<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 />
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