BackgroundWith about 120 colour morphs currently assigned to six nominal species, the genus Tropheus is an ideal model to study evolutionary divergence of populations in allopatry. The morphology of Tropheus has been described as relatively static, but reproductive constraints are sexually dimorphic due to mouthbrooding in females. We analysed phenotypic variation in six populations of T. moorii and one population of T. polli using geometric morphometrics to assess morphological differences among sexes in relation to the differentiation of populations and species.ResultsThe mean shapes differed significantly between sexes, populations, and species even though within-sex variation exceeded the divergence among populations. The first principal component of Procrustes shape coordinates revealed differences between populations and species in mouth position and ventral head shape. The second principal component reflected sex-specific shape differences, mainly comprising a relatively larger female viscerocranium and, in particular, a larger buccal area. While shape variation between populations and between sexes was primarily located in the cranial region, within-sex variation was relatively uniform across all landmarks.ConclusionsDeviations of the between-population and between-sex pattern of shape variation from that within sex indicate that the differences in head shape likely result from both adaptations to female mouthbrooding and population-specific foraging strategies.

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.

Cichlids are an excellent model to study explosive speciation and adaptive radiation. Their evolutionary success has been attributed to their ability to undergo rapid morphological changes related to diet, and their particular breeding biology. Relatively minor changes in morphology allow for exploitation of novel food resources. The importance of phenotypic plasticity and genetically based differences for diversification was long recognized, but their relationship and relative magnitude remained unclear. We compared morphology of individuals of four wild populations of the Lake Tanganyika cichlid Tropheus moorii with their pond-raised F1 offspring. The magnitude of morphological change via phenotypic plasticity between wild and pond-bred F1 fish exceeds pairwise population differences by a factor of 2.4 (mean Mahalanobis distances). The genetic and environmental effects responsible for among population differentiation in the wild could still be recognized in the pond-bred F1 fish. All four pond populations showed the same trends in morphological change, mainly in mouth orientation, size and orientation of fins, and thickness of the caudal peduncle. As between population differentiation was lower in the wild than differentiation between pond-raised versus wild fish, we suggest the narrow ecological niche and intense interspecific competition in rock habitats is responsible for consistent shape similarity, even among long-term isolated populations.Electronic supplementary materialThe online version of this article (doi:10.1007/s00114-010-0751-2) contains supplementary material, which is available to authorized users.

Lake Tanganyika harbors the ecologically, morphologically, and behaviorally most diverse species flock of cichlid fishes. It is comprised by substrate breeding and mouthbrooding species, most of which live in littoral habitats. Species communities are characterized by complex behavioral and trophic interactions, resulting in a dense pattern of partially overlapping territories, depending on the degree of ecological distinctness. We studied territorial behavior of breeding pairs in a substrate breeding species, with respect to territory size and defense behavior. The study species Variabilichromis moorii belongs to the tribe Lamprologini, the most species rich tribe of cichlids in Lake Tanganyika. Our study shows that breeding pairs of V. moorii can have highly complex territories, in which both parents hold separate subterritories which are shifted slightly according to the movements of the fry, but the outer borders are conjointly defended. The size of the total defended territory varied from \1 to almost 4 m 2 , averaging at about 2 m 2 . Depending on presence of competitors or fry-predators evoking agonistic interactions, the territory size varied quite substantially over the day. Attack rates and size of the defended area decreased with water depth. Agonistic behavior was observed toward heterospecifics as well as conspecifics, with heterospecific attacks mostly concerning territorial neighbors and potential fry-predators in about equal frequencies.

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.