<br />DISCRIMINATION OF COLORADO PLATEAU GILA SPP.
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<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;
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