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<br />.. <br /> <br />. the Colorado and Green rivers. This conclusion <br />cannot be confidently applied to certain popula- <br />tions that we have available but have not fully <br />studied from Lake Powell, however. Furthermore, <br />populations of Gila robusta from certain tribu- <br />taries were elegans and cYP4a are absent parallel <br />these species in many characteristics. We choose <br />a taxonomic treatment of this situation that em- <br />phasizes the specific distinction of the three <br />populations in the big-river habitat, but admit <br />the possibility that the species isolating mecha- <br />nisms may break down under disturbed (reservoir) <br />conditions and that the populations in tributary <br />streams may not be completely independent or iso- <br />lated from the three central forms. We refer to <br />the entire complex as the Gila robusta super- <br />species, including robusta, cypha, and elegans <br />as defined above, and including subspecies or <br />races of robusta, seminuda of the Virgin River, a <br />similar population in the San Juan River, and a <br />number of isolated populations in northern and <br />western Mexico. The complex in the Gila River <br />basin includes Gila intermedia and forms inter- <br />mediate between that species and robusta, i.e., <br />G. r. "grahami" (cf. Rinne, 1976). Gila inter- <br />media is inferred to be a part of the robusta <br />super species on the basis of shared characters <br />and possible gene exchange through "grahami", but <br />may be a sister phyletic line by its origins, as <br />indicated by its similarity to several of the <br />large-scaled, small-finned subgenera of Gila (cf. <br />Miller, 1945). <br /> <br />In spite of the uncertainty regarding peripher- <br />al populations, it is clear that robusta (s.s.), <br />elegans, and cypha have diverged in a number of <br />characters, and can be discriminated on the basis <br />of counts of fin rays, vertebrae, and gill rakers, <br />the relative depth and length of the caudal pe- <br />duncle, snout shape, fin position, and form of the <br />nuchal hump. The multivariate analysis shows that <br />cypha is intermediate between robusta and elegans <br />in general, but is extreme in several respects. <br />This suggests the possibility that cypha and <br />elegans were derived from robusta by separate <br />speciation events. Gila robusta shows the most <br />primitive characters of the three, and the sepa- <br />rate ways in which elegans and cypha are extreme <br />suggest that neither is likely to be ancestral to <br />either of the other two. <br /> <br />T~ack of coexistence of the three species <br />in s ler tributaries is in accordance with the <br />expect relationship between diversity and spatial <br />hetero neity. The species seem to be omnivorous <br />carnivo es with specializations related to habitat <br />--chacters associated with food processing (jaws, <br />teeth) are similar, except that robusta has fewer <br />gill rakers and elegans more (Table 2). In small <br />tributaries, such as the Virgin and possibly the <br />San Juan, a single species with intermediate morph- <br />ology seems to be selected for; in the larger <br />tributary systems of the Gila River, a bewilder- <br />ing mosaic of generalized forms occur (Rinne, 1976), <br />where in former times robusta and elegans lived <br />in the main channel. <br /> <br />The close apparent control of habitat size and <br />diversity over morphology and species diversity <br />suggests that if the habitat could be experimen- <br />tally changed, the populations should show pre- <br />dictable responses. Rather unfortunately, the <br />destruction of main-river habitat by Glen Canyon <br />Dam has created such an experiment. Early indi- <br />cations are that the two more specialized species <br />will not exist in the Lake Powell environment. <br />The three species seem to be breaking down locally <br />by hybridization. We predict that a single- <br />species of mixed origins and generalized charac- <br />teristics will appear. Given long enough, some <br />re-oriented diversification would develop, but <br />the reservoir is apparently silting in too rapidly <br /> <br />for the necessary stability to persist. <br /> <br />In a period of controversy following the poi- <br />soning of the upper Green River in 1962 (Miller <br />1963b), erroneous statements about the classifi- <br />cation, biology, and distribution of Colorado <br />River chubs appeared which have been destructive <br />to the understanding and management of these <br />fishes, and require correction. For example, <br />Stroud (1963:7) irresponsibly claimed that the <br />construction of Flaming Gorge Reservoir resulted <br />in making available "large numbers" of humpback <br />chubs, all of which were supposedly males. The <br />agency (U.S. Bureau of Sport Fisheries and Wild- <br />life) that "confirmed the presence and ready <br />availability of numerous humpback chubs in [the <br />Green River below Flaming Gorge Dam] . . ." <br />subsequently reported (Vanicek 1967; Vanicek et <br />al. 1970; Holden and Stalnaker 1975) the absence <br />of Gila cypha in the area indicated and great <br />rarity of this species below the mouth of the <br />Yampa River; only three specimens were taken in <br />1963, and none during 1964-1966, in the Green <br />River in Colorado and Utah (Kramer 1967:Table 2; <br />Vanicek et al. 1970:Table 4). <br /> <br />Vanicek and Kramer (1969:194-195) stated, <br />. . . c~iteria are not available for distinguish- <br />ing between young fish of the two morphological <br />variants" (Le., between what were then called <br />Gila r. robusta and G. r. elegans). "Consequent- <br />ly, specimens shorter than 200 rom total length <br />were combined in the present study under the <br />general taxon, Colorado chub.- Collections of <br />Gila robusta and G. elegans at The University of <br />Michigan, containing young as small as 22 to 40 rom <br />S.L., were identified as early as 1926; by 1968, <br />humpback chubs as small as 43 rom S.L. had been <br />determined. By the criteria reported here, we <br />have identified Gila robusta robusta (UMMZ 162818) <br />to 20 rom, Gila cypha (UMMZ 182415) to 54 rom, and <br />Gila elegans (UMMZ 162846) to 22 rom in total <br />length. Juveniles of three species are shown in <br />Figure 10. Holden and Stalnaker (1975) (citing <br />Minckley and Deacon, 1968, as authority) reported <br />Colorado squawfish (Ptychocheilus lucius Girard) <br />in the Grand Canyon, but the two specimens actu- <br />ally came from between Glen Canyon Dam and Lees <br />Ferry,. well above Gran d Canyon. The only valid <br />record (based on preserved material) known to us <br />of this species from the Colorado River in Grand <br />Canyon is represented by an adult (about 320 rom <br />S.L.) caught in 1975 by an unknown fisherman at <br />the mouth of Havasu Creek (ASU7087). However, <br />squawfish formerly moved up the Little Colorado <br />River in Grand Canyon to the base of Grand Falls <br />(Miller, 1963a:l, ftn.l). <br /> <br />622 <br /> <br />In our distribution map of the Gila robusta <br />complex (Figure 1), the record for G. elegans in <br />the Little Colorado River (at base of Grand Falls) <br />is based in part on the statement referred to <br />above,' in part on our conclusion as to the true <br />type locality for Gila elegans (and Gila robusta), <br />and in part on the absence of the bony tail from <br />the Little Colorado River above Grand Falls. The <br />holytype (USNM 251), as well as the three syntypes <br />of Gila robusta (USNM 246), were said to have <br />come from the "Zuni River, New Mexico" (Baird and <br />Girard, 1853, 1854; Girard, 1858:286-287). How- <br />ever, at the time of collection during the summer <br />(rainy season) of 1852, Zuni River was described <br />". . . as a mere rivulet, and not entitled to the <br />name of river; in most parts of our country it <br />would not be dignified with that of creek" <br />(Sitgreaves, 1854:5). This is hardly the habitat <br />of Gila elegans and, moreover, that species is <br />unknown from the Little Colorado River basin (to <br />which Zuni River is tributary in floods) above <br />Grand Falls, an impassible barrier 56 meters high <br />(Dryer, 1965). Furthermore, careful examination <br />of the channel of Zuni River in New Mexico con- <br />vinced us that (at least in recent centuries) <br />