It is often assumed that the mutation rate is an evolutionarily optimized property of a taxon. The relevant mutation rate is for mutations that affect fitness, U, but the strength of selection on the mutation rate depends on the average effect of a mutation. Determination of U is complicated by the possibility that mutational effects depend on the particular environmental context in which the organism exists. It has been suggested that the effects of deleterious mutations are typically magnified in stressful environments, but most studies confound genotype with environment, so it is unclear to what extent environmental specificity of mutations is specific to a particular starting genotype. We report a study designed to separate effects of species, genotype, and environment on the degradation of fitness resulting from new mutations. Mutations accumulated for .200 generations at 20°in two strains of two species of nematodes that differ in thermal sensitivity. Caenorhabditis briggsae and C. elegans have similar demography at 20°, but C. elegans suffers markedly reduced fitness at 25°. We find little evidence that mutational properties differ depending on environmental conditions and mutational correlations between environments are close to those expected if effects were identical in both environments.T HE importance of deleterious mutations to the evolutionary process is well appreciated (Morgan 1903;Haldane 1927;Fisher 1930;Sturtevant 1937;Kondrashov 1988), and much effort has been expended in understanding the processes by which new mutations arise and their effects on the phenotype and on fitness (reviewed by Simmons and Crow 1977;Drake et al. 1998;Keightley and Eyre-Walker 1999;Lynch et al. 1999;Houle and Kondrashov 2006). Drake, especially, has emphasized the remarkable consistency of the per-genome mutation rate across very broad taxonomic categories, but has also noted that there is considerable variation within those broad taxa (e.g., Drake et al. 1998). The idea that certain mutational properties vary between related species and even within species is venerable (Sturtevant 1937 and references therein), but there is as yet nothing approaching a comprehensive understanding of the variation in mutational properties-rate, distribution of effects, environmental sensitivity, molecular spectrum-of any species or group of closely related species, with the possible exception of the bacterium Escherichia coli (Matic et al. 1997;Sniegowski et al. 1997;Bjedov et al. 2003).An intriguing but almost completely unsubstantiated possibility (but see Nöthel 1987;Bjedov et al. 2003) is that mutation rates are themselves an evolutionarily optimized property (Fisher 1930;Sturtevant 1937;Kimura 1960Kimura , 1967Leigh 1970Leigh , 1973 Kondrashov 1995a,b;Dawson 1998). Because the vast majority of mutations with observable effects are deleterious, it is generally accepted that direct selection (almost) always favors a reduction in the mutation rate, with an optimal mutation rate of zero, at least in sexual taxa (e.g., Leig...