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Animal body coloration is often linked to social dominance and mating success. This is because it can carry information on an animal's body condition and competitive ability by reflecting the genetic quality of individuals or by responding to their current or past living conditions. The present study investigates genetic and environmental effects on a conspicuous color pattern of the cichlid fish Tropheus sp. black "Ikola," in which the size of a carotenoid-based yellow area on the body co-varies with social dominance. To examine environmental plasticity of the color pattern, we tested for effects of early-life stress, induced by reduced feeding of juveniles prior to color pattern formation, as well as effects of a stress treatment administered to fully colored adult fish. None of the stress treatments affected the color pattern as quantified by the width of the yellow bar. However, offspring bar width was correlated to parental values in mid-parent-mid-offspring regression analyses, and animal models estimated significant additive genetic effects on bar width, indicating heritability of the trait. Depending on the random effects structure of the animal models (i.e., whether including or excluding maternal and brood effects), narrow-sense heritability estimates for bar width ranged between 0.2 and 0.8, with the strongest statistical support for the highest estimate. In each of the alternative models, a large proportion of the total variance in bar width was explained by the included random effects, suggesting that bar width is strongly determined by genetic factors or shared maternal and brood environments, with limited scope for environmental influences later in life., Competing Interests: The authors have no conflicts of interest., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

Situated in the Eastern section of the European Alps, Austria encompasses a great diversity of different habitat types, ranging from alpine to lowland Pannonian ecosystems, and a correspondingly high level of species diversity, some of which has been addressed in various DNA barcoding projects. Here, we report a DNA barcode library of all the 476 species of Geometridae (Lepidoptera) that have been recorded in Austria. As far as possible, species were sampled from different Austrian regions in order to capture intraspecific genetic variation. In total, 2500 DNA barcode sequences, representing 438 species, were generated in this study. For complete coverage of Austrian geometrid species in the subsequent analyses, the dataset was supplemented with DNA barcodes from specimens of non-Austrian origin. Species delimitations by ASAP, BIN and bPTP methods yielded 465, 510 and 948 molecular operational taxonomic units, respectively. Congruency of BIN and ASAP partitions with morphospecies assignments was reasonably high (85% of morphospecies in unique partitions), whereas bPTP appeared to overestimate the number of taxonomic units. The study furthermore identified taxonomically relevant cases of morphospecies splitting and sharing in the molecular partitions. We conclude that DNA barcoding and sequence analysis revealed a high potential for accurate DNA-based identification of the Austrian Geometridae species. Additionally, the study provides an updated checklist of the geometrid moths of Austria., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Schattanek-Wiesmair et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

When one of two parents disappears in the midst of caring for offspring, the remaining parent is left with several options. They can either (1) desert the brood, (2) continue caring on their own and reject propositions from new potential partners, or (3) continue caring but remain receptive to re-mating opportunities. The presence of a brood may increase re-mating success of single parents, either because brood care is perceived as a signal of partner quality, or because prospective mates perceive the brood as potential energy source. In this field experiment, we used the socially monogamous, biparental cichlid fish Variabilichromis moorii to examine the re-mating strategy of males with or without dependent offspring after the loss of their female partner. Partner vacancies were filled quickly by new females, and these females engaged in high levels of affiliative behavior with the males. The new females engaged in territorial defense, but focused primarily against intruding conspecifics, likely as a means to repel rivals. The males, in turn, took over the majority of territorial defense against intruding heterospecifics. Interestingly, males that still had offspring from their previous partnerships did not show aggression toward their new female partners, even when those females were infanticidal and cannibalizing the males' current offspring. Overall, our experiment shows that single fathers of a biparental species will re-mate quickly even at the detriment to their current offspring., Competing Interests: The authors have no competing interest do declare., (© The Author(s) 2023. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology.)

Group-living animals are faced with the challenge of sharing space and local resources amongst group members who may be either relatives or non-relatives. Individuals may reduce the inclusive fitness costs they incur from competing with relatives by either reducing their levels of aggression toward kin, or by maintaining physical separation between kin. In this field study, we used the group-living cichlid Neolamprologus multifasciatus to examine whether within-group aggression is reduced among group members that are kin, and whether kin occupy different regions of their group's territory to reduce kin competition over space and local resources. We determined the kinship relationships among cohabiting adults via microsatellite genotyping and then combined these with spatial and behavioral analyses of groups in the wild. We found that aggressive contests between group members declined in frequency with spatial separation between their shelters. Female kin did not engage in aggressive contests with one another, whereas non-kin females did, despite the fact these females lived at similar distances from one another on their groups' territories. Contests within male-male and male-female dyads did not clearly correlate with kinship. Non-kin male-male and male-female dyads lived at more variable distances from one another on their territories than their corresponding kin dyads. Together, our study indicates that contests among group members can be mediated by relatedness in a sex-dependent manner. We also suggest that spatial relationships can play an important role in determining the extent to which group members compete with one another., Competing Interests: The authors have no conflicts of interest to declare., (© The Author(s) 2023. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology.)

Variation in fin shape is one of the most prominent features of morphological diversity among fish. Regulation of fin growth has mainly been studied in zebrafish, and it is not clear whether the molecular mechanisms underlying shape variation are equally diverse or rather conserved across species. In the present study, expression levels of 37 candidate genes were tested for association with fin shape in the cichlid fish Lamprologus tigripictilis . The tested genes included members of a fin shape-associated gene regulatory network identified in a previous study and novel candidates selected within this study. Using both intact and regenerating fin tissue, we tested for expression differences between the elongated and the short regions of the spade-shaped caudal fin and identified 20 genes and transcription factors (including angptl5 , cd63 , csrp1a , cx43 , esco2 , gbf1 , and rbpj ), whose expression patterns were consistent with a role in fin growth. Collated with available gene expression data of two other cichlid species, our study not only highlights several genes that were correlated with fin growth in all three species (e.g., angptl5 , cd63 , cx43 , and mmp9 ), but also reveals species-specific gene expression and correlation patterns, which indicate considerable divergence in the regulatory mechanisms of fin growth across cichlids., Supplementary Information: The online version contains supplementary material available at 10.1007/s10750-022-05068-4., Competing Interests: Conflict of interestThe authors declare that they have no competing interests., (© The Author(s) 2022.)

Mating patterns in animal populations can respond to environmental conditions and consequently vary across time. To examine this variation in nature, studies must include temporal replicates from the same population. Here, we report temporal variation in genetic parentage in the socially monogamous cichlid Variabilichromis moorii from Lake Tanganyika, using samples of broods and their brood-tending parents that were collected across five field trips from the same study population. The sampled broods were either spawned during the dry season (three field trips) or during the rainy season (two trips). In all seasons, we detected substantial rates of extra-pair paternity, which were ascribed to cuckoldry by bachelor males. Paternity shares of brood-tending males were consistently higher, and the numbers of sires per brood were consistently lower, in broods that were spawned in the dry seasons compared to broods from the rainy seasons. In contrast, the strength of size-assortative pairing in our V. moorii population did not vary temporally. Seasonal fluctuations in environmental conditions, such as water turbidity, are proposed as a mechanism behind variable cuckolder pressure. Our data demonstrate the utility of long-term monitoring to improve our understanding of animal mating patterns., Supplementary Information: The online version contains supplementary material available at 10.1007/s10750-022-05042-0., Competing Interests: Competing interestThe authors have no competing interests to declare that are relevant to the content of this article., (© The Author(s) 2022.)

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., (© 2022 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

Background: Sex-biased dispersal is a common and widespread phenomenon that can fundamentally shape the genetic structure of the social environments in which animals live. For animals that live in and move between social groups, sex-biased dispersal can result in an asymmetry in the degree of relatedness among cohabiting males and females, which can have strong implications for their social evolution. In this study, we measured the relatedness structure within and across groups of a wild population of Neolamprologus multifasciatus, a highly-social, shell-dwelling cichlid fish endemic to Lake Tanganyika, East Africa. In total, we genotyped 812 fish from 128 social groups at 20 microsatellite loci. Neolamprologus multifasciatus live at high densities, and also experience strong ecological constraints on free movement throughout their habitat. At the same time, they exhibit sex differences in the degree of reproductive competition within their groups and this makes them an excellent model system for studying the factors associated with sex-biased dispersal., Results: Social groups of N. multifasciatus consist of multiple males and females living together. We found that cohabiting females were unrelated to one another (Lynch-Ritland estimates of relatedness = 0.045 ± 0.15, average ± SD), while males shared much higher, albeit variable, levels of relatedness to other males in their groups (0.23 ± 0.27). We uncovered a pronounced decline in relatedness between males living in separate groups as the spatial separation between them increased, a pattern that was not evident in females. Female dispersal was also markedly constrained by the distribution and availability of nearby territories to which they could emigrate., Conclusions: Our results indicate female-biased dispersal in N. multifasciatus. Our study also highlights how the spatial distribution of suitable dispersal destinations can influence the movement decisions of animals. We also emphasize how sex-biased dispersal can influence the relatedness structure of the social environment in which individuals interact and compete with one another., (© 2022. The Author(s).)

Adverse conditions during early life can cause lasting body size deficits with effects on social and sexual competition, while an accelerated growth response can allow animals to catch up in body size but can be physiologically costly as well. How animals balance growth deficits and growth compensation is predicted to depend on the effects of each on lifetime fitness. We investigated the effects of experimental early-life food restriction on growth, body condition, and adult contest competition in a cichlid fish ( Tropheus sp.). Their longevity and aseasonal breeding suggest that, with view on lifetime reproductive success, temporarily growth-restricted Tropheus should rather invest extra time in reaching competitive body size than risk the potential costs of accelerated growth. However, size-selective predation pressure by gape size-limited piscivores may have favored the evolution of an accelerated growth response to early-life delays. Experimentally food-restricted fish temporarily reduced their growth rate compared to a control group, but maintained their body condition factor at the control level throughout the 80-week study period. There was no evidence for an accelerated growth response following the treatment, as the food-restricted fish never exceeded the size-specific growth rates that were measured in the control group. Food-restricted fish caught up with the body size of the control group several months after the end of the treatment period and were as likely as control fish to win size-matched contests over territories. Regardless of feeding regime, there were sex-specific differences in growth rates and in the trajectories of condition factors over time. Females grew more slowly than males but maintained their condition factors at a high level throughout the study period, whereas the males' condition factors declined over time. These differences may reflect sex-specific contributions of condition and body size to adult fitness that are associated with female mouthbrooding and male competition for breeding territories., Competing Interests: None declared., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The famously diverse body coloration of cichlid fish serves communicative functions in mating and social interactions including competition for resources. Here, we examined the effects of a color pattern trait-the width of a yellow bar on a black body-on territorial competition in males and females of a color variant ("Ikola") of the Lake Tanganyika cichlid Tropheus . First, measuring integumentary carotenoid concentrations in the yellow and black body regions, we established that wider yellow bars require more carotenoids allocated to body coloration. However, we also detected high carotenoid concentrations in the black body regions (> 100 µg/g fresh skin), raising questions about the function of non-displayed integumentary carotenoids. Behavioral experiments showed that fish with wider bars were quicker to explore an unfamiliar area of the tank. In experiments including presentations of fish dummies, the bar width of 'territorial' dummies had no effect on the latency time which test fish took to intrude into the dummies' territories. However, male test fish performed fewer aggressive acts against wide-barred than against narrow-barred dummy competitors. Our results suggest that intimidation by wide bars as well as correlations between bar width and explorative behavior may contribute to mediating success in territorial Tropheus "Ikola"., Competing Interests: Conflict of interestThe authors declare that they have no conflict of interest., (© The Author(s) 2020.)

Carotenoid pigments play a major role in animal body colouration, generating strong interest in the genes involved in the metabolic processes that lead from their dietary uptake to their storage in the integument. Here, we used RNA sequencing (RNA-Seq) to test for differentially expressed genes in a taxonomically replicated design using three pairs of related cichlid fish taxa from the genera Tropheus and Aulonocara . Within each pair, taxa differed in terms of red and yellow body colouration, and high-performance liquid chromatography (HPLC) analyses of skin extracts revealed different carotenoid profiles and concentrations across the studied taxa. Five genes were differentially expressed in all three yellow-red skin contrasts ( dhrsx , nlrc3 , tcaf2 , urah and ttc39b ), but only the tetratricopeptide repeat protein-coding gene ttc39b , whose gene product is linked to mammalian lipid metabolism, was consistently expressed more highly in the red skin samples. The RNA-Seq results were confirmed by quantitative PCR. We propose ttc39b as a compelling candidate gene for variation in animal carotenoid colouration. Since differential expression of ttc39b was correlated with the presence/absence of yellow carotenoids in a previous study, we suggest that ttc39b is more likely associated with the concentration of total carotenoids than with the metabolic formation of red carotenoids.

The otoliths of teleost fishes exhibit a great deal of inter- and intra-species shape variation. The ecomorphology of the saccular otolith is often studied by comparing its shape across species and populations inhabiting a range of environments. However, formal tests are often lacking to examine how closely variation in otolith shape follows the genetic drift of a neutral trait. Here, we examine patterns of saccular otolith shape variation in four species of African cichlid fishes, each sampled from three field sites. All four species showed the greatest level of otolith shape variation along two principal component axes, one pertaining to otolith height and another to the prominence of an anterior notch. Fish collected from the same site possessed similarities in saccular otolith shape relative to fish from other sites, and these 'site-difference' signatures were consistent across species and observable in both sexes. Sex-differences in saccular otolith shape differed in magnitude from site to site. Population differences in saccular otolith shape did not covary with neutral genetic differentiation between those populations. Otolith height, in particular, displayed large site similarities across species, weak correlation with neutral genetic variation, and strong sex differences, collectively suggesting that otolith shape represents a selectively non-neutral trait.

Background: Carotenoids contribute significantly to animal body coloration, including the spectacular color pattern diversity among fishes. Fish, as other animals, derive carotenoids from their diet. Following uptake, transport and metabolic conversion, carotenoids allocated to body coloration are deposited in the chromatophore cells of the integument. The genes involved in these processes are largely unknown. Using RNA-Sequencing, we tested for differential gene expression between carotenoid-colored and white skin regions of a cichlid fish, Tropheus duboisi "Maswa", to identify genes associated with carotenoid-based integumentary coloration. To control for positional gene expression differences that were independent of the presence/absence of carotenoid coloration, we conducted the same analyses in a closely related population, in which both body regions are white., Results: A larger number of genes (n = 50) showed higher expression in the yellow compared to the white skin tissue than vice versa (n = 9). Of particular interest was the elevated expression level of bco2a in the white skin samples, as the enzyme encoded by this gene catalyzes the cleavage of carotenoids into colorless derivatives. The set of genes with higher expression levels in the yellow region included genes involved in xanthophore formation (e.g., pax7 and sox10), intracellular pigment mobilization (e.g., tubb, vim, kif5b), as well as uptake (e.g., scarb1) and storage (e.g., plin6) of carotenoids, and metabolic conversion of lipids and retinoids (e.g., dgat2, pnpla2, akr1b1, dhrs). Triglyceride concentrations were similar in the yellow and white skin regions. Extracts of integumentary carotenoids contained zeaxanthin, lutein and beta-cryptoxanthin as well as unidentified carotenoid structures., Conclusion: Our results suggest a role of carotenoid cleavage by Bco2 in fish integumentary coloration, analogous to previous findings in birds. The elevated expression of genes in carotenoid-rich skin regions with functions in retinol and lipid metabolism supports hypotheses concerning analogies and shared mechanisms between these metabolic pathways. Overlaps in the sets of differentially expressed genes (including dgat2, bscl2, faxdc2 and retsatl) between the present study and previous, comparable studies in other fish species provide useful hints to potential carotenoid color candidate genes.

Blooms of gelatinous zooplankton can represent dramatic environmental perturbations for aquatic ecosystems. Yet, we still know little about how blooms impact fitness-related behaviours of fish caught within their areas of effect, especially for freshwater systems. Here, we documented the behavioural impacts of freshwater hydrozoan ( Limnocnida tanganjicae ) blooms on a territorial cichlid ( Variabilichromis moorii ), as well as on the wider community of cichlids in a shallow-water rocky habitat of Lake Tanganyika. Compared with non-bloom conditions, V. moorii individuals in the midst of blooms reduced their swimming and territory defence activities (each by approx. 50%) but not their foraging or affiliative behaviours. Despite this reduction in activity, V. moorii could not entirely avoid being stung and preferred to remain closer to the rocky substrata as opposed to the more open demersal zone. Many other fishes similarly hid among the benthic substrata, changing the composition of the fish community in the demersal zone during bloom conditions. Reductions in activity could have multiple fitness-related implications for individual fish. Establishing the consequences of these behavioural changes is important for understanding the effects of gelatinous zooplankton blooms in freshwater systems., Competing Interests: Prof. Kristina Sefc is a Board Member of Royal Society Open Science., (© 2019 The Authors.)

Background: Raising unrelated offspring is typically wasteful of parental resources and so individuals are expected to reduce or maintain low levels of parental effort when their parentage is low. This can involve facultative, flexible adjustments of parental care to cues of lost parentage in the current brood, stabilizing selection for a low level of paternal investment, or an evolutionary reduction in parental investment in response to chronically low parentage., Results: We studied parental care in Variabilichromis moorii, a socially monogamous, biparental cichlid fish, whose mating system is characterized by frequent cuckoldry and whose primary form of parental care is offspring defense. We combine field observations with genetic parentage analyses to show that while both parents defend their nest against intruding con- and hetero-specifics, males and females may do so for different reasons. Males in the study group (30 breeding pairs) sired 0-100% (median 83%) of the fry in their nests. Males defended less against immediate threats to the offspring, and more against threats to their territories, which are essential for the males' future reproductive success. Males also showed no clear relationship between their share of defense and their paternity of the brood. Females, on the other hand, were related to nearly all the offspring under their care, and defended almost equally against all types of threats., Conclusion: Overall, males contributed less to defense than females and we suggest that this asymmetry is the result of an evolutionary response by males to chronically high paternity loss in this species. Although most males in the current study group achieved high parentage in their nests, the average paternity in V. moorii, sampled across multiple seasons, is only about 55%. We highlight the importance and complexity of studying nest defense as a form of parental care in systems where defense may serve not only to protect current offspring, but also to ensure future reproductive success by maintaining a territory.

We report a hitherto unknown radiation within the clingfishes (Gobiesocidae), discovered in one of the best-studied marine biomes, the Mediterranean Sea. The monotypic genus Gouania is a Mediterranean endemic inhabiting the interstices of gravel beaches. Using geometric morphometric analyses, we identified two distinct morphotypes (characterized by a slender and a stout body shape, respectively) among Gouania willdenowi sampled from the three major Mediterranean basins (Eastern, Western and Adriatic). Slender and stout G. willdenowi occurred sympatrically in the Adriatic and the Eastern Mediterranean basins. Only the stout morphotype was found in the Western Mediterranean. Morphotypes were further distinguished by relative eye size and number of vertebrae. Based on mitochondrial and nuclear gene sequences, five highly divergent clades (COI-based K2P distances of 8 to 15%) were identified. The clades represented, respectively, stout Adriatic, slender Adriatic, stout Eastern, slender Eastern and stout Western Mediterranean Gouania. This suggests that the genus Gouania comprises at least five different species. Phylogenetic relationships among clades furthermore imply repeated evolution of convergent morphotypes. The onset of the Gouania radiation was dated to 3.23 (95% HPD 2.08-5.90) mya, and may therefore have overlapped with or followed the Messinian salinity crisis., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Comparative analyses of gene regulation inform about the molecular basis of phenotypic trait evolution. Here, we address a fin shape phenotype that evolved multiple times independently across teleost fish, including several species within the family Cichlidae. In a previous study, we proposed a gene regulatory network (GRN) involved in the formation and regeneration of conspicuous filamentous elongations adorning the unpaired fins of the Neolamprologus brichardi. Here, we tested the members of this network in the blockhead cichlid, Steatocranus casuarius, which displays conspicuously elongated dorsal and moderately elongated anal fins. Our study provided evidence for differences in the anatomy of fin elongation and suggested gene regulatory divergence between the two cichlid species. Only a subset of the 20 genes tested in S. casuarius showed the qPCR expression patterns predicted from the GRN identified in N. brichardi, and several of the gene-by-gene expression correlations differed between the two cichlid species. In comparison to N. brichardi, gene expression patterns in S. casuarius were in better (but not full) agreement with gene regulatory interactions inferred in zebrafish. Within S. casuarius, the dorsoventral asymmetry in ornament expression was accompanied by differences in gene expression patterns, including potential regulatory differentiation, between the anal and dorsal fin.

Introgression patterns between divergent lineages are often characterized by asymmetry in the direction and among-marker variation in the extent of gene flow, and therefore inform on the mechanisms involved in differentiation and speciation. In the present study, we test the hypothesis that unidirectional introgression between two phenotypically and genetically distinct lineages of the littoral, rock-dwelling cichlid fish Tropheus moorii across a wide sandy bay is linked to observed differences in mate preferences between the two lineages. This hypothesis predicts bi-directional nuclear gene flow and was rejected by congruent patterns of introgression in mtDNA, AFLP and microsatellite markers, with admixture confined to the populations west of the bay. This pattern can be explained on the basis of habitat changes in the course of lake level fluctuations, which first facilitated the development of a symmetric admixture zone including the area corresponding to the present sand bay and then shaped asymmetry by causing local extinctions and cessation of gene flow when this area became once more inhabitable. This conforms with previous assumptions that habitat dynamics are a primary determinant of population-level evolution in Tropheus . In this respect, Tropheus may be representative of species whose preferred habitat is subject to frequent re-structuring.

Background: In socially monogamous species, reproduction is not always confined to paired males and females. Extra-pair males commonly also reproduce with paired females, which is traditionally thought to be costly to the females' social partners. However, we suggest that when the relatedness between reproducing individuals is considered, cuckolded males can suffer lower fitness losses than otherwise expected, especially when the rate of cuckoldry is high. We combine theoretical modeling with a detailed genetic study on a socially monogamous wild fish, Variabilichromis moorii, which displays biparental care despite exceptionally high rates of extra-pair paternity., Results: We measured the relatedness between all parties involved in V. moorii spawning events (i.e. between males and females in social pairs, females and their extra-pair partners, and paired males and their cuckolders), and we reveal that males are on average more related to their cuckolders than expected by chance. Queller-Goodnight estimates of relatedness between males and their cuckolders are on average r = 0.038 but can range up to r = 0.64. This also increases the relatedness between males and the extra-pair offspring under their care. These intriguing results are consistent with the predictions of our mathematical model, which shows that elevated relatedness between paired males and their cuckolders can be adaptive for both parties when competition for fertilizations is strong., Conclusions: Our results show how cuckoldry by relatives can offset males' direct fitness losses with inclusive fitness gains, which can be substantial in systems where males face almost certain paternity losses.

In the absence of dispersal barriers, species with great dispersal ability are expected to show little, if at all, phylogeographic structure. The East African Great Lakes and their diverse fish faunas provide opportunities to test this hypothesis in pelagic fishes, which are presumed to be highly mobile and unrestricted in their movement by physical barriers. Here, we address the link between panmixis and pelagic habitat use by comparing the phylogeographic structure among four deepwater cichlid species of the tribe Bathybatini from Lake Tanganyika. We show that the mitochondrial genealogies (based on the most variable part or the control region) of the four species are very shallow (0.8-4% intraspecific divergence across entire distribution ranges) and that all species experienced recent population growth. A lack of phylogeographic structure in the two eupelagic species, Bathybates fasciatus and B. leo , was consistent with expectations and with findings in other pelagic cichlid species. Contrary to expectations, a clear phylogeographic structure was detected in the two benthopelagic species, B. graueri and Hemibates stenosoma . Differences in genetic diversity between eupelagic and benthopelagic species may be due to differences in their dispersal propensity, mediated by their respective predatory niches, rather than precipitated by external barriers to dispersal.

Selection arising from social competition over non-mating resources, i.e. resources that do not directly and immediately affect mating success, offers a powerful alternative to sexual selection to explain the evolution of conspicuous ornaments, particularly in females. Here, we address the hypothesis that competition associated with the territoriality exhibited by both males and females in the cichlid fish Tropheus selects for the display of a conspicuous colour pattern in both sexes. The investigated pattern consists of a vertical carotenoid-coloured bar on a black body. Bar width affected the probability of winning in size-matched female-female, but not male-male, contests for territory possession. Our results support the idea that the emergence of female territoriality contributed to the evolution of sexual monomorphism from a dimorphic ancestor, in that females acquired the same conspicuous coloration as males to communicate in contest competition., (© 2018 The Authors.)

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., (© 2018 The Authors. Molecular Ecology Published by John Wiley & Sons Ltd.)

Variation in fin shape and size contributes to the outstanding morphological diversity of teleost fishes, but the regulation of fin growth has not yet been studied extensively outside the zebrafish model. A previous gene expression study addressing the ornamental elongations of unpaired fins in the African cichlid fish Neolamprologus brichardi identified three genes (cx43, mmp9 and sema3d) with strong and consistent expression differences between short and elongated fin regions. Remarkably, the expression patterns of these genes were not consistent with inferences on their regulatory interactions in zebrafish. Here, we identify a gene expression network (GRN) comprising cx43, mmp9, and possibly also sema3d by a stepwise approach of identifying co-expression modules and predicting their upstream regulators. Among the transcription factors (TFs) predicted as potential upstream regulators of 11 co-expressed genes, six TFs (foxc1, foxp1, foxd3, myc, egr2, irf8) showed expression patterns consistent with their cooperative transcriptional regulation of the gene network. Some of these TFs have already been implicated in teleost fish fin regeneration and formation. We particularly discuss the potential function of foxd3 as driver of the network and its role in the unexpected gene expression correlations observed in N. brichardi.

The diversity of fin morphology within and across fish taxa offers great, but still largely unexplored, opportunities to investigate the proximate mechanisms underlying fin shape variation. Relying on available genetic knowledge brought forth mainly by the comprehensive study of the zebrafish caudal fin, we explored candidate molecular mechanisms for the maintenance and formation of the conspicuously elongated filaments adorning the unpaired fins of the East African "princess cichlid" Neolamprologus brichardi. Via qPCR assays, we detected expression differences of candidate genes between elongated and short regions of intact and regenerating fins. The identified genes include skeletogenic and growth factors (igf2b, fgf3, bmp2 and bmp4), components of the WNT pathway (lef1, wnt5b and wnt10) and a regulatory network determining fin ray segment size and junction (cx43, esco2 and sema3d), as well as other genes with different roles (mmp9, msxb and pea3). Interestingly, some of these genes showed fin specific expression differences which are often neglected in studies of model fish that focus on the caudal fin. Moreover, while the observed expression patterns were generally consistent with zebrafish results, we also detected deviating expression correlations and gene functions.

Morphological differentiation among closely related species provides opportunities to study mechanisms shaping natural phenotypic variation. Here, we address variation in the orientation of melanin-colored body stripes in three cichlid species of the tribe Haplochromini. Melanochromis auratus displays a common pattern of dark, straight horizontal body stripes, whereas in Aristochromis christyi and Buccochromis rhoadesii , oblique stripes extend from the anterior dorsal to the posterior mid-lateral trunk. We first validated a stably reference gene, and then, investigated the chromatophore distribution in the skin by assessing the expression levels of the iridophore and melanophore marker genes, ltk and slc24a5 , respectively, as well as pmel , a melanophore pigmentation marker gene. We found anterior-posterior differences in the expression levels of the three genes in the oblique-striped species. The higher anterior expression of ltk , indicates increased iridophore density in the anterior region, i.e., uneven horizontal distribution of iridophores, which coincides with the anterior dorsalization of melanophore stripe in these species. The obliqueness of the horizontal body stripes might be a result of distinct migratory or patterning abilities of melanophores in anterior and posterior stripe regions which could be reflected by variation in the expression of genes involved in melanophore patterning. To address this, we investigated anterior-posterior expression levels of a primary set of candidate target genes with known functions in melanophore migration and stripe patterning in the adult zebrafish, and their related gene regulatory network. Among these genes, those with differences in anterior-posterior expression showed only species-specific differential expression, e.g., sdf1a , col14a1a , ifitm5 , and agpat3 , with the exception of fbxw4/hagoromo (differentially expressed in an oblique-and the straight-striped species). In summary, distinct anterior-posterior gradients in iridophore density found to be more similar characteristic between the two oblique-striped species. Furthermore, the species-specific differential expression of genes involved in stripe patterning might also implicate distinct molecular processes underlying the obliqueness of body stripe in two closely related cichlid species., Competing Interests: The authors declare there are no competing interests.

The establishment of hybrid taxa relies on reproductive isolation from the parental forms, typically achieved by ecological differentiation. Here, we present an alternative mechanism, in which shifts in the strength and location of dispersal barriers facilitate diversification by hybridisation. Our case study concerns the highly diverse, stenotopic rock-dwelling cichlids of the African Great Lakes, many of which display geographic colour pattern variation. The littoral habitat of these fish has repeatedly been restructured in the course of ancient lake level fluctuations. Genetic data and an experimental cross support the hybrid origin of a distinct yellow-coloured variant of Tropheus moorii from ancient admixture between two allopatric, red and bluish variants. Deficient assortative mating preferences imply that reproductive isolation continues to be contingent on geographic separation. Linking paleolimnological data with the establishment of the hybrid variant, we sketch a selectively neutral diversification process governed solely by rearrangements of dispersal barriers., (© 2017 John Wiley & Sons Ltd/CNRS.)

Fish color patterns are among the most diverse phenotypic traits found in the animal kingdom. Understanding the molecular and cellular mechanisms that control in chromatophore distribution and pigmentation underlying this diversity is a major goal in developmental and evolutionary biology, which has predominantly been pursued in the zebrafish model system. Here, we apply results from zebrafish work to study a naturally occurring color pattern phenotype in the fins of an African cichlid species from Lake Tanganyika. The cichlid fish Neolamprologus meeli displays a distinct dorsal color pattern, with black and white stripes along the edges of the dorsal fin and of the dorsal half of the caudal fin, corresponding with differences in melanophore density. To elucidate the molecular mechanisms controlling the differences in dorsal and ventral color patterning in the fins, we quantitatively assessed the expression of 15 candidate target genes involved in adult zebrafish pigmentation and stripe formation. For reference gene validation, we screened the expression stability of seven widely expressed genes across the investigated tissue samples and identified tbp as appropriate reference. Relative expression levels of the candidate target genes were compared between the dorsal, striped fin regions and the corresponding uniform, grey-colored regions in the anal and ventral caudal fin. Dorso-ventral expression differences, with elevated levels in both white and black stripes, were observed in two genes, the melanosome protein coding gene pmel and in igsf11 , which affects melanophore adhesion, migration and survival. Next, we predicted potential shared upstream regulators of pmel and igsf11 . Testing the expression patterns of six predicted transcriptions factors revealed dorso-ventral expression difference of irf1 and significant, negative expression correlation of irf1 with both pmel and igsf11 . Based on these results, we propose pmel , igsf11 and irf1 as likely components of the genetic mechanism controlling distinct dorso-ventral color patterns in N. meeli fins., Competing Interests: The authors declare there are no competing interests.

A recent study of North American canids by Rutledge et al. (Biol. Lett. 11, 20150303 (doi:10.1098/rsbl.2015.0303)) refutes the hypothesized hybrid origin of the eastern wolf (EW) based on genomic evidence against very recent hybridization. However, the analyses do not rule out the possibility of more ancient hybridization. Claims to have resolved the evolutionary origin of the EW are therefore inappropriate. Importantly, though, we plead that uncertainty about the ancient history of the taxon should not affect current conservation policy., (© 2016 The Author(s).)

Mating system variability is known to exist between and within species, often due to environmental influences. An open question is whether, vice versa, similar environmental conditions entail congruent mating behavior, for example in terms of multiple paternity, in species or populations sharing largely comparable breeding modes. This study employed microsatellite markers to investigate the incidence of multiple paternity in Cyprichromis coloratus and Cyprichromis leptosoma, two sympatric, closely related, mouthbrooding Lake Tanganyika cichlids with similar ecological and behavioral characteristics including the formation of open-water schools. Mouthbrooding females of both species were collected from the same mixed-species breeding school at the same time, minimizing environmental variation during courtship and mating. In C. coloratus, four of 12 broods had more than one sire, with a mean of 1.33 reconstructed sires per brood. C. leptosoma exhibited multiple paternity in 18 of 22 broods, with a mean of 2.59 or 2.86 reconstructed sires per brood according to the programs gerud and colony, respectively. In addition, two broods were found to contain offspring transplanted from another brood. There was no significant difference in brood size between species, but mean sire number did differ significantly. Hence, substantial similarity in reproductive behavior along with shared environmental conditions during courtship and spawning did not lead to equal rates of polyandry or sneaking in the two species.

The homoplasy excess test (HET) is a tree-based screen for hybrid taxa in multilocus nuclear phylogenies. Homoplasy between a hybrid taxon and the clades containing the parental taxa reduces bootstrap support in the tree. The HET is based on the expectation that excluding the hybrid taxon from the data set increases the bootstrap support for the parental clades, whereas excluding non-hybrid taxa has little effect on statistical node support. To carry out a HET, bootstrap trees are calculated with taxon-jackknife data sets, that is excluding one taxon (species, population) at a time. Excess increase in bootstrap support for certain nodes upon exclusion of a particular taxon indicates the hybrid (the excluded taxon) and its parents (the clades with increased support).We introduce a new software program, hext, which generates the taxon-jackknife data sets, runs the bootstrap tree calculations, and identifies excess bootstrap increases as outlier values in boxplot graphs. hext is written in r language and accepts binary data (0/1; e.g. AFLP) as well as co-dominant SNP and genotype data.We demonstrate the usefulness of hext in large SNP data sets containing putative hybrids and their parents. For instance, using published data of the genus Vitis (~6,000 SNP loci), hext output supports V. × champinii as a hybrid between V. rupestris and V. mustangensis .With simulated SNP and AFLP data sets, excess increases in bootstrap support were not always connected with the hybrid taxon (false positives), whereas the expected bootstrap signal failed to appear on several occasions (false negatives). Potential causes for both types of spurious results are discussed.With both empirical and simulated data sets, the taxon-jackknife output generated by hext provided additional signatures of hybrid taxa, including changes in tree topology across trees, consistent effects of exclusions of the hybrid and the parent taxa, and moderate (rather than excessive) increases in bootstrap support. hext significantly facilitates the taxon-jackknife approach to hybrid taxon detection, even though the simple test for excess bootstrap increase may not reliably identify hybrid taxa in all applications.

Population histories depend on the interplay between exogeneous and endogeneous factors. In marine species, phylogeographic and demographic patterns are often shaped by sea level fluctuations, water currents and dispersal ability. Using mitochondrial control region sequences ( n = 120), we infer phylogeographic structure and historic population size changes of a common littoral fish species, the black-faced blenny Tripterygion delaisi (Perciformes, Blennioidei, Tripterygiidae) from the north-eastern Adriatic Sea. We find that Adriatic T. delaisi are differentiated from conspecific populations in the remaining Mediterranean, but display little phylogeographic structure within the Adriatic basin. The pattern is consistent with passive dispersal of planktonic larvae along cyclonic currents within the Adriatic Sea, but limited active dispersal of adults. Demographic reconstructions are consistent with recent population expansion, probably triggered by rising sea levels after the last glacial maximum (LGM). Placing the onset of population growth between the LGM and the warming of surface waters (18 000-13 000 years BP) and employing a novel expansion dating approach, we inferred a substitution rate of 2.61-3.61% per site per MY. Our study is one of only few existing investigations of the genetic structure of animals within the Adriatic basin and is the first to provide an estimate for mitochondrial control region substitution rates in blennioid fishes.

The largely endemic cichlid species flocks of the East African Great Lakes are among the prime examples for explosive speciation and adaptive radiation. Speciation rates differ among cichlid lineages, and the propensity to radiate has been linked to intrinsic and extrinsic factors such as sexual selection and ecological opportunity. Remarkably, only one cichlid tribe-the Boulengerochromini-comprises just a single species, Boulengerochromis microlepis , a predominantly piscivorous endemic of Lake Tanganyika and the world's largest cichlid. While the lineage diverged from its closest relatives at the onset of the Lake Tanganyika radiation >8 MYA, mitochondrial control region sequences collected in this study dated the most recent common ancestor of B. microlepis to ~60-110 KYA. There was no evidence of phylogeographic structure in the lake-wide sample. Patterns of genetic diversity and demographic analyses were consistent with slow and steady population growth throughout the reconstructed timescale. Additionally, the shallow divergence within the species may be related to a possibly large variance in reproductive success in this highly fecund species. Trophic niche space restriction by sympatric piscivores, lack of geographic structure, low potential for sexual selection arising from the monogamous mating system and extinction may have contributed to keeping the lineage monotypic.

Assortative mating promotes reproductive isolation and allows allopatric speciation processes to continue in secondary contact. As mating patterns are determined by mate preferences and intrasexual competition, we investigated male-male competition and behavioral isolation in simulated secondary contact among allopatric populations. Three allopatric color morphs of the cichlid fish Tropheus were tested against each other. Dyadic male-male contests revealed dominance of red males over bluish and yellow-blotch males. Reproductive isolation in the presence of male-male competition was assessed from genetic parentage in experimental ponds and was highly asymmetric among pairs of color morphs. Red females mated only with red males, whereas the other females performed variable degrees of heteromorphic mating. Discrepancies between mating patterns in ponds and female preferences in a competition-free, two-way choice paradigm suggested that the dominance of red males interfered with positive assortative mating of females of the subordinate morphs and provoked asymmetric hybridization. Between the nonred morphs, a significant excess of negative assortative mating by yellow-blotch females with bluish males did not coincide with asymmetric dominance among males. Hence, both negative assortative mating preferences and interference of male-male competition with positive assortative preferences forestall premating isolation, the latter especially in environments unsupportive of competition-driven spatial segregation.

Female mate preferences for male traits and resource characteristics affect trait evolution and diversification. Here, we test the effects of male body traits and territory characteristics on within-population female preferences and on population-assortative mating in the cichlid Tropheus moorii . Within-population preferences of females were independent of male body size, coloration and territory size but were strongly dependent on territory quality and co-varied with male courtship activity. Courtship activity of individual males was contingent on the quality of their assigned territory, and therefore, courtship may not only indicate intrinsic male quality. On the basis of these results we suggest that female preferences for high-quality territories reinforce the outcome of malemale competition and ensure male mating success. Mating preferences of females for males of their own color variant (ascertained in a previous experiment) were not overturned when males of another color variant were presented in a superior territory, indicating that within- and between-population mate preferences of females depend on different cues.

Non-sexual social selection can underlie the evolution of sexually monomorphic phenotypes. A causal relationship between territorial competition and sexual monomorphism predicts that male and female competitors should employ similar contest behavior and that contest outcome should depend on the same traits in males and females. We test this prediction in a sexually monomorphic cichlid fish of the genus Tropheus, in which males and females defend individual feeding territories. Lineages basal to Tropheus are sexually dimorphic and have non-territorial females, suggesting that a switch to female territoriality and loss of sexual dimorphism occurred in the Tropheus lineage. We compare rates of agonistic behavior and the effects of body size asymmetries on competitive success between male-male and female-female contests in an experimental setup. Body size asymmetry had the same effect in male and female contests, being negatively correlated with contest duration and positively correlated with the probability of winning. Male and female winners employed the same rates of frontal and lateral displays as well as charges against their opponents. Contest duration was longer in females. In tied contests, females displayed more than males. Our data suggest that intraspecific contest competition for territories selects for large body size in both sexes and support a link between the evolution of female territoriality and the loss of sexual size dimorphism in Tropheus .

This study examines the performance of DNA barcodes (mt cytochrome c oxidase 1 gene) in the identification of 1004 species of Lepidoptera shared by two localities (Finland, Austria) that are 1600 km apart. Maximum intraspecific distances for the pooled data were less than 2% for 880 species (87.6%), while deeper divergence was detected in 124 species. Despite such variation, the overall DNA barcode library possessed diagnostic COI sequences for 98.8% of the taxa. Because a reference library based on Finnish specimens was highly effective in identifying specimens from Austria, we conclude that barcode libraries based on regional sampling can often be effective for a much larger area. Moreover, dispersal ability (poor, good) and distribution patterns (disjunct, fragmented, continuous, migratory) had little impact on levels of intraspecific geographic divergence. Furthermore, the present study revealed that, despite the intensity of past taxonomic work on European Lepidoptera, nearly 20% of the species shared by Austria and Finland require further work to clarify their status. Particularly discordant BIN (Barcode Index Number) cases should be checked to ascertain possible explanatory factors such as incorrect taxonomy, hybridization, introgression, and Wolbachia infections.

Phylogenetic analyses of rapid radiations are particularly challenging as short basal branches and incomplete lineage sorting complicate phylogenetic inference. Multilocus data of presence-absence polymorphisms such as obtained by AFLP genotyping overcome some of the difficulties, but also present their own intricacies. Here we analyze >1000 AFLP markers to address the evolutionary history of the Limnochromini, a cichlid fish lineage endemic to Lake Tanganyika, and to test for potential effects of outgroup composition on tree topology. The data support previous mitochondrial evidence on the tribe's taxonomy by confirming the polyphyly of the genus Limnochromis and - in contradiction to a recent taxonomic revision - nesting the genus Greenwoodochromis within the Limnochromini. Species relationships suggest that ecological segregation occurred during the rapid basal radiation of the Limnochromini. The large phylogenetic distance between candidate outgroup taxa and the Limnochromini radiation caused random outgroup effects. Bootstrap support for ingroup nodes was lower in outgroup-rooted than in midpoint-rooted trees, and root positions and ingroup tree topologies varied in response to the composition of the outgroup. These observations suggest that the predisposition for homoplastic evolution makes AFLP-based phylogenetic analyses particularly susceptible to random biases introduced by too-distant outgroup taxa., (Copyright © 2013 The Authors. Published by Elsevier Inc. All rights reserved.)

Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to complex social behaviour and morphological versatility, they are characterised by extensive diversity in colouration, both within and between species. Here, we review the cellular and molecular mechanisms underlying colour variation in this group and the selective pressures responsible for the observed variation. We specifically address the evidence for the hypothesis that divergence in colouration is associated with the evolution of reproductive isolation between lineages. While we conclude that cichlid colours are excellent models for understanding the role of animal communication in species divergence, we also identify taxonomic and methodological biases in the current research effort. We suggest that the integration of genomic approaches with ecological and behavioural studies, across the entire cichlid family and beyond it, will contribute to the utility of the cichlid model system for understanding the evolution of biological diversity., (Copyright © 2013 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Whether premating isolation is achieved by male-specific, female-specific or sex-independent assortative preferences often depends on the underlying evolutionary processes. Here we test mate preferences of males presented with females of different allopatric colour variants of the cichlid fish Tropheus sp., a Lake Tanganyika endemic with rich geographical colour pattern variation, in which the strength of sexual isolation varies between populations. We conducted two-way mate choice experiments to compare behaviour of males of a red-bodied morph (population Moliro) towards females from their own population with behaviour towards females from four allopatric populations at different stages of phylogenetic and phenotypic divergence. Males courted same-population females significantly more intensely than females of other populations, and reduced their heteromorphic courtship efforts both with increasing genetic and increasing phenotypic distinctness of the females. In particular, females of a closely related red-bodied population received significantly more courtship than either genetically distinct, similarly coloured females ('Kirschfleck' morph) or genetically related, differently coloured females ('yellow-blotch' morph), both of which were courted similarly. Genetically and phenotypically distinct females (Tropheus polli) were not courted at all. Consistent with previous female-choice experiments, female courtship activity also decreased with increasing genetic distance from the males' population. Given successful experimental and natural introgression between colour morphs and the pervasive allopatry of related variants, we consider it unlikely that assortative preferences of both sexes were driven by direct selection during periods of secondary contact or, in turn, drove colour pattern differentiation in allopatry. Rather, we suggest that sexual isolation evolved as by-product of allopatric divergence., (© 2012 The Authors. Journal of Evolutionary Biology © 2012 European Society For Evolutionary Biology.)

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.

Genome scan-based tests for selection are directly applicable to natural populations to study the genetic and evolutionary mechanisms behind phenotypic differentiation. We conducted AFLP genome scans in three distinct geographic colour morphs of the cichlid fish Tropheus moorii to assess whether the extant, allopatric colour pattern differentiation can be explained by drift and to identify markers mapping to genomic regions possibly involved in colour patterning. The tested morphs occupy adjacent shore sections in southern Lake Tanganyika and are separated from each other by major habitat barriers. The genome scans revealed significant genetic structure between morphs, but a very low proportion of loci fixed for alternative AFLP alleles in different morphs. This high level of polymorphism within morphs suggested that colour pattern differentiation did not result exclusively from neutral processes. Outlier detection methods identified six loci with excess differentiation in the comparison between a bluish and a yellow-blotch morph and five different outlier loci in comparisons of each of these morphs with a red morph. As population expansions and the genetic structure of Tropheus make the outlier approach prone to false-positive signals of selection, we examined the correlation between outlier locus alleles and colour phenotypes in a genetic and phenotypic cline between two morphs. Distributions of allele frequencies at one outlier locus were indeed consistent with linkage to a colour locus. Despite the challenges posed by population structure and demography, our results encourage the cautious application of genome scans to studies of divergent selection in subdivided and recently expanded populations., (© 2012 Blackwell Publishing Ltd.)

Uniparental maternal brood care often coincides with multiple paternity and single maternity of broods, possibly reflecting benefits of polyandry and costs of uniparental care. Genetic data from the maternally mouthbrooding cichlid fish Simochromis pleurospilus revealed the opposite pattern--low polyandry and allomaternal care. More than 70% of the investigated females had mated with a single male, and 14% of the females had unrelated fry in their broods. Broods with foreign fry were in the late stage of brood care, in which females guard free-swimming fry and recall the broods into their mouths for protection. With one exception, fostering females were related to their adopted fry at the level of first cousins (R(QG) > 0.12), but relatedness between fosters and adopted fry was not significantly higher than between fosters and fry tended by other females. Relatedness among breeders extended to the level of first-order relatives. Mean relatedness among contemporaneously breeding dams (R(QG) = 0.08) was significantly higher than among dams breeding in different seasons (R(QG) = -0.04), which suggests a temporal or spatial concentration of mouthbrooding relatives. Indeed, females sometimes brood in small groups. This behaviour may reduce brood predation but will increase the risk of brood mixing, which is possibly mitigated by low costs of brood care and indirect benefits accrued by relatedness among the breeders in the group. Remarkably, the apparent inbreeding potential did not give rise to bet-hedging polyandry or active avoidance of relatives, as half of the mated individuals were related at R(QG)  > 0.13 and polyandry did not coincide with high within-pair relatedness., (© 2012 Blackwell Publishing Ltd.)

Discriminating female mate preferences enhance the variance in reproductive success among males of a population and create a potential for sexual selection, which can account for trait evolution and diversification. Fish color patterns are among the prime targets of mate choice-driven sexual selection. Populations of the cichlid Tropheus from Lake Tanganyika display remarkable geographic color pattern variation, but the role of female choice in their rapid and rich phenotypic diversification is unclear. Males and females establish a pair bond prior to spawning monogamously, but as brood care is strictly maternal, female investment in reproduction is high and the operational sex ratio is male-biased. Therefore, variance in male reproductive success can accrue if individual males succeed repeatedly in securing a mate. To test this prediction in the red colored Tropheus moorii "Chimba", four pairs of males were presented to a series of females and female mate preferences were inferred from pairwise interactions. There was a significant difference in mating success between the males of each pair ( P < 0.001 over all trials), as-with one exception-females shared preferences for the same males. Male courtship activity was strongly correlated with female choice. Our experiment suggests that female choice contributes to the variance in male reproductive success in the tested population.

Hybridization among littoral cichlid species in Lake Tanganyika was inferred in several molecular phylogenetic studies. The phenomenon is generally attributed to the lake level-induced shoreline and habitat changes. These allow for allopatric divergence of geographically fragmented populations alternating with locally restricted secondary contact and introgression between incompletely isolated taxa. In contrast, the deepwater habitat is characterized by weak geographic structure and a high potential for gene flow, which may explain the lower species richness of deepwater than littoral lineages. For the same reason, divergent deepwater lineages should have evolved strong intrinsic reproductive isolation already in the incipient stages of diversification, and, consequently, hybridization among established lineages should have been less frequent than in littoral lineages. We test this hypothesis in the endemic Lake Tanganyika deepwater cichlid tribe Bathybatini by comparing phylogenetic trees of Hemibates and Bathybates species obtained with nuclear multilocus AFLP data with a phylogeny based on mitochondrial sequences. Consistent with our hypothesis, largely congruent tree topologies and negative tests for introgression provided no evidence for introgressive hybridization between the deepwater taxa. Together, the nuclear and mitochondrial data established a well-supported phylogeny and suggested ecological segregation during speciation.

Phylogenetic analysis identified haplochromine cichlids from isolated water bodies in the eastern branch of the East African Rift system as an ancient lineage separated from their western sister group in the course of the South Kenyan-North Tanzanian rift system formation. Within this lineage, the close phylogenetic relatedness among taxa indicates a recent common ancestry and historical connections between now separated water bodies. In connection with a total lack of local genetic diversity attributable to population bottlenecks, the data suggest cycles of extinction and colonization in the unstable habitat provided by the small lakes and rivers in this geologically highly active area., (© 2011 The Authors. Journal of Fish Biology © 2011 The Fisheries Society of the British Isles.)

The conditions of phenotypic and genetic population differentiation allow inferences about the evolution, preservation and loss of biological diversity. In Lake Tanganyika, water level fluctuations are assumed to have had a major impact on the evolution of stenotopic littoral species, though this hypothesis has not been specifically examined so far. The present study investigates whether subtly differentiated colour patterns of adjacent Tropheus moorii populations are maintained in isolation or in the face of continuous gene flow, and whether the presumed influence of water level fluctuations on lacustrine cichlids can be demonstrated in the small-scale population structure of the strictly stenotopic, littoral Tropheus. Distinct population differentiation was found even across short geographic distances and minor habitat barriers. Population splitting chronology and demographic histories comply with our expectation of old and rather stable populations on steeper sloping shore, and more recently established populations in a shallower region. Moreover, population expansions seem to coincide with lake level rises in the wake of Late Pleistocene megadroughts ~100 KYA. The imprint of hydrologic events on current population structure in the absence of ongoing gene flow suggests that phenotypic differentiation among proximate Tropheus populations evolves and persists in genetic isolation. Sporadic gene flow is effected by lake level fluctuations following climate changes and controlled by the persistence of habitat barriers during lake level changes. Since similar demographic patterns were previously reported for Lake Malawi cichlids, our data furthermore strengthen the hypothesis that major climatic events synchronized facets of cichlid evolution across the East African Great Lakes., (© 2011 Blackwell Publishing Ltd.)

Cichlid fishes of Lake Tanganyika display a variety of mating and parental care behaviors, including polygamous and monogamous mouthbrooding and substrate breeding, cooperative breeding, as well as various alternative reproductive tactics such as sneaking and piracy. Moreover, reproductive behaviors sometimes vary within species both in space and in time. Here, I survey reports on mating and parenting behaviors of Lake Tanganyika cichlid species and address the evolution of mating and parental care patterns and sexual dimorphism. Notes on measures of sexual selection intensity and the difficulties of defining mating systems and estimating selection intensities at species level conclude the essay.