contrasted by a relatively poor albeit biogeographically clearly structured species diversity in rivers. The present study analyzed the genetic structure and phylogeographic history of species and populations of the genus Pseudocrenilabrus in Zambian rivers that span two major African drainage systems, the Congo-and the Zambezi-system. The mtDNA phylogeny identifies four major lineages, three of which occur in the Congo-system and one in the Zambezi system. Two of the Congo-clades (Lake Mweru and Lunzua River) comprise distinct albeit yet undescribed species, while the fish of the third Congo-drainage clade (Chambeshi River and Bangweulu swamps), together with the fish of the Zambezi clade (Zambezi and Kafue River) are assigned to Pseudocrenilabrus philander. Concerning the intraspecific genetic diversity observed in the sampled rivers, most populations are highly uniform in comparison to lacustrine haplochromines, suggesting severe founder effects and/or bottlenecking during their history. Two bursts of diversification are reflected in the structure of the linearized tree. The first locates at about 3.9% mean sequence divergence and points to an almost simultaneous colonization of the sampled river systems. Subsequent regional diversification (with about 1% mean sequence divergence) occurred contemporaneously within the Kafue River and the Zambezi River. The clear-cut genetic biogeographic structure points to the dominance of geographic speciation in this lineage of riverine cichlid fishes, contrasting the importance of in situ diversification observed in lake cichlids.
Tetracycline was used as a chemical tag in a mark-recapture study to examine the pattern of increment formation in the otoliths of Tropheus moorii, a rock-dwelling cichlid from Lake Tanganyika. A total of 256 fish were captured by divers and injected with tetracycline. Of these, nine were recaptured after either 1 or 2 years at liberty and eight retained tags within their otoliths. Comparison of the number of growth increments formed after the tag and the time at liberty demonstrated that increments were deposited on an annual basis in the otoliths of this species. Furthermore, there was a strong relationship between otolith mass and age suggesting that otoliths grew at a predictable rate throughout the life of the fish. Validation of an annual pattern of increment deposition allowed age and growth information to be derived from otoliths. This showed that T. moorii grew rapidly to attain adult size by 3 years of age. Males grew faster than females and also attained a larger size than females (8Á74 v. 7Á91 cm L S respectively). The longevity of some of these small freshwater fish was surprising; the oldest individual had an age of 10 years, while the average age of adults was 4 years. # 2004 The Fisheries Society of the British Isles
In the extremely species-rich haplochromine cichlid fishes of the East African Great Lakes, prezygotic isolation between closely related species is often maintained by color-assortative mating. In 1998, local fisherman working for the ornamental fish trade released different color morphs of the cichlid genus Tropheus into a small harbor basin in the southern part of Lake Tanganyika. This artificial amalgamation of color morphs provides a unique possibility to study mating patterns in cichlids in a natural environment over time. In a precursor study, we analyzed genotypes and phenotypes of almost 500 individuals sampled between 1999 and 2001 and uncovered a marked degree of color-assortative mating, which depended on the level of color pattern dissimilarity between morphs. Twelve years after introduction of nonindigenous morphs, we again sampled Tropheus individuals from the harbor basin and an adjacent, originally pure population and analyzed phenotypes (coloration) and genotypes (mitochondrial control region and 9 microsatellite loci) to assess the current status of the admixed population. Principal component analyses of color score data and population assignment tests demonstrate an increasing level of introgressive hybridization between morphs but also some ongoing color-assortative mating within morphs. The observed mating pattern might have been influenced by fluctuating environmental conditions such as periodic algal blooms or increased sedimentation causing turbid conditions in an otherwise clear lake.
Lake Tanganyika comprises the oldest and most diverse species flock of cichlid fishes. Many species are subdivided into numerous populations, often classified as geographical races, colour morphs or sister species. Unlike younger species flocks, in which speciation is accompanied by ecomorphological diversification that of Lake Tanganyika is at a mature stage with little further morphological change, most probably caused by stabilizing selection. This study addresses body shape differences among three genetically distinct but morphologically similar populations of Tropheus moorii, by focusing on bony structures of the cichlid head. We test by means of geometric morphological methods whether shape changes in the cichlid head are based on particular osteological differences. Specimens were disarticulated enzymatically, and standardized digital images of the disarticulated bony elements were taken. A landmark system was established for the dentary, articular, premaxilla, quadrate and the preopercle. Only the dentary shows significant differentiation among the three populations. Since all three populations live in similar cobble habitats and occupy the same trophic niche, the observed difference in the shape of the dentary can either be explained by different directional selection due to subtle habitat differences, or by neutral drift constrained by borders enforced through stabilizing selection. Lack of difference might indicate stabilizing selection on bone shape.
Stenotopic specialization to a fragmented habitat promotes the evolution of genetic structure. It is not yet clear whether small-scale population structure generally translates into large-scale intraspecific divergence. In the present survey of mitochondrial genetic structure in the Lake Tanganyika endemic Altolamprologus (Teleostei, Cichlidae), a rock-dwelling cichlid genus comprising A. compressiceps and A. calvus, habitat-induced population fragmentation contrasts with weak phylogeographic structure and recent divergence among genetic clades. Low rates of dispersal, perhaps along gastropod shell beds that connect patches of rocky habitat, and periodic secondary contact during lake level fluctuations are apparently sufficient to maintain genetic connectivity within each of the two Altolamprologus species. The picture of genetic cohesion was interrupted by a single highly divergent haplotype clade in A. compressiceps restricted to the northern part of the lake. Comparisons between mitochondrial and nuclear phylogenetic reconstructions suggested that the divergent mitochondrial clade originated from ancient interspecific introgression. Finally, 'isolation-with-migration' models indicated that divergence between the two Altolamprologus species was recent (67-142 KYA) and proceeded with little if any gene flow. As in other rock-dwelling cichlids, recent population expansions were inferred in both Altolamprologus species, which may be connected with drastic lake level fluctuations.
Group-living animals are often faced with complex reproductive decisions, namely how to partition within-group reproduction, how to obtain extra-group reproduction and how these two means of reproduction should be balanced. The solutions to these questions can be difficult to predict because ecological conditions can affect the scopes for within-group and extra-group reproduction in complex ways. For example, individuals that are restricted from moving freely around their habitats may have limited extra-group reproductive opportunities, but at the same time, groups may live in close proximity to one another, which could potentially have the opposite effect.The group-living cichlid fish Neolamprologus multifasciatus experiences such ecological conditions, and we conducted an intensive genetic parentage analysis to investigate how reproduction is distributed within and among groups for both males and females. We found that cohabiting males live in "high-skew" societies, where dominant males monopolize the majority of within-group reproduction, while females live in "low-skew" societies, where multiple females can produce offspring concurrently.Despite extremely short distances separating groups, we inferred only very low levels of extra-group reproduction, suggesting that subordinate males have very limited reproductive opportunities. A strength of our parentage analysis lies in its inclusion of individuals that spanned a wide age range, from young fry to adults. We outline the logistical circumstances when very young offspring may not always be accessible to parentage researchers, and present strategies to overcome the challenges of inferring mating patterns from a wide age range of offspring.
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