What, if anything, is a Tilapia?-mitochondrial ND2 phylogeny of tilapiines and the evolution of parental care systems in the African cichlid fishes.

Abstract

We estimated a novel phylogeny of tilapiine cichlid fish (an assemblage endemic to Africa and the Near East) within the African cichlid fishes on the basis of complete mitochondrial NADH dehydrogenase subunit 2 (ND2) gene sequences. The ND2 (1,047 bp) gene was sequenced in 39 tilapiine cichlids (38 species and 1 subspecies) and in an additional 14 nontilapiine cichlid species in order to evaluate the traditional morphologically based hypothesis of the respective monophyly of the tilapiine and haplochromine cichlid fish assemblages. The analyses included many additional cichlid lineages, not only the so-called tilapiines, but also lineages from Lake Tanganyika, east Africa, the Neotropics and an out-group from Madagascar with a wide range of parental care and mating systems. Our results suggest, in contrast to the historical morphology-based hypotheses from Regan (1920, 1922 ), Trewavas (1983), and Stiassny (1991), that the tilapiines do not form a monophyletic group because there is strong evidence that the genus Tilapia is not monophyletic but divided into at least five distinct groups. In contrast to this finding, an allozyme analysis of Pouyaud and Agnèse (1995), largely based on the same samples as used here, found a clustering of the Tilapia species into only two groups. This discrepancy is likely caused by the difference in resolution power of the two marker systems used. Our data suggest that only type species Tilapia sparrmanii Smith (1840) should retain the genus name TILAPIA: One particular group of tilapiines (composed of genera Sarotherodon, Oreochromis, Iranocichla, and Tristramella) is more closely related to an evolutionarily highly successful lineage, the haplochromine cichlids that compose the adaptive radiations of cichlid species flocks of east Africa. It appears that the highly adaptable biology of tilapiines is the ancestral state for all African cichlids and that the more stenotypic lifestyle of the haplochromine cichlids is derived from this condition. We reconstructed the evolution of the highly variable parental care systems on the basis of the most inclusive composite phylogeny to date of the African, Neotropical, and Madagascan cichlids with special emphasis on a group of tilapiines comprising the substrate-spawning genus Tilapia, and the mouthbrooding genera Sarotherodon and OREOCHROMIS: We demonstrate several independent origins of derived mouthbrooding behaviors in the family Cichlidae.