Evolutionary biology: evidence for sympatric speciation?

Sympatric speciation is difficult to demonstrate in nature and remains a hotly debated issue. Barluenga et al. present a case of putative sympatric speciation for two cichlid species in the Nicaraguan crater lake Apoyo, but they overlook or reinterpret some key published information on the system. Although sympatric speciation is possible in theory, we show here that, when this information is taken into account, the results of Barluenga et al. do not provide conclusive evidence for sympatric speciation: this is because the null hypothesis of multiple invasion with introgression cannot be rejected.

Evolution of NADH dehydrogenase subunit 2 in east African cichlid fish.

The complete sequence (1047 bp) of the mitochondrially encoded ND2 gene was obtained from 31 species of cichlid fishes to investigate the evolutionary history of the species flocks of the East African lakes. The observed pattern of nucleotide substitution in this sequence is typical of mitochondrial genes, showing a high transition bias and rapid mutational saturation, especially at codon positions where base frequencies are unequal. The base composition of the third position of codons is heterogeneous among species, suggesting frequent shifts in the pattern of substitution. Phylogenetic analysis of the sequences shows that the mtDNA variation in Lake Malawi cichlids is nested monophyletically within the range of variation shown by Tanganyikan cichlids. The closest Tanganyikan relatives of the Malawi flock are members of the tribe Tropheini. Classifications based on morphology are generally supported by the mtDNA data, with some significant exceptions in the Tropheini and Lamprologini. Because of an apparently rapid radiation of the Tanganyikan lineages, it is difficult to assess the basal topology of the Tanganyikan radiation at this time. Divergences among tribes are consistent with an intralacustrine radiation within the past 10 million years.

Similar morphologies of cichlid fish in Lakes Tanganyika and Malawi are due to convergence.

The species flocks of cichlid fishes in the lakes of East Africa are the most spectacular example of adaptive radiation among living vertebrates. Similar highly derived morphologies are found among species in different lakes. These similarities have been variously interpreted either as evidence for migration of ancestral species between the lakes, or of striking convergence of morphology. To distinguish among these competing hypotheses we sequenced a portion of the mitochondrial DNA control region from six pairs of morphologically similar taxa from Lakes Malawi and Tanganyika. Our results clearly indicate a separate origin of these morphologies in the two lakes. They also suggest that the Tanganyikan radiation is relatively old, and that the species flocks in lakes Malawi and Victoria are derived from one of the ancient lineages found in Lake Tanganyika. These findings have important implications for understanding the pattern of morphological and behavioral evolution in these fish.

Fish predation as a factor in the distribution of Lake Malawi gastropods.

Gastropod densities in Lake Malawi, Africa drop dramatically below 4.5 m water depth. Weed beds disappear and the number of molluscivorous cichlids increases below this depth. When molluscivores were excluded from open sand habitat by cages, due to migration and reduced predation, the density of snails increased by 40-60% within a week. Alternatively, when adult Cyrtocara placodon were placed into the cages, snail density equaled the controls. The molluscivores disproportionately consumed snails of the genus Bulinus relative to those of the more heavily armoured genus, Melanoides. Cichlid molluscivores are hypothesized to be responsible for preventing the thin shelled bilharzia vector snail, Bulinus globosus, from successfully invading the open shore areas of Lake Malawi. Lake Malawi may be relatively free of this human disease because of snail predation by cichlids.

The effect of luminance upon the distribution and behavior of the Eleotrid fish Gobiomorus dormitor, and its prey.

Gobiomorus dormitor success in capturing prey, cichlids, poeciliids and atherinids, is correlated with low luminance. Both prey and predator populations inhabiting rocky areas in Lake Jiloa, Nicaragua shift to deeper water during the dry season when luminance is greater. Predator and prey populations are also deeper over bright sandy areas than over rocks. Predators, during the middle of the day, attack prey significantly less in water above 3 m than in water between 3-12 m, but there is a significant increase in the number of attacks occurring above 3 m at twilight. The predator becomes more active, and the prey disappear into holes at twilight. These data suggest that conclusions concerning the importance of light in structuring the community and behavior of marine fishes can be generalized to tropical freshwater systems.