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<br />108 <br /> <br />RUTH ET AL. <br /> <br />TABLE 5.-Comparative measures of presumably in- <br />trogressed genes in flannel mouth and razorback suckers, <br />based on relevant loci (Ck-A and M-Icdh-A). <br /> <br />Locality <br /> <br />Flannelmouth sucker <br /> <br />(Theoretical maximum) <br />Virgin River <br />Paria River <br />Little Colorado River <br />Upper Colorado River <br /> <br />Proportion <br />of genes <br />from ftan- <br />nelmouth <br />sucker <br /> <br />(1.000) <br />0.950 <br />1.000 <br />1.000 <br />0.993 <br /> <br />Razorback sucker <br /> <br />(Theoretical maximum) <br />Lake Mohave (adults, biased sample) <br />Lake Mohave (juveniles. unbiased sample) <br />Senator Wash Reservoir <br />Dexler Hatchery <br /> <br />(0.000) <br />0.02& <br />0.015 <br />0.000 <br />0.010 <br /> <br />are eliminated, even though all were surely ex- <br />changed in the initial hybridization event. Evi- <br />dence for introgression in flannelmouth sucker is <br />found both in the upper Colorado River sample <br />and in a sample (Virgin River) from the lower <br />Colorado (Table 5). However, although introgres- <br />sive hybridization may be an ongoing biological <br />interaction between these two species, it appar- <br />ently is rare enough for the species to maintain <br />the integrity of their respective gene pools. A sim- <br />ilar si.tuation has recently been reported for the <br />sympatric common shiner Notropis cornU/us and <br />striped shiner N. chrysocephalus by Dowling and <br />Moore (1984). No F, hybrids were identified in <br />any sample of razorback sucker because these in- <br />dividuals would be expected to be heterozygous <br />at all four marker loci: Ck-A, G6pi-Al, M-fcdh-A, <br />and S-Sod-Al. Introgressed individuals are, at best, <br />the products ofF2 backcrosses. The question might <br />be raised as to why the species-specific allelic dis- <br />tributions at G6pi-Al and S-Sod-Al do not also <br />supply some evidence of introgressive hybridiza- <br />tion as do Ck-A and M-fcdh-A. Selection against <br />the hybrid heterozygous conditions at the former <br />two loci may be the reason. However, we caution <br />against overinterpretation of this possibility given <br />the rarity of Ck-A and M-fcdh-A heterozygotes in <br />both flannelmouth and razorback suckers; it is <br />possible that variation of introgressive origin at <br />G6pi-Al and S-Sod-Al was not observed because <br />of the relatively small number of specimens ex- <br />amined. <br />In summary, although we cannot rule out the <br />possibility that rare marker alleles were present in <br /> <br />the common ancestor of these species, we believe <br />the data are in accord with the hypothesis of in- <br />trogressive hybridization. However, the frequency <br />of introgressive hybridization is quite low. If the <br />introgressive interaction between these:two catos- <br />tomid species is related to habitat medification <br />rather than to simple variation in the complete- <br />ness of reproductive isolating mechanisms, future <br />increased measures of introgression might be ex- <br />pected as the Colorado River system experiences <br />continued modification. <br />The Lake Mohave sample (I a) used in this study <br />was not randomly selected. Many specimens were <br />available and those suspected of being hybrids of <br />razorback and flannel mouth suckers, based on their <br />morphology, were selected for study. Our allo- <br />zyme findings thus raise three questions. (I) Is the <br />morphology of razorback suckers a poor indicator <br />of introgression? (2) Are introgressed morpholog- <br />ical and allozyme characters decoupled and dif- <br />ferentially selected? (3) Is introgression in Lake <br />Mohave at such a low level that the individuals <br />selected for study provide the only evidence for <br />this kind of genetic interaction? The first two ques- <br />tions can be addressed with some difficulty via <br />long-term breeding experiments. The more per- <br />tinent third question is addressed by the allozyme <br />data from the larger, randomly sampled collection <br />of juvenile razorback suckers from the same lo- <br />cality (sample 1 b; Table 5). If introgression, as <br />measured via allozyme expression, is indeed rare <br />in Lake Mohave, parental stock could be taken <br />with a random selection strategy without the need <br />of electrophoretic screening of such stock. <br /> <br />Evaluation of Current Hatchery Stock <br /> <br />Genes presumed to be introgressed from tlan- <br />nelmouth suckers are present at very low levels in <br />razorback suckers in both the hatchery stock and <br />the samples from Lake Mohave, the source of <br />hatchery breeding stock (Table 5). The current <br />hatchery stock has not been appreciably affected <br />by sampling error in favor of introgressed genes <br />from tlannelmouth suckers. Indeed, the hatchery <br />stock has fewer introgressed genes than the pre- <br />sumed parental stock from Lake Mohave. Thus, <br />in terms of introgressed genes, the hatchery stock <br />is at least as "good" as, if not "better" than the <br />Lake Mohave natural population. <br />The maintenance of variability in refugium <br />populations or hatchery stocks has been discussed <br />recently by Turner (1984) and Meffe (1986). Ge- <br />netic variability in razorback suckers (Table 3) <br />might be viewed as low when compared to the <br />