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<br />INTROGRESSION IN COLORADO RIVER CATOSTOMIDS <br /> <br />Razorback sucker. - Eleven of 2 I loci were <br />polymorphic in at least one of the samples of ra- <br />zorback sucker (Table J). In tests for conformance <br />with Hardy-Weinberg equilibrium expectations, <br />all but one sample lacked significance (P > 0.05); <br />the genotype array at the S-Mdh-A! locus in the <br />Dexter Hatchery sample exhibited a significant <br />departure (P < 0.00 I) from equilibrium expec- <br />tations. This deviation was the result of a defi- <br />ciency of heterozygotes and an excess in both ho- <br />mozygote categories. Both of the other samples <br />(la and 2) contained this diallelic polymorphism <br />at this locus but both yielded genotype arrays con- <br />forming to equilibrium expectations. Measures of <br />genetic variability were comparable among the <br />samples of razorback suckers (Table 2). <br />The contingency chi-square analysis of samples <br />la and 2 revealed agreement with equilibrium <br />conditions for all polymorphic loci. No genetic <br />differentiation was calculated by Nei's (1978) <br />method between the two native samples of razor- <br />back sucker and between the hatchery sample and <br />either of these native samples; i.e., 1= 1.000, and <br />D = 0.000. Positive mean F-statistics for the na- <br />tive samples (Table 4) indicated a small degree of <br />overall heterozygote deficiency within and be- <br />tween samples (mostly due to S-Mdh-A'). An FST <br />value of 0.011 calculated between the nalive sam- <br />ples is considered to reflect "little" differentiation <br />(sensu Hartl 1981). <br /> <br />Interspecific Differentiation <br /> <br />Divergence between species is often marked by <br />complete allelic differentiation at one or more loci <br />(A vise 1976). These "marker" loci can be ex- <br />tremely useful in studies of biochemical identifi- <br />cation and hybridization. Four loci, Ck-A. G6pi- <br />A!. M-Icdh-A. and S-Sod-A!. exhibited allelic <br />distributions that were species-specific, or nearly <br />so, and could be used to separate flannelmouth <br />suckers from razorback suckers. In addition, many <br />population-specific minor alleles at other loci <br />served to distinguish these taxa (Table I). The <br />average unbiased genetic distance and genetic <br />identity coefficients (Nei 1978) computed between <br />flannelmouth sucker and razorback sucker were <br />D = 0.220 and 1= 0.802, respectively. <br />The primary interest of this study was to ex- <br />amine flannelmouth and razorback suckers for dif- <br />ferences in gene expression as well as for differ- <br />ences in allelic composition at homologous loci. <br />The catostomid fishes are ancestrally tetraploid <br />(Uyeno and Smith J 972) and would, in theory, be <br />expected to express duplications (beyond the dip- <br /> <br />107 <br /> <br />loid condition) at all loci (Ohno 1970) whose <br />products can be resolved via electrophoretic <br />methods. However, with multiple (duplicated) loci, <br />one is free to vary (to acquire mutatio~s that might <br />have been selected against in the single-locus con- <br />dition and may even be "silenced" or "turned off" <br />in a regulatory sense) as long as the other locus <br />remains functional (Ohno 1970). Thus, ancestral- <br />ly tetraploid catostomid species can differ in the <br />number of genes controlling given enzyme sys- <br />tems (e.g., Ferris and Whitt 1977). Such differ- <br />ences have been used in phylogenetic studies (Fer- <br />ris and Whitt 1978b; Buth 1979, 1980). However, <br />the number of genes controlling the enzyme sys- <br />tems examined proved to be identical for razor- <br />back and flannel mouth suckers, eliminating this <br />character as one useful for the identification of <br />hybrids. <br /> <br />Assessment of Possible Introgression <br /> <br />Allozymic evidence for introgressive hybridiza- <br />tion between flannefmouth suckers and razorback <br />suckers can be found in the examination of the <br />distribution of the gene products of those marker <br />loci previously discussed. Two of these loci, Ck-A <br />and M-/cdh-A. have allelic distributions suggest- <br />ing some gene flow between the two species. In- <br />traspecific variation involving creatine kinase (Ck- <br />A) is rare (Ferris and Whitt 1978a) and the reso- <br />lution of Ck-A heterozygosity in both species in <br />this study would not be expected. Both alleles are <br />present in both species and the predominant allele <br />in each species (e.g., Ck-A[aJ in the flannel mouth <br />sucker) is present in low frequency in the other <br />species. There is, of course, the possibility that <br />both alleles at both loci were present in the com- <br />mon ancestor of these catostomids and that they <br />have drifted differentially or been selected for, al- <br />though not to fixation, in these two species. The <br />hypothesis of the sharing of these alleles as prim- <br />itive traits can only be addressed through the ex- <br />amination of additional, closely related, species of <br />Catostomus (see Murphy et al. 1984). The situa- <br />tion involving M-Icdh-A is somewhat similar and <br />is summarized in Table I. <br />If introgressive hybridization is accepted as the <br />means to account for the sharing of the other <br />species' alleles at Ck-A and M-/cdh-A. one can <br />observe that gene flow is proceeding both from <br />flannelmouth to razorback sucker (with M-/cdh-A <br />favored) and from razorback to flannelmouth <br />sucker (with Ck-A favored). Here, "favored" <br />means that an introgessed allele remains in the <br />new gene pool while other "nonfavored" alleles <br />