RESUMO
The magnitudes of phenotypic variances in peripherally isolated populations of common chaffinches (Fringilla coelebs) in the Azores, Madeira, and Canaries relative to their continental source populations in Iberia and Morocco have played a pivotal role in the formulation of competing hypotheses of phenotypic evolution in these isolates. Because Van Valen's niche variation hypothesis and Grant's model of island evolution were based on museum skins prone to measurement error and temporal variation, we re-examined the patterns of phenotypic variability using more precise skeletal measurements from freshly collected specimens. Levene's tests showed that univariate character variances were homogeneous in all island and continental populations, although there was a consistent trend for the magnitude of the variances to be lower for all characters in all Canary island populations. Multivariate Levene's tests, however, revealed significantly reduced total variances in the Hierro and Madeira populations compared to some Azores and continental populations. The Azores and continental populations did not differ in variability, and lower variances in the Canaries were not related to the presence or absence of the congeneric blue chaffinch (F. teydea), contrary to the predictions of Van Valen's niche variation hypothesis. Population variability was not inversely related to differentiation or isolation within the Azores or Canaries archipelagoes, opposite to the association reported by Grant. Our results also differ from both previous studies which reported much larger differences in population variabilities, and this likely reflects the use of heterogeneous samples of museum specimens, less precise external characters, and the use of tests sensitive to sample size. Differentiation among populations has been markedly greater in the Canary islands, implicating founder events and possibly historically stronger directional selection as determinants of this enhanced divergence relative to the Azores. These variance-reducing processes are unlikely to explain current lower levels of phenotypic variability because there has been sufficient time since colonization for replenishment of variability in polygenic characters. Average heterozygosities at putatively neutral allozyme loci are 1.6 times higher in the Azores compared to the Canaries, and support the view that effective population sizes are smaller in the latter archipelago. We argue that reduced variances in the Canary island populations represent lower equilibrium levels maintained by drift and mutation in populations with smaller long-term effective sizes, consistent with Lynch and Hill's neutral model of phenotypic evolution. Although episodes of selection in the past may have been interspersed with long periods of effective neutralism and drift, adaptationist hypotheses invoking a primary role for variance-reducing selection appear to be unwarranted.
RESUMO
Peripherally isolated populations of common chaffinches (Fringilla coelebs) in the Canaries, Madeira, and Azores were compared genetically with their putative ancestral stock in Iberia and Morocco, and with a population of blue chaffinches (F. teydea) from Tenerife, using protein electrophoresis of 42 loci. The continental populations are only weakly differentiated genetically (FST = 0.092), despite distinctive subspecific differences in plumage and morphometrics between Iberia and Morocco populations. Estimated levels of gene flow among continental populations are high enough to account for their relative genetic homogeneity, and it is unlikely that homogenizing selection is operating to mimic the effects of gene flow. In contrast, the Atlantic island populations are well differentiated genetically (FST = 0.321), and have diverged considerably from their continental conspecifics. The development of significant genetic differentiation within the Canaries but not the Azores likely results from smaller population sizes, very restricted gene flow, and enhanced random drift in the former populations. There is no convincing evidence in support of stronger directional selection acting on genotypes or phenotypes to reduce within-population variability in Canaries populations as proposed by Grant (1979), although other tenets of his model of island evolution are supported by our analysis. Although genetic variability is reduced in four of the Canaries populations, only the Hierro population appears to have encountered a severe bottleneck. Yet it has not differentiated markedly from the La Palma population to which it is subspecifically allied. We conclude that gradual divergence in isolated populations of small to moderate size is the most plausible explanation for the evolution of intraspecific and interspecific diversity in Atlantic island chaffinches.
