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The mean time to shared haplotype ancestry for mtDNA genes can-be derivecTsimilarly (Avise et al. 1988), but m theory is only one-fourth as large as for autosoma! nuclear genes, the differ­ ence being attributable to a twofold effect due to the haploid transmission of mtDNA and another twofold effect due to mtDNA's normal pattern of uni­ parental transmission. The above theory assumes that times to common ancestry for allelic pairs are independent Therefore, in interpreting empirical data for any particular species against these expectations (see Chapter 61, caution must be exercised : because the history of lineage coalescence within a real population imposes a severe correlation on pairwise comparisons (Ball et al.

1992). ). In another such meta-analysis of protein electrophoretic stud­ ies, Britten (1996) concluded that "selection, including overdominance, has at most a weak effect at allozyme loci, and [this] casts some doubt on the widely held notion that heterozygosity and individual fitness are strongly correlated/' This conclusion generally echoed a sentiment expressed 20 years earlier by Selander (1976): "Notwithstanding the immense amount of effort expended in surveying (allozyme) variation ... , the sample sizes of loci are generally inadequate for satisfactory analyses ...

A harmonic mean is closer to the smaller rather than to the larger of a series of numbers being averaged, so can be much lower than most population censuses. A severe reduction in population size, called a "popula­ tion bottleneck," can greatly depress evolutionary Ne. 3. Combination o f separate sexes and fluctuating population size: If census popula- vs; tion sizes of males and females are known across multiple generations, joint The History of Interest in Genetic Variation effects of the above factors on Ne can be.

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