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7
Native Fishes of the Grand Canyon
Region: An Obituary?
W. L. MINCKLEY, Arizona State University, Tempe, Arizona
The highly endemic fish fauna of the Colorado River downstream from
Glen Canyon Dam has been largely extirpated. Damming, diversion. and
other human regulation stabilized the system, which enhanced introduced
nonnative fishes and proved detrimental to native species. Historical ac-
counts are used to (1) reconstruct the ecology of fishes in a pristine Colo-
rado River, (2) document changes resulting in the decline of native species,
and (3) indict the introduction and establishment of alien fishes as the ulti-
mate factor in extirpation of the natives. Despite legislation and dedicated
efforts by scientists and fisheries managers in the 1970s and early 1980s,
trends toward extinction of native fishes were not reversed. Deemphasis of
environmental issues because of economic and political pressures for devel-
opment in the later 1980s makes survival of this unique fauna doubtful.
INTRODUCTION
The U.S. National Park Service was founded in 1916 with a mandate to
preserve resources Placed under its care "unimpaired for the enjoyment of
future generations." Shortly thereafter, in 1919, Grand Canyon National
Park was established to protect the natural attributes of spectacular gorges
cut by the Colorado River. These efforts failed for some of the native
aquatic biota. Construction and operation of dams and reservoirs outside the
park resulted in a chain reaction of environmental change that forced its
124
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NATIVE FISlIES...
125
native fishes to local extinction (Dolan et al., 1974; Johnson and Carothers,
1987).
This paper concerns the decline and disappearance of indigenous fishes
from the lower Colorado River, including the Grand Canyon region. To
document this, the fishes are first placed in the context of a pristine ecosys-
tem, reconstructed logically and with a modicum of speculation from his-
toric data. Demise of the fauna started just after 1900, as the river was
progressively harnessed for water supply, flood control, and power produc-
tion. The fish fauna collapsed from downstream to upstream, in the same
sequence as the river was regulated. Reasons for this ecological catastrophe
are discussed for individual species and for the fauna as a whole. Finally,
the grim prognosis for native fishes is tempered by a brief review of legal
statutes, scientific facts, and moral obligations, all of which exist to save
them. But before this happens, it is clear that society must develop an ethic
sufficient to realize this goal.
THE PRISTINE HABITAT
The Colorado River collects water, sediments, and dissolved solids from
more than 600,000 km2 of mountains, plateaus, and basins (Graf, 1985~.
From north to south, the river flows through the Rocky Mountain, Colorado
plateau, and basin and range physiographic provinces. High elevations pro-
vide spring and early summer runoff in the form of snowmelt, and most of
the water yield is from that source. Historically, low flow predominated in
summer through winter, with late summer spates in the south, where a
bimodal pattern of winter rains and late summer monsoons prevails.
After leaving the Rockies, the nver winds through desert, where signifi-
cant amounts of water are lost to evaporation. Ions are concentrated and
added by inflowing salt springs; as a result, total dissolved solids increase.
Along with this material are sediments from the headwaters, joined by even
more stripped from the sparsely vegetated deserts and plateaus by runoff of
infrequent, often violent rains.
The course of the river almost defies description. After the steep headwa-
ters, it alternatively meanders through broad, aggraded valleys and then
slices through bedrock uplifts. Most valley reaches are now modified for
agriculture. The river is constrained and channelized from its original state
of flowing through braided channels, over bars of shifting sand and gravel,
and between alluvial banks. Runs and riffles were smooth and strong, and
oxbow lakes, backwaters behind sandbars, and other lentic habitats were
common. The canyons, if not used for a dam site or drowned by a reservoir,
remain much the same as they were. They vary in morphology, with steep
or sloping walls. Some are deeper than 1,500 m, others are less deep. A few
are relatively straight, while most are sinuous. Rapids are common, with
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126
COLORADO RIVER ECOLOGY AND DAM MANAGEMENT
high waves, whirlpools, and other turbulence creating "whitewater" (in the
jargon of the river), which reflects underwater obstructions such as bedrock
dikes, stony debris from side canyons, or rockfalls from the canyon walls
(Leopold, 1969; Dolan et al., 1978~. Between rapids are "pools," where
deep, strong currents, or flatwaters, flow through unobstructed channels.
Most accounts of the river stress its variability in discharge and sediment
load. In a -40-year, pre-dam period of record, discharge at Yuma, Arizona,
varied over five orders of magnitude, from 0.5 to 7,100 m3 so (Dill, 1944~.
Variations in transported sediment were also substantial. On an annual ba-
sis, materials accumulated during low discharge, to again be moved by high
water in spring. Longer-term cycles also existed. From 45.4 to 455 million
metric tons years of silt was transported through Grand Canyon between
1922 and 1935 (Howard, 1947; Howard and Dolan, 1981), while records at
Yuma varied from 36.3 to 326 million metric tons in the same period. Thus,
a significant percentage was aggrading on the floodplain and along the
channel. Bedload must have varied similarly, but measurements then, as
now, were unreliable. Degradation of these deposits ocurred during episodic
realignment and cutting of valley deposits by major floods, when virtual
slurries of sand, gravel, and boulders must have ground inexorably through
canyons. Such processes are graphically described in pre-dam accounts of
building and slouching of banks and bars along the lower river (Sykes,
1937; Smith and Crampton, 1987; Beer, 1988~.
Although impressive to humans, the dangers posed for desert fishes by
upper limits of these variations are more imagined than real. The physical
force of flooding rarely harms either individuals or populations (Meffe,
1984; Meffe and Minckley, 1987; Minckley and Meffe, 1987~. Fishes ap-
parently move to clear the surface along shore even in canyons, thereby
avoiding both the force of currents and molar action of transported bedload,
as well as higher turbidities in deeper, turbulent channels. In wide places,
they simply swim onto floodplains and then swim back as water recedes.
Coarse, suspended sediments are similarly innocuous except, perhaps, through
abrasion. High concentration of clays may suffocate fishes by clogging their
branchial chambers (Wallen, 1951), but such has rarely been observed in
western streams (Minckley, 1973~.
A major seasonal problem may have been the food supply. Food may
have been limiting during periods of high turbidity and bedload transport.
As detailed in this chapter by Blinn and Cole, lower parts of large erosive
streams are notoriously depauperate of fish foods. Algae and other primary
producers are limited by turbidity and scour, and plankton is rare. Inverte-
brates need primary producers, detrital materials, or other animals to eat,
and most are excluded from unstable bottoms anyway.
Conditions are quite different at low water. Coarse particles settle quickly,
water clears to expose boulders and canyon walls to sunlight for consider-
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lIATIVE FISlIES...
127
able depths, and bars are stabilized and armored with gravel as declining
discharges sweep away the sand. Such places provide substrate for biologi-
cal colonization.
No studies are available for large unmodified rivers of the lower Colo-
rado basin, but primary producers and stream invertebrates in small desert
streams are remarkably resilient (Meffe and Minckley, 1987; Grimm and
Fisher, 1989~. Diatoms reappear a day after flash flooding, and complex
algal communities reestablish in weeks (Fisher et al., 1982; Fisher, 1986~.
Insects with life cycles of days or weeks recolonize in a few days, and those
with longer generation times reappear over a month or two (Bruns and
Minckley, 1980; Gray, 1981; Gray and Fisher, 1981; Minckley, 1981~. Many
feed on detritus, finely ground from leaves, branches, and trunks of trees
being carried to the sea (Minckley and Rinne, 19851. Thus, considering the
predictably long periods of low discharge in the river, algae and inverte-
brates may have been in ample supply for much of the year. Added to this
were terrestrial invertebrates (e.g., Tyus and Minckley, 1988) and diverse
inputs from tributaries.
Despite these benefits of low discharge, drought is the single most dan-
gerous time for fishes. Many western fishes appear to require little more
than water to survive, but they do need water. In valley reaches, infiltration
into deep, coarse alluvial fill robs the channel of surface flow, and intermittence
or desiccation may result. High temperatures and oxygen depletion in the
remnant pools become critical and lethal. On the other hand, these massive
alleviated valleys, many of which are actually structural intermontane ba-
sins, also act as vast subterranean reservoirs, dammed by the bedrock through
which the canyons are cut.
These reservoirs were the key to fish survival. They leaked downslope
into canyon-bound reaches, where pools held water in scoured holes and
shaded undercuts along cliffs. Subterranean water percolating through coarse
alluvium tends to be cool, mixed, and rich in nutrients (Grimm et al., 1981~;
thus, canyons often provide salubrious, highly productive habitat if suffi-
cient sunlight is available. Bedrock reaches are also rich in springs, adding
security to the system. In the large and complex Colorado River basin,
mainstream fishes had many canyon refugia and after a few weeks, at most,
would have been saved from even the greatest regional drought by runoff
from somewhere. Clearly it worked over the millennia, since some of the
fishes have persisted since Miocene (Minckley, et al., 1986~.
UNIQUENESS OF THE FISH FAUNA
The special nature of the Colorado River fish fauna was recognized early.
Evermann and Rutter (1895, p. 475) listed 5 families, 18 genera, and 32
indigenous species, and wrote:
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128
COLORADO RIVER ECOLOGY AND DAM MANAGEMENT
Though the families and species . . . are very few, they are of unusual
interest to the student of geographic distribution . . . over 78 percent of
the species of fishes now known from the Colorado Basin are peculiar
to it . . . a larger percentage of species peculiar to a single river basin
than is found elsewhere In North America.
Subsequent work has not altered these conclusions. Miller (1959) re-
ported 31 freshwater species, while Minckley et al. (1986) documented 32.
Recognized endemism at the species level still hovers near 75%. and Carlson
and Muth (1989) compiled 44 taxa, 93% endemic when undescribed forms
and subspecies were included. Clearly, as a result of long isolation and
special conditions, the Colorado River basin supports the most distinctive
ichthyofauna in North America.
Larger fishes of the Colorado River mainstream share an intriguing, number
of features (Hubbs, 1941; Miller and Webb, 1986~. Their adult body sizes
are large, and some species live exceptionally long lives. (All fish measure-
ments reported here are for total length, from the extreme tip of the snout to
the distal end of the caudal fin.) Body shapes are more fusiform and stream-
lined than most, with streamlining carried to extremes to include small
depressed skulls, large predorsal humps or keels (or both), and elongated,
pencil-thin caudal peduncles. Their eyes are small, the fins are expansive
and often falcate (sickle shaped) with strong leading rays, and the skins are
thick and leathery, especially on the head, anterior body, and leading edges
of fins. Finally, the scales are tiny, deeply embedded in the skin, or some-
times essentially absent.
Most of these special features have been related to severity of habitat.
Body and fin shape, structure, and size are construed as hydrodynamic
adaptations for maintaining position and maneuvering in swift, turbulent
currents. Large bodies and fins may, however, be as much a necessity to
generate sufficient power to negotiate such places. Leathery skins with small,
embedded, or reduced scales may minimize friction, counteract sediment
abrasion, or provide a rigid external sheath to maximize muscle efficiency.
Reduced eyes may be another way to avoid abrasion. Finally, long life
seems adaptive to unpredictable environments, since the ability to repro-
duce spans decades rather than only a few years. On the other hand, large
size (correlated with long life) scarcely seems adaptive to seasonally low
discharge. Despite alternative possibilities, common trends in morphology
across a number of unrelated taxa speak for commonality in other than
phylogeny, and the river's harsh selective arena seems a reasonable choice.
HISTORIC REVIEW
Early Records and Surveys
Faunal remains in archaeological sites show that Colorado River fishes
were caught and eaten by Indians (Miller, 1955; Euler, 1978; Miller and
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NATIVE FISHES...
129
Smith, 1984~. Early canyon explorers ate them too, and valuable records
appear in the accounts of John Wesley Powell (1875), Stegner, (1954),
Robert Brewster Stanton's railway surveys (Smith and Crampton, 1987),
the adventures of Ellsworth and Emery Kolb (Kolb and Kolb, 1914; Kolb,
1989), and others.
The earliest scientific collectors sampled tributaries (Baird and Girard,
1853a-c; Girard, 1857, 1859; Cope and Yarrow, 1875; Kirsch, 1889), the
Colorado mainstream near Yuma (Abbott, 1860; Evermann and Rutter, 1895;
Gilbert and Scofield, 1898; Chamberlain, 1904; Snyder, 1915), and at up-
stream crossings on the Grand (= Colorado3 and Green Jordan, 1891; Evermann
and Rutter, 1895) rivers. Canyons remained inhospitable to humans until
rubberized boats made whitewater rafting safe and reliable. Even with mod-
ern equipment, sampling is difficult at best, and the faunas of canyon-bound
reaches of the river remain the least understood of all. Most species had
nonetheless been collected and described before 1900 (Minckley and Dou-
glas, 1991~. The early surge of inquiry declined over almost four decades of
complacency that followed Jordan and Evermann's (1896-1900) monograph
"The Fishes of North and Middle America."
EVIDENCES OF FAUNAL CHANGE
Carl L. Hubbs and Robert R. Miller reinitiated studies of Colorado River
fishes in the late 1930s. They began by producing revisionary works and
compilations of records (Hubbs and Miller, 1941; Miller, 1943), descrip-
tions of new taxa and life stages (Miller, 1946a; Winn and Miller, 1954;
Miller and Hubbs, 1960), and records of hybrids (Hubbs et al., 1942; Hubbs
and Miller, 1953), and they moved with time to authoritative biogeographic
accounts (Hubbs and Miller, 1948; Miller, 1959~. Scattered throughout these
works were notes on faunal declines, and Miller (1946b) published an early
plea for study of native fishes before further alterations of the large western
rivers were undertaken.
Dill's (1944) survey of the lower Colorado River was the first to provide
insight on both native and introduced fishes downstream from the new Boulder
(= Hoover) and Parker dams. He noted reductions in native species attrib-
uted to environmental changes associated with damming. Wallis (1951) considered
the fauna "in urgent need for further research, because so many forces are
fast exterminating it." Jonez et al. (1951) and Jonez and Sumner (1954) also
expressed concern for native fishes in the face of rapid environmental change.
Alien fishes attracted early attention (Dill, 1944; Beland, 1953b; Hubbs,
1954~. Reservoirs changed the river in ways that enhanced lentic-adapted,
nonnative species aonez et al., 1951; Beland, 1953a; Kimsey, 1958; Nicola,
1979), and reservoir spor~sheries became important regional resources (Moffett,
1942, 1943; Wallis, 1951; Jonez and Sumner, 1954~. A remarkable array of
both native and nonnative species were used as bait (Miller, 1952), and bait
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130
COLORADO RIVER ECOLOGY AND DAM MANAGEMENT
and forage fishes escaped or were intentionally stocked to join and feed
expanding game fish populations (Hubbs, 1954; Kimsey et al., 1957; LaRivers,
1962; U.S. Fish and Wildlife Service, 1980, 1981~.
By the 1960s, although essentially invisible to the untrained eye, indig-
enous fishes of the lower Colorado River had been largely replaced by exotic
species. Miller (1961) wrote of dramatic changes in abundance and distribu-
tion and of local extirpation of native forms. At this same time, Glen Canyon,
Flaming Gorge, Navajo, and other major dams authorized by the Colorado
River Storage Project Act of 1956 were also nearing completion. These were
far from invisible. The magnitude of change was finally realized, and alarms
began to sound in an emerging conservation community. The Colorado River
had indeed been tamed, its wildness was lost except in the most isolated and
inaccessible reaches, and the public was stirred to react. Resistance against
other dams was led by the Sierra Club, and conservationists voiced their
opposition to further modifications (Fradkin, 1981~.
In addition to this controversy, almost 700 km of the Green River system
above the new Flaming Gorge Dam was poisoned in 1962 to clear the way
for an introduced trout fishery (Holder, 1991~. Some of the targets were
native fishes. The operation went awry, and fishes were killed far down-
stream through Dinosaur National Monument (Miller, 1963a, 1964~. This
event, possibly more than anything else, solidified the resolve of those
protective of native fishes.
In addition to making available funds for reservoir planning and con-
struction, the storage act provided for evaluation of-the archaeological sig-
nificance of areas to be inundated. Preimpoundment surveys conducted along
with archaeological salvage operations included biological collections for
use in interpreting paleo-Indian ecology. Studies were made in the Flaming
Gorge Reservoir basin (Dibble and Stout, 1960; Woodbury, 1963), Navajo
basin (Pendergast and Stout, 1961), and the future basin of Lake Powell
(Woodbury, 1959; McDonald and Dotson, 1960~. Marble and Grand can-
yons were ignored since they were not to be directly affected. By the time
Miller (1968) and Suttkus and colleagues (Suttkus et al., 1976; Suttkus and
Clemmer, 1979) began sampling in 1968 and 1970, respectively, the down-
stream impacts of Glen Canyon Dam were already evident.
In 1963, partially in response to public outcry over the poisoning, agen-
cies such as the National Park Service, Fish and Wildlife Service, and state
conservation departments began studies of Green River fishes (Vanicek,
1967; Vanicek and Kramer, 1969; Vanicek et al., 1970~. Such research
expanded elsewhere as new legislation came to bear in the late 1960s (Holder,
1973; Holden and Stalnaker, 1975~. Disappearing species became rallying
points for the Endangered Species Protection Act of 1966, which evolved to
the Endangered Species Act of 1973. Recognition that habitat was being
lost at an unacceptable rate stimulated the National Environmental Protec-
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NATIVE FISHES...
131
tion Act of 1969, which mandated assessment and disclosure of impacts of
federal projects. After 1966, lists of species in jeopardy served to focus
research and management efforts and agencies such as the Bureau of Recla-
mation became involved to satisfy legal requirements for project construc-
tion and operations.
Ongoing research has since documented the status and ecology of fishes
targeted by official listings. Some resulted in major reports (Minckley, 1979;
Carothers and Minckley, 1981; Miller et al., 1982b-c; Maddux et al., 1987;
Tyus et al., 1987; Ohmart et al., 1988), only excerpts of which have appeared
in the open literature. Smaller, more specific projects dealt with species' biol-
ogy or habitat; many of these are cited in recovery plans (U.S. Fish and
Wildlife Service, 1989b-d) or status reports for rare species (Seethaler, 1978;
McAda and Wydoski, 1980; Minckley, 1983; Kaeding and Osmundson, 1988;
Minckley et al., 1991; Tyus, 1991~. Reviews varied from annotated bibliogra-
phies (Wydoski et al., 1980), through edited symposia (Spofford et al., 1980;
Miller et al., 1982a; Adams and Lamarra, 1983; Minckley and Deacon, 1991),
to contributions dealing with the fauna (Deacon, 1968, 1979; Joseph et al.,
1977; Holden, 1979; Behnke and Benson, 1983; Stanford and Ward, 1986c)
and its ecosystem (Deacon and Minckley, 1974; Ono et al., 1983; Williams et
al., 1985; Stanford and Ward, 1986a,b; Carlson and Muth, 1989~. In short, a
wealth of information now exists on Colorado River fishes.
NATIVE FISHES
Six of eight fishes native to Grand Canyon National Park (Table 7-1) are
endemic. Speckled dace and roundtail chub are known from adjacent rivers,
and the latter, known only from a few specimens (C. O. Minckley, 1980), is
excluded from further consideration. Four of the remainder, humpback chub,
bonytail, Colorado squawfish, and razorback sucker, are listed or proposed
as endangered by the Department of the Interior (U.S. Fish and Wildlife
Service 1989a, 1990~. Of these, only the humpback chub (Figure 7-1) per-
sists as a reproducing population. The other three are extirpated or exceed-
ingly rare. Speckled dace, flannelmouth sucker, and bluehead sucker remain
relatively common (Minckley, 1985~.
Most early accounts of fishes in the Colorado River were for large spe-
cies sought for food, e.g., "Colorado River salmon" (squawfish) was on the
menu of Christmas dinner for the Stanton Party in 1889 (Measeles, 1981;
Smith and Crampton, 1987~. Most records were more informal. When Kolb
and Kolb (1914, p. 123) heard splashing near their camp at the Little Colo-
rado River in 1911. they wrote:
. . . the fins and tails of numerous fish could be seen above the water.
The striking of their tails had caused the noise we had heard. The 'bony
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132
COLORADO RIVER ECOLOGY AND DAM MANAGEMENT
TABLE 7-1 Common and scieniific names of naiive and introduced
fishes recorded from Grand Canyon National Park, Arizona. Those taxa
marked with an asterisk (*) are listed or proposed for listing as
endangered by the U.S. Deparunent of the Interior.
CLUPEIDAE, shads introduced
Threadfin shad
SALMONIDAE, salmons and trout all introduced
Apaehe trout
Cutthroat trout
Silver Salmon
Rainbow trout
Brown trout
Brook trout
CYPRINIDAE, minnows
Native species
*Humpback ehub
*Bonytail
Roundtail ehub
*Colorado squawfish
Speekled daee
Introdueed species
Common earp
Red shiner
Golden shiner
Fathead minnow
Redside shiner
CATOSTOMIDAE, suckers all native
Flannelrnouth sucker
Bluchead sucker
*Razorback sucker
ICTALURIDAE, bullhead catfishes all introduced
Blaek bullhead
Channel catfish
FUNDULIDAE, killifishes introduced
Plains killifish
POECILIIDAE, livebearers- introdueed
Mosquitofish
CENTRARCHIDAE, sunfishes—all introduced
Green sunfish
Bluegill
Largemouth bass
PERCICHTHYIDAE, temperate basses introduced
Striped bass
Dorosoma petenense GHnther
Oncorhynchus apache Miller
O. clarki Riehardson
O. kisutck Walbaum
O. mykiss Walbaum
Salmo trutta Linnaeus
Salvelinus fontinalis Mitehill
Gila cypha Miller
G. elegans Baird and Girard
G.r. robusta Baird and Girard
Ptychocheilus lucius Girard
Rhinichthys osculus Girard
Cyprinus carpio Linnaeus
Cyprinella lutrensis Baird and Girard
Notemigonus crysoleucus Mitchill
Pimephales promelas Rafinesque
Richardsonius balteatus Richardson
Catostomus latipinnis Baird and Girard
Pantosteus discobolus Cope
Xyrauchen texanus Abbott
Ameiurus melas Rafinesque
fctalurus punctatus Rafinesque
Fundulus zebrinus Jordan and Gilbert
Gambusia affinis Baird and Girard
Lepomis cyanellus Rafinesque
L. machrochirus Rafinesque
Micropterus salmoides Lacepede
Morone saxatilis Walbaum
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NATIVE FISHES...
a:
133
FIGURE 7-1 Male humpback chub, cat 38 cm long, captured from the Little
Colorado River, Arizona; photograph by J. N. Rinne at Willow Beach National Fish
Hatchery.
tail' were spawning .... The Colorado is full of them; so are many
other muddy streams of the Southwest. They seldom exceed 16 inches
in length, and are silvery white in color. With a small flat head some-
what like a pike, the body swells behind it to a large hump.
Suttkus and Clemmer (1977) believed the fish to be humpback chubs
rather than bonytail, which seems likely. True bonytail (Figure 7-2) were
better described by Dellenbaugh (1984, p. 15) from the Green River:
. . . a fish about ten to sixteen inches long, slim with fine scales and
large fins. Their heads came down with a sudden curve to the mouth,
and their bodies tapered off to a very small circumference just before
the tail spread out.
Another quotation from Dellenbaugh (1984, p. 98) introduces the Colo-
rado squawfish (Figure 7-3 and 7-4~. The incident occulted near the present
town of Green River, Utah, in 1871; Minckley (1973, p. 124) erred in
attributing it to Grand Canyon in 1872:
He [a member of the second Powell Expedition] thought his precious
hook was caught on a snag. Pulling gently in order not to break his line
the snag lifted with it and presently he was astounded to see, not the
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134 COLORADO RIVER ECOLOGY AND DAM MANAGEMENT
FIGURE 7-2 Female bonytail, cat 50 cm long, trammel-netted from Lake Mohave,
Arizona-Nevada; photograph by W. L. Minckley.
branch of a tree or a waterlogged stick, but the head of an enormous
fish appear above the surface .... Casting again another of the same
kind came forth and then a third. The longest . . . was at least thirty or
thirty-six inches with a circumference of fifteen inches. The others were
considerably shorter but nevertheless very large fish.
Colorado squawfish were familiar to all of the early canyon explorers.
Stanton recorded "large" or"huge" fish caught in the Green and Colorado
rivers (Smith and Crampton, 1987~. Dellenbaugh (1984, facing p. 102) pub-
lished a photograph of two large ones taken by the Stanton party in 1889. In
southern Wyoming in 1911, Kolb (1989, p. 15)
. . . caught sight of fish gathered in a deep pool, under the foliage of a
cottonwood tree which had fallen into the river. Our most tempting bait
failed to interest them; so Emery, ever clever with hook and line, 'snagged'
a catfish . . . for salmon bait, a fourteen-pound specimen rewarding our
attempt .... They sometimes weigh twenty-five or thirty pounds, and
are common at twenty pounds; being stockily built fish, with large, flat
heads.
I found no references to other species in this kind of older literature,
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NATIVE FISHES...
167
Gray, L. J. 1981. Species composition and life histories of aquatic insects in a lowland Sonoran
Desert stream. American Midland Naturalist 106:229-242.
Gray, L. J., and S. G. Fisher. 1981. Postflood recolonization pathways of macroinvenebrates in
a lowland Sonoran Desert stream. American Midland Naturalist 106:149-157.
Grimm, N. B., and S. G. Fisher. 1989. Stability of periphyton and macroinvertebrates to
disturbance by flash floods in a desert stream. Journal of the North American Benthological
Society 8:293-307.
Grimm, N. B., S. G. Fisher, and W. L. Minckley. 1981. Nitrogen and phosphorus dynamics in
hot desert streams of southwestern U.S.A. Hydrobiologia 83:303-312.
Gustaveson, A. W., H. R. Maddux, and B. L. Bonebrake. 1990. Assessment of a forage fish
introduction into Lake Powell. Utah Department of Natural Resources, Division of Wildlife
Resources, Salt Lake. 51 p.
Hamman, R. L. 1981. Spawning and culture of Colorado squawfish in raceways. Progressive
Fish-Culturist 43: 173-177.
Hamman, R. L. 1982a. Culture of endangered Colorado River fishes. Section II. Induced
spawning and culture of the humback chub. P. 158-167, in W. H. Miller, J. J. Valentine, D.
L. Archer, H. M. Tyus, R. A. Valdez, and L. R. Kacding, eds., Colorado River Fisheries
Project, Part 3. Contracted Studies. Final Report, U.S. Bureau of Reclamation Contract 9-
07-40-L-1016, and U.S. Bureau of Land Management Memorandum of Understanding CO-
910-MU9-933. U.S. Fish and Wildlife Service, Salt Lake City, Utah.
Hamman, R. L. 1982b. Spawning and culture of humpback chub. Progressive Fish-Culturist
44:213-216.
Hendricks, L. J. 1961. The striped mullet, Mugil cephalus Linnaeus. P. 93-94, in B. W. Walker,
ea., The ecology of the Salton Sea, California, in relation to the sportf~shery. California
Department of Fish and Game Fish Bulletin 113.
Holden, P. B. 1968. Systematic studies of the genus Gila (Cyprinidae) of the Colorado River
Basin. Unpublished Master's Thesis, Utah State University, Logan. 68 p.
Holden, P. B. 1973. Distribution, abundance, and life history of the fishes of the upper
Colorado River Basin. Unpublished Doctoral Dissertation, Utah State University, Logan. 59 p.
Holden, P. B. 1979. Ecology of riverine fishes in regulated stream systems, with emphasis on
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