Invisible pair bonds detected by molecular analyses

A focus on pair bonds between males and females is fundamental to study the evolution of social organization. Because pair bonds are generally identified from direct observations of pairs that maintain physical proximity, pair bonds may have been overlooked in animals that do not exhibit such visible pairs. The Lake Tanganyika cichlid fish Xenotilapia rotundiventralis forms schools that consist of mouthbrooding and non-brooding adults in mid-water, and visible pairs are not recognized. A previous study suggested that mouthbrooding females transfer fractions of the young to males when the young become large. However, it remains a mystery whether the mating pairs maintain pair bonds so that the females can transfer the young to their mates. To answer this question, we conducted a parentage analysis using 10 microsatellite markers. The analysis showed that the mouthbrooding adults were most likely genetic fathers and mothers of the young in their mouths. This finding suggests that the female-to-male shift of young takes place between mating partners, and thus the mating pairs maintain pair bonds at least until the shift of young. The present study is the first to detect pair bonds in animals in which physical proximity has not been observed.     

Field studies of the behavioral ecology and agonistic behavior of Cichlasoma meeki (Pisces: Cichlidae)

Synopsis The behavior and ecology of the firemouth cichlid were investigated in southern Mexico. Observations were conducted primarily at Laguna Bacalar, Quintana Roo. The fish bred throughout the year, nesting in small rocky crevices. Territories were established either by wandering pairs or by lone males which subsequently attracted mates. Stationary territories were maintained for two to three weeks by both pair members until the young were free-swimming, after which the parents defended the young as they foraged. Brood care lasted approximately three months. The distances at which fish were attacked and the behavior performed varied with the age of the defended young and the sex of the parent, as well as the species and size of the intruder. The sexes differed in their reproductive and agonistic activities. Predation on young was rapid if the parents were diverted from their defense. Serious injuries to adults were not observed to result from intraspecific contests. Such interactions were very common, but were ritualized with physical contact between animals rare. Color patterning varied with breeding interval, sex and behavior.     

Variations in age and size at maturity of female Nile tilapia, Oreochromis niloticus, populations from man-made lakes of Côte d'Ivoire

The reproductive characteristics of Oreochromis niloticus females were studied in two large hydroelectric dams and 6 small agropastoral reservoirs of Côte d'Ivoire, selected for their diversity of environmental conditions and sizes (6 to 80000 ha). Comparative analysis of age and size at maturity revealed large differences between the populations with early maturity, late maturity, and intermediate situations. All the populations matured in their first year, between 5.6 and 10 months. Age and size at maturity were positively correlated with the reservoir area. The range of variation in age at maturity between populations was far greater than that of size at maturity. However, size at maturity also appears to be very plastic as intra-population variations of 2 to 3 cm were found between consecutive years. A comparison of age at maturity for a population in 1994 and 1996 showed that this trait is also likely to vary significantly at the intra-population level. Environmental factors which could potentially affect age and size at maturity are discussed. Growth differences are the probable explanation for the observed patterns of variation between populations. Rapid changes in age and size at maturity at inter or intra-population levels suggest that the observed variations might be explained by differences in environmental variables (phenotypic plasticity) rather than by genetic differences.     

Evolution of the tribe Tropheini from Lake Tanganyika: synchronized explosive speciation producing multiple evolutionary parallelism

One of the most surprising outcomes of recent molecular studies on cichlid fishes of the three Great East African Lakes Victoria, Malawi and Tanganyika, was the stunning rapidity of speciation and cladogenesis at early stages of adaptive radiation. Despite their rapid pace, speciation events were so far intuitively assumed to proceed in a bifurcating and tree-like fashion, even if they could not be resolved by gene phylogenies due to a lack of resolution. On the basis of phylogenetic analyses of the Tropheini, a lineage of endemic rock-dwelling cichlid fishes from Lake Tanganyika, we suggest a pathway of explosive speciation that accounts for a non-bifurcating manner of cladogenesis. This pattern is likely to be the result of the contemporaneous origin of a multitude of founder populations in geographically isolated rock habitats among which gene flow was interrupted simultaneously by a major change of the lake habitat in the form of a rapid rise of the lake level. As a consequence, all new species arising from that vicariance event must exhibit almost equal genetic distances to each other, within the scope of genetic diversity of the founder population(s), even if the actual processes of subsequent speciation and eco-morphological diversification followed independent routes. Our phylogeny also suggests a high frequency of parallel evolution of equivalent trophic specialization in the Tropheini. This phenomenon seems to be an inherent feature of this pathway of speciation, due to the action of similar selective forces on the same set of species colonizing isolated habitats of the same type. Explosive speciation via synchronization of genetic divergence triggered by rapid environmental changes seems to be particularly likely to occur at advanced stages of adaptive radiation, when species are already adapted to particular habitats and have a reduced ability for dispersal.     

The role of sex ratios and resource availability on the pre-mating behavior of a monogamous fish

Convict cichlid fish (Archocentrus nigrofasciatus) are one of the few monogamous species where the female appears more colorful than the male. We examined whether this sexual dimorphism was reflected in a female-biased actual sex ratio (ASR) and whether convict cichlid females would exhibit behavioral traits typified in monogamous males when they possessed the extravagant colors and/or morphology. Our field observations revealed that females are marginally, but not significantly, more abundant than males but they did initiate more intersexual social interactions (i.e. approach behavior) than males. They also showed more intrasexual aggression. Furthermore, males more commonly chased females rather than vice versa. Our laboratory experiment also indicated that differences in intrasexual aggression were not related to differences in the ASR but did appear with the addition of a breeding site. Thus, while intrasexual competition was more frequently observed between females in the field it was probably related to the availability of breeding sites rather than our estimates of the ASR.     

Elevated mtDNA diversity in introduced populations of Cynotilapia afra (Günther 1894) in Lake Malawi National Park is evidence for multiple source populations and hybridization

Genetic variation in many invasive species shows little or no signs of a founder event, suggesting that high genetic diversity may facilitate establishment success. The rocky‐shore, plankton‐feeding cichlid fish Cynotilapia afra is endemic to Lake Malawi, but naturally absent from many suitable sites. In the 1960s, this species was introduced to the southern areas of the lake, presumably as a result of the aquarium fish trade. It has now become established on a number of rocky areas within the Lake Malawi National Park. Here, we analysed DNA sequence variation in the mitochondrial control region of six native and four introduced populations of C. afra, and three populations of the closely‐related and hybridizing Pseudotropheus zebra. In contrast to previous studies of Lake Malawi rock dwelling cichlids, network analyses suggested that native populations of C. afra showed high levels of lineage sorting in mtDNA. Introduced populations showed higher sequence and haplotype diversity than their native counterparts. Our analyses suggested that the elevated gene diversity was largely attributed to the fact that the introduced C. afra populations were derived from several genetically distinct and geographically separate populations, and to a lesser extent because of introgressive hybridization with native P. zebra. The establishment and spread of C. afra may be partly because of its ability to occupy a vacant ecological niche, but it may also have been facilitated by its enhanced genetic diversity.     

Social fishes and single mothers: brain evolution in African cichlids

As with any organ, differences in brain size--after adequate control of allometry--are assumed to be a response to selection. With over 200 species and an astonishing diversity in niche preferences and social organization, Tanganyikan cichlids present an excellent opportunity to study brain evolution. We used phylogenetic comparative analyses of sexed adults from 39 Tanganyikan cichlid species in a multiple regression framework to investigate the influence of ecology, sexual selection and parental care patterns on whole brain size, as well as to analyse sex-specific effects. First, using species-specific measures, we analysed the influence of diet, habitat, form of care (mouthbrooding or substrate guarding), care type (biparental or female only) and intensity of sexual selection on brain size, while controlling for body size. Then, we repeated the analyses for male and female brain size separately. Type of diet and care type were significantly correlated with whole brain size. Sex-specific analyses showed that female brain size correlated significantly with care type while male brain size was uncorrelated with care type. Our results suggest that more complex social interactions associated with diet select for larger brains and further that the burden of uniparental care exerts high cognitive demands on females.     

Adult plasticity in African cichlids: Rapid changes in opsin expression in response to environmental light differences

Phenotypic plasticity allows organisms to adapt quickly to local environmental conditions and could facilitate adaptive radiations. Cichlids have recently undergone an adaptive radiation in Lake Malawi where they inhabit diverse light environments and tune their visual sensitivity through differences in cone opsin expression. While cichlid opsin expression is known to be plastic over development, whether adults remain plastic is unknown. Adult plasticity in visual tuning could play a role in cichlid radiations by enabling survival in changing environments and facilitating invasion into novel environments. Here we examine the existence of and temporal changes in adult visual plasticity of two closely related species. In complementary experiments, wild adult Metriaclima mbenji from Lake Malawi were moved to the lab under UV‐deficient fluorescent lighting; while lab raised M. benetos were placed under UV‐rich lighting designed to mimic light conditions in the wild. Surprisingly, adult cichlids in both experiments showed significant changes in the expression of the UV‐sensitive single cone opsin, SWS1, in only 3 days. Modeling quantum catches in the light environments revealed a possible link between the light available to the SWS1 visual pigment and SWS1 expression. We conclude that adult cichlids can undergo rapid and significant changes in opsin expression in response to environmental light shifts that are relevant to their habitat and evolutionary history in Lake Malawi. This could have contributed to the rapid divergence characteristic of these fantastic fishes.     

Aggressive biases towards similarly coloured males in Lake Malawi cichlid fishes

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