Sexually divergent selection, allometric constraints, and the evolution of sexual dimorphism in cichlids from Lake Tanganyika.

The evolution of sexual dimorphism is widely acknowledged as manifestation of sex-specific genetic architecture. Although empirical studies suggested that sexual dimorphism evolves as a joint consequence of constraints arising from the genetic architecture and sexually divergent selection, it remains unclear whether and how these established microevolutionary processes scale up to the macroevolutionary patterns of sexual dimorphism among taxa. Here, we studied how sexual selection and parental care drive sexual dimorphism in cichlid fishes from Lake Tanganyika. We found that male-male competition, female choice, and maternal mouthbrooding are associated with sexual dimorphism in body length, body color, and head length, respectively, despite strong allometric relationships between body length and head length. Within-species (static) allometry of head length on body length evolved as sex-specific responses to mouthbrooding where females evolved higher intercepts while males evolved steeper slopes. Thus, selection to increase mouth size in mouthbrooders may have broken down and reorganized the pattern of allometric constraints that are inherently strong and concordant between sexes. Furthermore, sex-specific responses to mouthbrooding left a remarkably clear signature on the macroevolutionary pattern, resulting in a decoupling of co-evolution in parameters of static allometries observed exclusively within maternal mouthbrooders. Our study provides multiple lines of evidence that are consistent with the idea that macroevolutionary patterns of sexual dimorphism in Lake Tanganyika cichlids result from sexually divergent selection. Our approach illustrates that an examination of within-population phenotypic variance in the phylogenetic comparative framework may facilitate nuanced understandings of how macroevolutionary patterns are generated by underlying microevolutionary processes.

Genomic Microevolution of Vibrio cholerae O1, Lake Tanganyika Basin, Africa.

Africa's Lake Tanganyika basin is a cholera hotspot. During 2001-2020, Vibrio cholerae O1 isolates obtained from the Democratic Republic of the Congo side of the lake belonged to 2 of the 5 clades of the AFR10 sublineage. One clade became predominant after acquiring a parC mutation that decreased susceptibility to ciprofloxacin.

Sustainable Management of the African Great Lake Coastal Areas: Motivations and Perspectives of Community Citizen Scientists.

The long-term sustainability of the African Great Lakes is strongly connected to the management and monitoring of their coastal areas. Yet, the communities that live in these areas are rarely involved in monitoring and have limited influence on key management issues. Furthermore, regulatory activities and knowledge sharing in these transnational ecosystems are strongly limited by funding and infrastructure limitations. Citizen science has great potential to advance both scientific and public understanding of the state of the environment. However, there remains a limited understanding of participants' motivations and expectations, especially in developing countries, where citizen science has great potential to complement regulatory monitoring. The present study explores the motivations of citizen scientists in villages along Lake Tanganyika's northern coast and their potential to take a more active role in lake management. Motivations were examined through qualitative interviews, focus groups, and quantitative surveys with 110 citizen scientists and 110 non-citizen scientists from participating villages. Key motivational factors identified were the desire to contribute to scientific research and local knowledge, as well as aspects of financial compensation. The results confirm that participation in citizen science provides many benefits to participants beyond their role as data aggregators and final knowledge users. However, the incentives to participation varied to those typically considered in citizen science programs conducted in developed countries. To create sustainable long-term community based environmental monitoring, these motivations should be incorporated in the program design and participant recruitment.

Explosive networking: The role of adaptive host radiations and ecological opportunity in a species-rich host-parasite assembly.

Many species-rich ecological communities emerge from adaptive radiation events. Yet the effects of adaptive radiation on community assembly remain poorly understood. Here, we explore the well-documented radiations of African cichlid fishes and their interactions with the flatworm gill parasites Cichlidogyrus spp., including 10,529 reported infections and 477 different host-parasite combinations collected through a survey of peer-reviewed literature. We assess how evolutionary, ecological, and morphological parameters determine host-parasite meta-communities affected by adaptive radiation events through network metrics, host repertoire measures, and network link prediction. The hosts' evolutionary history mostly determined host repertoires of the parasites. Ecological and evolutionary parameters predicted host-parasite interactions. Generally, ecological opportunity and fitting have shaped cichlid-Cichlidogyrus meta-communities suggesting an invasive potential for hosts used in aquaculture. Meta-communities affected by adaptive radiations are increasingly specialised with higher environmental stability. These trends should be verified across other systems to infer generalities in the evolution of species-rich host-parasite networks.

Variable hybridization between two Lake Tanganyikan cichlid species in recent secondary contact.

Closely related taxa frequently exist in sympatry before the evolution of robust reproductive barriers, which can lead to substantial gene flow. Post-divergence gene flow can promote several disparate trajectories of divergence ranging from the erosion of distinctiveness and eventual collapse of the taxa to the strengthening of reproductive isolation. Among many relevant factors, understanding the demographic history of divergence (e.g. divergence time and extent of historical gene flow) can be particularly informative when examining contemporary gene flow between closely related taxa because this history can influence gene flow's prevalence and consequences. Here, we used genotyping-by-sequencing data to investigate speciation and contemporary hybridization in two closely related and sympatrically distributed Lake Tanganyikan cichlid species in the genus Petrochromis. Demographic modelling supported a speciation scenario involving divergence in isolation followed by secondary contact with bidirectional gene flow. Further investigation of this recent gene flow found evidence of ongoing hybridization between the species that varied in extent between different co-occurring populations. Relationships between abundance and the degree of admixture across populations suggest that the availability of conspecific mates may influence patterns of hybridization. These results, together with the observation that sets of recently diverged cichlid taxa are generally geographically separated in the lake, suggest that ongoing speciation in Lake Tanganyikan cichlids relies on initial spatial isolation. Additionally, the spatial heterogeneity of admixture between the Petrochromis species illustrates the complexities of hybridization when species are in recent secondary contact.

Material gradients in gastropod radulae and their biomechanical significance: a combined approach on the paludomid Lavigeria grandis.

The radula, a chitinous membrane spiked with teeth, is the molluscan autapomorphy for the gathering and processing of food. The teeth, as actual interfaces between the organism and the ingesta, act as load transmitting regions and have to withstand high stresses during foraging - without structural failure or high degrees of wear. Mechanisms contributing to this were studied previously in paludomid gastropods from Lake Tanganyika. For some species, gradients in hardness and Young's modulus along the teeth were detected, enabling the bending and relying of teeth onto the next row, distributing the stresses more equally. The here presented study on one of them - Lavigeria grandis - aims at shedding light on the origin of these functional gradients. The mechanical properties were identified by nanoindentation technique and compared to the elemental composition, determined by elemental dispersive X-ray spectroscopy (EDX, EDS). This was done for the complete radular (mature and immature tooth rows), resulting in overall 236 EDX and 700 nanoindentation measurements. Even though teeth showed regional differences in elemental composition, we could not correlate the mechanical gradients with the elemental proportions. By applying confocal laser scanning microscopy (CLSM), we were finally able to relate the mechanical properties with the degree of tanning. CLSM is a common technique used on arthropod cuticle, but was never applied on radular teeth before. In general, we found that nanoindentation and CLSM techniques complement one another, as for example, CLSM is capable of revealing heterogeneities in material or micro-gradients, which leads to a better understanding of the functionalities of biological materials and structures.

Micro-cracks and micro-fractures reveal radular tooth architecture and its functional significance in the paludomid gastropod Lavigeria grandis.

Most molluscan taxa forage with their radula, a chitinous membrane with embedded teeth. The teeth are the actual interfaces between the animal and its ingesta and serve as load-transmitting regions. During foraging, these structures have to withstand high stresses without structural failure and without a high degree of wear. Mechanisms contributing to this failure- and wear-resistance were well studied in the heavily mineralized teeth of Polyplacophora and Patellogastropoda, but for the rather chitinous teeth of non-limpet snails, we are confronted with a large gap in data. The work presented here on the paludomid gastropod Lavigeria grandis aims to shed some light on radular tooth composition and its contribution to failure- and wear-prevention in this type of radula. The teeth were fractured and the micro-cracks studied in detail by scanning electron microscopy, revealing layers within the teeth. Two layers of distinct fibre densities and orientations were detected, covered by a thin layer containing high proportions of calcium and silicon, as determined by elemental dispersive X-ray spectroscopy. Our results clearly demonstrate the presence of failure- and wear-prevention mechanisms in snail radulae without the involvement of heavy mineralization-rendering this an example of a highly functional biological lightweight structure. This article is part of the theme issue 'Nanocracks in nature and industry'.

Mitochondrial genetic diversity and divergence dating of Angolan colobus monkeys (Colobus angolensis) in the eastern forests of Kenya and Tanzania: Implications for subspeciation and reconstructing historical biogeography.

Whether the Colobus angolensis that reside in the fragmented forests in eastern Kenya and Tanzania represent one subspecies or two has been debated for 50 years. Morphological and more recent genetic and ecological studies suggest that these populations represent two subspecies, C. a. palliatus and C. a. sharpei. However, their distribution of mitochondrial variation remains unresolved since the genetic study only characterized four populations at the range ends. Therefore, we characterized five populations in the area of the hypothesized subspecies divide. We identified eight new haplotypes which, combined with those previously identified, provided 26 haplotypes from nine populations for analysis. Haplotypes found south of the Rufiji River cluster together but separately from northern haplotypes. The largest sequence differences within cytochrome b occur between population pairs representing opposite sides of the river; their mean difference (1.5%) is more than that of other primate subspecies. Analysis of molecular variance attributes most of the variation to that north versus south of the river. These results support the previous subspecies distinction between C. a. palliatus (northern) and C. a. sharpei (southern), divided by the Rufiji River. The estimated time of the most recent common ancestor of all haplotypes indicates that the subspecies have been isolated from each other for approximately 550,000 years. The common ancestor of northern and southern haplogroups was 370,000 and 290,000 years ago, respectively. Nevertheless, the correlation between genetic and geographic distances suggests that isolation-by-distance contributed to population structuring. Significant variation among populations, with only three haplotypes shared between populations, also indicates that an extended period of isolation drove population distinctiveness. Considering these results, we evaluate hypotheses about the founding and differentiation of these subspecies during Pleistocene climatic fluctuations and propose a novel, more direct migration route from Central Africa to their current range navigating Lake Tanganyika, the central Tanzanian corridor, and the Rufiji River.

Citizen scientist monitoring accurately reveals nutrient pollution dynamics in Lake Tanganyika coastal waters.

Several studies in Lake Tanganyika have effectively employed traditional methods to explore changes in water quality in open waters; however, coastal monitoring has been restricted and sporadic, relying on costly sample and analytical methods that require skilled technical staff. This study aims in validating citizen science water quality collected data (nitrate, phosphate and turbidity) with those collected and measured by professional scientists in the laboratory. A second objective of the study is to use citizen scientist data to identify the patterns of seasonal and spatial variations in nutrient conditions and forecast potential changes based on expected changes in population and climate (to 2050). The results showed that the concentrations of nitrate and phosphate measured by citizen scientists nearly matched those established by professional scientists, with overall accuracy of 91% and 74%, respectively. For total suspended solids measured by professional and turbidity measured by citizen scientists, results show that, using 14 NTU as a cut-off, citizen scientist measurements of Secchi tube depth to identify lake TSS below 7.0 mg/L showed an accuracy of 88%. In both laboratory and citizen scientist-based studies, all measured water quality variables were significantly higher during the wet season compared to the dry season. Climate factors were discovered to have a major impact on the likelihood of exceeding water quality restrictions in the next decades (2050), which could deteriorate lake conditions. Upscaling citizen science to more communities on the lake and other African Great Lakes would raise environmental awareness, inform management and mitigation activities, and aid long-term decision-making.

Mapping of quantitative trait loci underlying a magic trait in ongoing ecological speciation.

BACKGROUND: Telmatochromis temporalis is a cichlid fish endemic to Lake Tanganyika. The normal and dwarf morphs of this fish are a clear example of ongoing ecological speciation, and body size plays an important role in this speciation event as a magic trait. However, the genetic basis underlying this trait has not been studied. RESULTS: Based on double-digested restriction-site associated DNA (ddRAD) sequencing of a hybrid cross between the morphs that includes F0 male, F0 female, and 206 F2 individuals, we obtained a linkage map consisting of 708 ddRAD markers in 22 linkage groups, which corresponded to the previously reported Oreochromis niloticus chromosomes, and identified one significant and five suggestive quantitative trait loci (QTL) for body size. From the body-size distribution pattern, the significant and three of the five suggestive QTL are possibly associated with genes responsible for the difference in body size between the morphs. CONCLUSIONS: The QTL analysis presented here suggests that multiple genes, rather than a single gene, control morph-specific body size. The present results provide further insights about the genes underlying the morph specific body size and evolution of the magic trait during ecological speciation.

Transcriptomics unravels molecular players shaping dorsal lip hypertrophy in the vacuum cleaner cichlid, Gnathochromis permaxillaris.

BACKGROUND: Teleosts display a spectacular diversity of craniofacial adaptations that often mediates ecological specializations. A considerable amount of research has revealed molecular players underlying skeletal craniofacial morphologies, but less is known about soft craniofacial phenotypes. Here we focus on an example of lip hypertrophy in the benthivorous Lake Tangnayika cichlid, Gnathochromis permaxillaris, considered to be a morphological adaptation to extract invertebrates out of the uppermost layer of mud bottom. We investigate the molecular and regulatory basis of lip hypertrophy in G. permaxillaris using a comparative transcriptomic approach. RESULTS: We identified a gene regulatory network involved in tissue overgrowth and cellular hypertrophy, potentially associated with the formation of a locally restricted hypertrophic lip in a teleost fish species. Of particular interest were the increased expression level of apoda and fhl2, as well as reduced expression of cyp1a, gimap8, lama5 and rasal3, in the hypertrophic lip region which have been implicated in lip formation in other vertebrates. Among the predicted upstream transcription factors, we found reduced expression of foxp1 in the hypertrophic lip region, which is known to act as repressor of cell growth and proliferation, and its function has been associated with hypertrophy of upper lip in human. CONCLUSION: Our results provide a genetic foundation for future studies of molecular players shaping soft and exaggerated, but locally restricted, craniofacial morphological changes in fish and perhaps across vertebrates. In the future, we advocate integrating gene regulatory networks of various craniofacial phenotypes to understand how they collectively govern trophic and behavioural adaptations.

A new linkage group involved in sex determination in haplotilapiine cichlids.

The high diversity of sex chromosomes and sex determination systems among haplotilapiines suggests that this large cichlid clade is a good model for investigating the evolution of genetics of sex determination. Nonetheless, information about sex determination in this clade remains sparse. The present study reports a microsatellite marker that is closely associated with sex in Xenotilapia rotundiventralis from Lake Tanganyika, Africa. This study is the first to suggest the role of linkage group 17 in sex determination in haplotilapiine cichlids.

Structural manipulations of a shelter resource reveal underlying preference functions in a shell-dwelling cichlid fish.

Many animals can modify the environments in which they live, thereby changing the selection pressures they experience. A common example of such niche construction is the use, creation or modification of environmental resources for use as nests or shelters. Because these resources often have correlated structural elements, it can be difficult to disentangle the relative contribution of these elements to resource choice, and the preference functions underlying niche-construction behaviour remain hidden. Here, we present an experimental paradigm that uses 3D scanning, modelling and printing to create replicas of structures that differ with respect to key structural attributes. We show that a niche-constructing, shell-dwelling cichlid fish, Neolamprologus multifasciatus, has strong open-ended preference functions for exaggerated shell replicas. Fish preferred shells that were fully intact and either enlarged, lengthened or had widened apertures. Shell intactness was the most important structural attribute, followed by shell length, then aperture width. We disentangle the relative roles of different shell attributes, which are tightly correlated in the wild, but nevertheless differentially influence shelter choice and therefore niche construction in this species. We highlight the broad utility of our approach when compared with more traditional methods (e.g. two-choice tasks) for studying animal decision-making in a range of contexts.

Structural genomic variation leads to genetic differentiation in Lake Tanganyika's sardines.

Identifying patterns in genetic structure and the genetic basis of ecological adaptation is a core goal of evolutionary biology and can inform the management and conservation of species that are vulnerable to population declines exacerbated by climate change. We used reduced-representation genomic sequencing methods to gain a better understanding of genetic structure among and within populations of Lake Tanganyika's two sardine species, Limnothrissa miodon and Stolothrissa tanganicae. Samples of these ecologically and economically important species were collected across the length of Lake Tanganyika, as well as from nearby Lake Kivu, where L. miodon was introduced in 1959. Our results reveal differentiation within both S. tanganicae and L. miodon that is not explained by geography. Instead, this genetic differentiation is due to the presence of large sex-specific regions in the genomes of both species, but involving different polymorphic sites in each species. Our results therefore indicate rapidly evolving XY sex determination in the two species. Additionally, we found evidence of a large chromosomal rearrangement in L. miodon, creating two homokaryotypes and one heterokaryotype. We found all karyotypes throughout Lake Tanganyika, but the frequencies vary along a north-south gradient and differ substantially in the introduced Lake Kivu population. We do not find evidence for significant isolation by distance, even over the hundreds of kilometres covered by our sampling, but we do find shallow population structure.

Brood parasitism of an open-water spawning cichlid by the cuckoo catfish.

The cuckoo catfish Synodontis multipunctatus and S. grandiops are endemic to Lake Tanganyika and the only known nonavian vertebrates that exhibit obligate interspecific brood parasitism. Seven maternal mouth-brooding cichlid fish species are reported to be natural hosts of the parasitic catfish and share a common reproductive behaviour that the catfish exploits: cichlid females spawn eggs on the bottom, allowing the catfish female to place her eggs near the cichlid eggs, and the cichlid females collect the catfish eggs by mouth together with their own eggs. However, so far it has not been reported that the cuckoo catfish exploit different spawning behaviours. The genus Cyprichromis consists of five maternal mouth-brooding species endemic to Lake Tanganyika, most of which spawn and collect eggs in open water. This study reports that the cuckoo catfish also parasitizes the open-water spawning Cyprichromis coloratus, although it may not be a regular host.

A functional trade-off between trophic adaptation and parental care predicts sexual dimorphism in cichlid fish.

Although sexual dimorphism is widespread in nature, its evolutionary causes often remain elusive. Here we report a case where a sex-specific conflicting functional demand related to parental care, but not to sexual selection, explains sexual dimorphism in a primarily trophic structure, the gill rakers of cichlid fishes. More specifically, we examined gill raker length in a representative set of cichlid fish species from Lake Tanganyika featuring three different parental care strategies: (i) uni-parental mouthbrooding, whereby only one parental sex incubates the eggs in the buccal cavity; (ii) bi-parental mouthbrooding, whereby both parents participate in mouthbrooding; and (iii) nest guarding without any mouthbrooding involved. As predicted from these different parental care strategies, we find sexual dimorphism in gill raker length to be present only in uni-parental mouthbrooders, but not in bi-parental mouthbrooders nor in nest guarders. Moreover, variation in the extent of sexual dimorphism among uni-parental mouthbrooders appears to be related to trophic ecology. Overall, we present a previously unrecognized scenario for the evolution of sexual dimorphism that is not related to sexual selection or initial niche divergence between sexes. Instead, sexual dimorphism in gill raker length in uni-parental mouthbrooding cichlid fish appears to be the consequence of a sex-specific functional trade-off between a trophic function present in both sexes and a reproductive function present only in the brooding sex.

Do Scales of the Cichlid Altolamprologus compressiceps in Lake Tanganyika Function as a Morphological Defense Against Scale-Eating?

Many cichlid species in the shallow-shore of Lake Tanganyika suffer damage from attacks by the scale-eater Perissodus microlepis. Many prey fish engage in warning behaviors to this predator. It has been hypothesized that, if prey fish have difficulty employing such behavioral tactics, morphological defenses against scale-eating, such as hard scales, will evolve. The shrimp-eating cichlids, Altolamprologus compressiceps (Ac) and Neolamprologus fasciatus (Nf), exhibit hunting behaviors in which they remain motionless for up to 10 seconds while aiming at prey, when they are vulnerable to scale-eating predators; thus, these fish have likely evolved morphological defenses against "scale-attacks". We tested this hypothesis in Ac and Nf, as well as three other predatory fish, Lamprologus callipterus, Lepidiolamprologus elongatus and Lep. attenuatus, that are not motionless for such a long time. Under natural conditions, Ac and Nf were rarely attacked, while the other three species were attacked frequently. When freshly killed specimens of these five species were displayed underwater in the presence of P. microlepis, Ac was rarely attacked, while Nf and the three other species were attacked frequently. Among the five fish species, the force required to tear off scales was highest for Ac, and this force was negatively correlated with the frequency of attacks on the displayed fish. These results support the hypothesis that the hard scales of Ac function as an anti-scale-attack measure, although it remains unclear why free-swimming Nf were rarely attacked while aiming at prey, despite the fact that the force required to tear off its scales was not large.

Climate warming reduces fish production and benthic habitat in Lake Tanganyika, one of the most biodiverse freshwater ecosystems.

Warming climates are rapidly transforming lake ecosystems worldwide, but the breadth of changes in tropical lakes is poorly documented. Sustainable management of freshwater fisheries and biodiversity requires accounting for historical and ongoing stressors such as climate change and harvest intensity. This is problematic in tropical Africa, where records of ecosystem change are limited and local populations rely heavily on lakes for nutrition. Here, using a ∼1,500-y paleoecological record, we show that declines in fishery species and endemic molluscs began well before commercial fishing in Lake Tanganyika, Africa's deepest and oldest lake. Paleoclimate and instrumental records demonstrate sustained warming in this lake during the last ∼150 y, which affects biota by strengthening and shallowing stratification of the water column. Reductions in lake mixing have depressed algal production and shrunk the oxygenated benthic habitat by 38% in our study areas, yielding fish and mollusc declines. Late-20th century fish fossil abundances at two of three sites were lower than at any other time in the last millennium and fell in concert with reduced diatom abundance and warming water. A negative correlation between lake temperature and fish and mollusc fossils over the last ∼500 y indicates that climate warming and intensifying stratification have almost certainly reduced potential fishery production, helping to explain ongoing declines in fish catches. Long-term declines of both benthic and pelagic species underscore the urgency of strategic efforts to sustain Lake Tanganyika's extraordinary biodiversity and ecosystem services.

Brood-tending males in a biparental fish suffer high paternity losses but rarely cuckold.

Extra-pair paternity within socially monogamous mating systems is well studied in birds and mammals but rather neglected in other animal taxa. In fishes, social monogamy has evolved several times but few studies have investigated the extent to which pair-bonded male fish lose fertilizations to cuckolders and gain extra-pair fertilizations themselves. We address this gap and present genetic paternity data collected from a wild population of Variabilichromis moorii, a socially monogamous African cichlid with biparental care of offspring. We show that brood-tending, pair-bonded males suffer exceptionally high paternity losses, siring only 63% of the offspring produced by their female partners on average. The number of cuckolders per brood ranged up to nine and yet, surprisingly, brood-tending males in the population were rarely the culprits. Brood-tending males sired very few extra-pair offspring, despite breeding in close proximity to one another. While unpaired males were largely responsible for the cuckoldry, pair-bonded males still enjoyed higher fertilization success than individual unpaired males. We discuss these results in the context of ecological and phenotypic constraints on cuckoldry and the fitness payoffs of alternative male tactics. Our study provides new insights into how pair-bonded males handle the trade-off between securing within-pair and extra-pair reproduction.

Disentangling Incomplete Lineage Sorting and Introgression to Refine Species-Tree Estimates for Lake Tanganyika Cichlid Fishes.

Adaptive radiation is thought to be responsible for the evolution of a great portion of the past and present diversity of life. Instances of adaptive radiation, characterized by the rapid emergence of an array of species as a consequence to their adaptation to distinct ecological niches, are important study systems in evolutionary biology. However, because of the rapid lineage formation in these groups, and occasional gene flow between the participating species, it is often difficult to reconstruct the phylogenetic history of species that underwent an adaptive radiation. In this study, we present a novel approach for species-tree estimation in rapidly diversifying lineages, where introgression is known to occur, and apply it to a multimarker data set containing up to 16 specimens per species for a set of 45 species of East African cichlid fishes (522 individuals in total), with a main focus on the cichlid species flock of Lake Tanganyika. We first identified, using age distributions of most recent common ancestors in individual gene trees, those lineages in our data set that show strong signatures of past introgression. This led us to formulate three hypotheses of introgression between different lineages of Tanganyika cichlids: the ancestor of Boulengerochromini (or of Boulengerochromini and Bathybatini) received genomic material from the derived H-lineage; the common ancestor of Cyprichromini and Perissodini experienced, in turn, introgression from Boulengerochromini and/or Bathybatini; and the Lake Tanganyika Haplochromini and closely related riverine lineages received genetic material from Cyphotilapiini. We then applied the multispecies coalescent model to estimate the species tree of Lake Tanganyika cichlids, but excluded the lineages involved in these introgression events, as the multispecies coalescent model does not incorporate introgression. This resulted in a robust species tree, in which the Lamprologini were placed as sister lineage to the H-lineage (including the Eretmodini), and we identify a series of rapid splitting events at the base of the H-lineage. Divergence ages estimated with the multispecies coalescent model were substantially younger than age estimates based on concatenation, and agree with the geological history of the Great Lakes of East Africa. Finally, we formally tested the three hypotheses of introgression using a likelihood framework, and find strong support for introgression between some of the cichlid tribes of Lake Tanganyika.