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<br />DISCRIMINATION OF COLORADO PLATEAU GILA SPP. <br /> <br />169 <br /> <br />than G. elegans. It is also worth noting that depth <br />of frontal depression (a direct measure of nuchal <br />concavity) was not a major component of species <br />segregation. However, a more indirect measure <br />(head depth at occiput) was. <br />Gila robusta and G. cypha have been a focus for <br />the majority of morphometric research accom- <br />plished on this group to date. Douglas et al. (1989) <br />used qualitative characteristics to separate these <br />species in the Yampa River. Kaeding et al. (1990) <br />compared field identification at Black Rocks, Col- <br />orado, with taxonomic assignment based on a prin- <br />cipal component analysis (PCA). In the former, the <br />complex relationships between anal fin and caudal <br />peduncle, eye and jaw, as well as snout, occiput, <br />and breast-nuchal scalation, were deemed impor- <br />tant; in the latter, the most important characters <br />were depth of nuchal depression and caudal pe- <br />duncle depth. McElroy and Douglas (1995) ex- <br />amined population-level differentiation of these <br />species from eight sites in the upper Colorado Riv- <br />er basin, and McElroy et al. (1997) differentiated <br />the two species on four morphometric characters <br />(eye diameter, caudal peduncle, orbit-jaw relation, <br />and skull depression-nuchal hump). <br />In this study, morphometric characters (Table I) <br />best segregated the two species, and depth of fron- <br />tal depression was an important character, which <br />was also the case for Kaeding et al. (1990) and <br />McElroy et al. (1997). A similar level of discrim- <br />ination is maintained in the field subset, even <br />though the former character was removed (albeit <br />with the addition of two additional morphometric <br />characters). However, other morphometric char- <br />acters listed in Table 1 have not been previously <br />noted as discriminatory. For example, two of four <br />field characters (anal fin base and head depth at <br />eye) relate more to qualitative characters depicted <br />in Douglas et al. (1989: Figure 2) than to any oth- <br />ers. Two remaining field characters (dorsal fin base <br />and pectoral insertion to pelvic insertion) have not <br />been previously identified as important. Their des- <br />ignation supports Douglas et al. (1989), who ar- <br />gued for application of new approaches and new <br />characters in resolving this complex problem. <br />Smith et al. (1979) evaluated phenotypic rela- <br />tionships among all three study species using PCA <br />and noted that characters best contributing to their <br />separation were dorsal and anal rays, gill raker and <br />vertebral number, and depth and length of caudal <br />peduncle. Those meristic characters indicated by <br />Smith et al. (1979) also discriminated well in our <br />analyses (Table 1). In addition, caudal peduncle <br />depth was an influential morphometric character, <br /> <br />as were several others not mentioned by these au- <br />thors. Discriminating field characters pertained to <br />the peduncle, its relation to the anal fin, the snout, <br />its relation to the occiput, and the dorsal fin base. <br />All three data sets discriminated greater than 95%. <br />It is clear from this study that a variety of char- <br />acters differ significantly among these species, and <br />various combinations of characters provide excel- <br />lent discrimination. The high kappa value (P < <br />0.00001) and failure of the jackknife to improve <br />the original segregation underscore this point. Fur- <br />thermore, misclassified individuals, both within <br />each species-pair and among all three species, <br />were rarely shared among the three data subsets, <br />again suggesting that a variety of characters ade- <br />quately discriminate. We thus reject the hypothesis <br />that the three species are phenotypically indistin- <br />guishable. <br />It is also clear that G. elegans is the most dif- <br />ficult of the three to assign to group. Of four mis- <br />identified individuals in the discriminant analysis <br />with field data, three were G. elegans (Figure 2). <br />Similarly, when sample sizes were equilibrated <br />among species and within-group covariances were <br />used in the analysis, two (of three) misidentified <br />individuals were G. elegans. <br /> <br />Assignment of Juveniles and Putative Hybrids to <br />Species <br /> <br />Overall, assignment of juveniles to species <br />seemed reasonable, with 88% correctly allocated. <br />However, this percentage is misleading. The pre- <br />ponderance of juveniles were G. robusta (N = 106; <br />75%), and these were correctly assigned at greater <br />than 97%. Correct assignment of juveniles to G. <br />cypha and G. elegans were only 60% and 66%, <br />respectively. Furthermore, if one considers that <br />several correct assignments of juveniles to these <br />species were actually borderline with respect to <br />score, then the number of unambiguous assign- <br />ments is even weaker. This suggests that although <br />field characters (Table 1) may be satisfactory for <br />designating juvenile G. robusta to species, they <br />appear marginal at best in assigning juvenile G. <br />cypha and G. elegans. Quite possibly, the occiput <br />and nuchal hump have not as yet developed ade- <br />quately enough to serve as the diagnostic tool these <br />structures provide in adults (Table 1). Given these <br />mixed results, we cannot reject our second hy- <br />pothesis, that adult characters also segregate ju- <br />veniles. However, this does not mean that juveniles <br />of G. cypha and G. elegans cannot be differentiated <br />phenotypically. Composites of several different <br />quantitative characters may instead be useful; <br />