Captive breeding for the conservation of cichlid fishes

Cichlid fishes are by far the largest familial group of endangered vertebrates, especially the haplochromines. This paper concerns the organization and management of captive breeding of haplochromine cichlids. The setting up of a small-scale laboratory programme for the conservation of endangered species is described in terms of funding, staffing, installation and livestock husbandry. Breeding is discussed in the context of the selection of broodstock, basic reproductive biology, rearing, disease, pathological disorders and the arrangements necessary for the transfer and documentation of progeny. There are already indications in Africa and elsewhere that the dramatic decline and demise of cichlid taxa in Lake Victoria will not be an isolated phenomenon. There is no prospect that the captive breeding of cichlids can alone resolve such large-scale problems in fisheries management and ecology, or prevent the loss of taxa in nature. Nevertheless, captive breeding provides conservation options which are otherwise limited or unavailable regarding the saving of individual 'heritage' species, restocking to the wild, fundamental laboratory research and, finally, public education on the grave issue of actual and prospective mass extinctions of cichlids and other rare fishes.     

Ecological and evolutionary consequences of the trophic polymorphism in Cichlasoma citrinellum (Pisces: Cichlidae)

The neotropical cichlid fish Cichlasoma citrinellum is polymorphic in the structure of its pharyngeal jaw apparatus and external morphology. The pharyngeal jaws are either gracile and bear slender, pointed teeth (papilliform) or robust with strong, rounded teeth (molariform). Molariform morphs have a ‘benthic’, and papilliform morphs a ‘limnetic’ body form. Furthermore, this species is also polychromatic, with yellow and black morphs. The molariform morphology of the pharyngeal jaw apparatus adapts the fish for cracking and feeding on snails. Based on analysis of stomach contents, 94% of the molariform morph ate snails whereas only 19%, of the papilliform morph did so. This result suggests that the morphs occupy different ecological niches. The morphology of the pharyngeal jaw apparatus does not correlate significantly with sex, but it does with body colouration (P<0.005). Cichlasoma citrinellum mate assortatively with their own colour; therefore a mating preference for colour may lead to genetic isolation of trophic morphs. The frequency of the molariform morph differs strikingly among populations of five Nicaraguan lakes and its abundance is correlated with the abundance of snails, the fishes' principal prey item. Among populations the frequency of molariform morphs decreases in the dry season. Morphology possibly changes reversibly within particular individuals between seasons. These results suggest that phenotypic plasticity and polymorphisms may be an adaptive characteristic of cichlid fishes. Patterns of intraspecific morphological variation match patterns of interspecific morphological diversification which suggests that universal developmental mechanisms canalize the possible expressions of morphology. The ability to respond morphologically to environmental shifts, in conjunction with genetically determined trophic polymorphisms and sexual selection via mate choice, could be the basis for speciation through intermediate stages of polymorphism of the impressive adaptive radiation of cichlid fishes.     

Morphological and temporal variation in early embryogenesis contributes to species divergence in Malawi cichlid fishes

The cichlid fishes comprise the largest extant vertebrate family and are the quintessential example of rapid “explosive” adaptive radiations and phenotypic diversification. Despite low genetic divergence, East African cichlids harbor a spectacular intra- and interspecific morphological diversity, including the hyper-variable, neural crest (NC)-derived traits such as coloration and craniofacial skeleton. Although the genetic and developmental basis of these phenotypes has been investigated, understanding of when, and specifically how early, in ontogeny species-specific differences emerge, remains limited. Since adult traits often originate during embryonic development, the processes of embryogenesis could serve as a potential source of species-specific variation. Consequently, we designed a staging system by which we compare the features of embryogenesis between three Malawi cichlid species—Astatotilapia calliptera, Tropheops sp. ‘mauve’ and Rhamphochromis sp. “chilingali”—representing a wide spectrum of variation in pigmentation and craniofacial morphologies. Our results showed fundamental differences in multiple aspects of embryogenesis that could underlie interspecific divergence in adult adaptive traits. First, we identified variation in the somite number and signatures of temporal variation, or heterochrony, in the rates of somite formation. The heterochrony was also evident within and between species throughout ontogeny, up to the juvenile stages. Finally, the identified interspecific differences in the development of pigmentation and craniofacial cartilages, present at the earliest stages of their overt formation, provide compelling evidence that the species-specific trajectories begin divergence during early embryogenesis, potentially during somitogenesis and NC development. Altogether, our results expand our understanding of fundamental cichlid biology and provide new insights into the developmental origins of vertebrate morphological diversity.     

The importance of carotenoids in signaling during aggressive interactions between male firemouth cichlids (Cichlasoma meeki)

Male firemouth cichlids, Cichlasoma meeki, have red pigmentationcovering large areas of their ventral surface, which is displayedduring aggressive interactions. We manipulated the levels ofred pigmentation by assigning the fish to one of two diets,which were as similar as possible except that one was high incarotenoids while the other was low in carotenoids. During diadictrials under white light, fish kept on the high carotenoid dietwon a higher proportion of contests than fish kept on the lowcarotenoid diet Under green light, where differences in rednesscannot be discriminated, there was no effect of diet on theoutcome of contests. These experiments demonstrate that it isthe effect of the diet on red pigmentation that is importantrather than some confounding variable such as differential growthrates. The weight of the two fish was also important; therewas a tendency for the heavier fish to win more contests. Themass effect was subordinate to color under white light but wasthe dominant factor under green light The nature of the contestsunder the different light conditions also varied; the displayin which the red pigmentation is most obvious was not used undergreen light, but was common under white light This suggeststhat the display strategies are flexible and can be alteredaccording to which displays are most effective in a given environmentPrevious studies of other species of fish and birds have shownthat the degree of redness influences mate choice and is affectedby parasite infestations. We propose that carotenoid pigmentationis likely to reflect a general quality, influenced by severalfactors, rather than a context-specific quality such as fightingability.     

Experimental analysis of some factors affecting parental expenditure and investment inCichlasoma nigrofasciatum (Cichlidae)

Synopsis Parental investment is the cost of providing parental care. The short-term cost of parental care was measured in the biparental substrate nesting cichlid,Cichlasoma nigrofasciatum, by comparing the expected future survival (measured indirectly as energy content of the body), time taken to breed again and, among females, the number of eggs produced at a subsequent spawning of parental and non-parental pairs. In comparison with non-parental pairs, parental pairs took significantly longer to respawn. Body condition and female fecundity were unaffected after a single parental cycle. The effect on parental cost and expenditure of factors likely to stress the parental fish was also investigated. Removing the male parent had no effect on female parental cost. Exposing pairs to potential predators of offspring increased the time taken by pairs to respawn. In parental males, reducing the level of feeding gave rise to a reduction in some care behaviours.     

Gene expression remodelling and immune response during adaptive divergence in an African cichlid fish

Variation in gene expression contributes to ecological speciation by facilitating population persistence in novel environments. Likewise, immune responses can be of relevance in speciation driven by adaptation to different environments. Previous studies examining gene expression differences between recently diverged ecotypes have often relied on only one pair of populations, targeted the expression of only a subset of genes or used wild‐caught individuals. Here, we investigated the contribution of habitat‐specific parasites and symbionts and the underlying immunological abilities of ecotype hosts to adaptive divergence in lake–river population pairs of the cichlid fish Astatotilapia burtoni. To shed light on the role of phenotypic plasticity in adaptive divergence, we compared parasite and microbiota communities, immune response, and gene expression patterns of fish from natural habitats and a lake‐like pond set‐up. In all investigated population pairs, lake fish were more heavily parasitized than river fish, in terms of both parasite taxon composition and infection abundance. The innate immune response in the wild was higher in lake than in river populations and was elevated in a river population exposed to lake parasites in the pond set‐up. Environmental differences between lake and river habitat and their distinct parasite communities have shaped differential gene expression, involving genes functioning in osmoregulation and immune response. Most changes in gene expression between lake and river samples in the wild and in the pond set‐up were based on a plastic response. Finally, gene expression and bacterial communities of wild‐caught individuals and individuals acclimatized to lake‐like pond conditions showed shifts underlying adaptive phenotypic plasticity.     

Species‐specific ontogenetic diet shifts among Neotropical Crenicichla: using stable isotopes and tissue stoichiometry

Ontogenetic diet shifts were compared among five sympatric pike cichlids Crenicichla in a subtropical South American stream using stable C and N isotopes and tissue stoichiometry (C:N). Within species, stable N isotopes were positively related to body size while C:N showed negative relationships. Stable C isotopes, however, were not related to body size in any species. By modelling the switch to piscivory using gut content‐isotope‐body size relationships, diet shifts were shown to be species‐specific with regard to both rate and degree of piscivory. Compared to other piscivorous lineages, Crenicichla appear to be unusually small‐bodied (based on maximum body size). Because of their diversity, abundance and dynamic size‐structured functional roles, Crenicichla may exert broad and complex predation pressures on the aquatic community.     

Genetic interactions controlling sex and color establish the potential for sexual conflict in Lake Malawi cichlid fishes

Sex-determining systems may evolve rapidly and contribute to lineage diversification. In fact, recent work has suggested an integral role of sex chromosome evolution in models of speciation. We use quantitative trait loci analysis of restriction site-associated DNA -tag single nucleotide polymorphisms to identify multiple loci responsible for sex determination and reproductively adaptive color phenotypes in Lake Malawi cichlids. We detect a complex epistatic sex system consisting of a major female heterogametic ZW locus on chromosome 5, two separate male heterogametic XY loci on chromosome 7, and two additional interacting loci on chromosomes 3 and 20. Our data support the known chromosomal linkage between orange blotch color and ZW, as well as novel genetic associations between male blue nuptial color and two sex determining regions (an XY and ZW locus). These results provide further empirical evidence for a complex antagonistic sex–color system in this species flock and suggest a possible role for, and effect of, polygenic sex-determining systems in rapid evolutionary diversification.