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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, sunfishesall 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 the Colorado River. P. 57-74, in J. V. Ward and J. A. Stanford, eds., The Ecology of Regulated Streams. Plenum Press, New York. Holden, P. B. 1991. Ghosts of the Green River: Impacts of Green River poisoning on manage- ment of native Sshes. In press, in W. L. Minckley and J. E. Deacon, eds., Battle Against Extinction: Native Fish Management in the American West. University of Arizona Press, Tucson. Holden, P. B., and C. B. Stalnaker. 1970. Systematic studies of the cyprinid genus Gila in the upper Colorado River basin. Copeia 1970:409-429. Holden, P. B., and C. B. Stalnaker. 1975. Distribution and abundance of mainstream fishes of the middle and upper Colorado River basin, 1967-1973. Transactions of the American Fisheries Society 104:217-231. Holden, P. B., and E. J. Wick. 1982. Life history and prospects for recovery of Colorado squawfish. P. 98-108, in W. H. Miller, H. M. Tyus, and C. A. Carlson, eds., Fishes of the Upper Colorado River System: Present and Future. Westem Division, American Fisheries Society, Bethesda, Md. Howard, A. S., and R. Dolan. 1981. Geomorphology of the Colorado River in the Grand Canyon. Journal of Geology 89:269-298. Howard, C. S. 1947. Suspended sediment in the Colorado River. U.S. Geological Survey Water-Supply Paper 998:1-165. Hubbs, C. L. 1941. The relation of hydrological conditions to speciation in fishes. P. 182-195, in A Symposium on Hydrobiology. University of Wisconsin Press, Madison.

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168 COLORADO RIVER ECOLOGY ACID DAM MANAGEMENT Hubbs, C. L. 1954. Establishment of a forage fish, the red shiner (Notropis lutrensis), in the lower Colorado River system. California Fish and Game 40:287-294. Hubbs, C. L., L. C. Hubbs, and R. C. Johnson. 1942. Hybridization in nature between species of catostomid fishes. Contributions of the University of Michigan Laboratory of Vertebrate Biology 22:1-76. Hubbs, C. L., and R.R. Miller. 1941. Studies of the fishes of the order Cyprinodontes. XVII. Genera and species of the Colorado River system. Occasional Papers of the University of Michigan Museum of Zoology 433:1-9. Hubbs, C. L., and R. R. Miller. 1948. The zoological evidence: Correlation between fish distribution and hydrographic history in the desert basins of western United States. Bulletin of the University of Utah 30:17-166. Hubbs, C. L., and R. R. Miller. 1953. Hybridization in nature between the fish genera Catostomus and Xyrauchen. Papers of the University of Michigan Museum of Zoology 38:207-233. Janisch, J. L. 1985. Evaluation of Lees Ferry fishery and future management. Arizona Game and Fish Department Publications 85-3:1-23. Johnson, J. E., and B. L. Jensen. 1991. Hatcheries for endangered freshwater fishes. In press, in W. L. Minckley and J. E. Deacon, eds., Battle Against Extinction: Native Fish Manage- ment in the American West. University of Arizona Press, Tucson. Johnson, J. E., R. R. Kramer, E. Larsen, and L. Bonebrake. 1987. Flaming Gorge fisheries investigations: Trout growth, survival, and microhabitat selection in the Green River, Utah. 1978-82. Utah Division Wildlife Publication 87-13:1-185. Johnson, R. R., and S. W. Carothers. 1987. External threats: The dilemma of resource manage- ment on the Colorado River in Grand Canyon National Park, USA. Environmental Manage- ment 11:99-107. Jonez, A., R. D. B eland, G. Duncan, and R. A. Wagner. 1951. Fisheries report of the lower Colorado River. Fisheries Task Group-Colorado River. Great Basin Field Commission. Arizona Game and Fish Commission., Phoenix. 89 p. Jonez, A., and R. C. Sumner. 1954. Lakes Mead and Mohave investigations, a comparative study of an established reservoir as related to a newly created impoundment. Final Report, Federal Aid Wildlife Restoration (Dingell-Johnson) Project F-1-R, Nevada Game and Fish Commission, Carson City. 186 p. Jordan, D. S. 1891. Repon of explorations in Utah and Colorado during the summer of 1889, with an account of the fishes found in each of the river basins examined. Bulletin of the U.S. Fish Commissioner 9:1-40. Jordan, D. S., and B. W. Evermann. 1896-1900. lathe fishes of North and Middle America. Bulletin of the U.S. National Museum 47(4 pans):1-3313. Joseph, T. W., J. A. Sinning, R. J. Behnke, and P. B. Holden. 1977. An evaluation of the status, life history, and habitat requirements of endangered and threatened fishes of the upper Colorado River system. U.S. Fish Wildlife Service FWS/OBS-77062:1-169. Kaeding, L. R., B. D. Burdick, P. A. Schrader, and W. R. Noonan. 1986. Recent capture of a bonytail (Gila elegans) and observations on this nearly extinct cyprinid from the Colorado River. Copeia 1986:1021-1023. Kaeding, L. R., and D. B. Osmundson. 1988. Interaction of slow growth and increased early- life mortality: A hypothesis on the decline of Colorado squawfish in the upstream regions of its historic range. Environmental Biology of Fishes 22:287-298. Kaeding, L. R., and M. A. Zimmerman. 1982. Life history and ecology of the humpback chub in the Little Colorado and Colorado rivers of the Grand Canyon, Arizona. P. 281-320, in W. H. Miller, J. J. Valentine, D. L. Archer, H. M. Tyus, R. A. Valdez, and L. R. Kaeding, eds., Colorado River Fishery Project, Part II. Field Investigations. Final Report, U.S. Bureau of Reclamation Contract 9-07-40-L-1016, and U.S. Bureau of Land Management

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NATIVE FISHES... 169 Memorandum of Understanding CO-910-MU9-933. U.S. Fish and Wildlife Service, Salt Lake City, Utah. Kaeding, L. R., and M. A. Zimmerman. 1983. Life history and ecology of the humpback chub in the Little Colorado and Colorado rivers of Grand Canyon. Transactions of the American Fisheries Society 112:577-594. Karp, C.A., and H. M. Tyus. 1990. Behavioral interactions between young Colorado squawfish and six fish species. Cope u: 1990:25-34. Kimsey, J. B. 1958. Fisheries problems in impounded waters of Califomia and the lower Colorado River. Transactions of the American Fisheries Society 87:319-332. Kimsey, J. B., R. H. Hagy, and G. W. McCammon. 1957. Progress report on the Mississippi threadfin shad, Dorosorna petenense atchafaylae (sic.) in the Colorado River for 1956. California Department of Fish and Game Inland Fisheries Administrative Report 57-23:1- 48. Kirsch, P. H. 1889. Notes on a collection of fishes obtained in the Gila River at Fort Thomas, Arizona. Proceedings of the U.S. National Museum 11:555-558. Kolb, E. L. 1989 (originally published 1914). Through the Grand Canyon from Wyonung to Mexico. University of Arizona Press, Tucson. 344 p. Kolb, E. L., and E. Kolb. 1914. Experience in the Grand Canyon. National Geographic Maga- zine 26(2):99-184. Kondolf, G. M., S. S. Cook, H. R. Maddux, and W. R. Persons. 1989. Spawning gravels of rainbow trout in Glen and Grand Canyons, Arizona. Journal of the Arizona-Nevada Acad- emy of Science 23:19-28. Kubly, D. M. 1990. The endangered humpback chub (Gila cyp).a) in Arizona: A review of past studies and suggestions for future research. Repon, U.S. Bureau of Reclamation, Salt Lake City. Arizona Game and Fish Department, Phoenix. 138 p. (draft, February 1990). Kubly, D. M., and G. A. Cole. 1979. The chemistry of the Colorado River and its tributaries in Marble and Grand canyons. P. 565-572, in Proceedings of the First Annual Conference on Scientific Research in the National Parks. U.S. National Park Service Transactions and Proceedings Series 5. LaRivers, I. 1962. Fish and Fisheries of Nevada. Nevada State Printing Office, Carson City. 782 p. Leopold, L. B. 1969. The rapids and poolsGrand Canyon. U.S. Geological Survey Profes- sional Paper 669d:131-145. Maddux, H. R., and W. G. Kepner. 1988. Spawning of bluehead sucker in Kanab Creek, Arizona (Pisces: Catostomidae). The Southwestern Naturalist 33:364-365. Maddux, H. R., D. M. Kubly, J. C. DeVos, Jr., W. M. Persons, R. Stacdicke, and R. L. Wright. 1987. Effects of varied flow regimes on aquatic resource sof Glen and Grand canyons. Final Report, U.S. Bureau of Reclamation Contract 4-AG-40-01810. Arizona Game and Fish Department, Phoenix. 291 p. Marsh, P. C. 1985. Effects of incubation temperature on survival of embryos of native Colo- rado River fishes. The Southwestern Naturalist 30:129-140. Marsh, P. C. 1987. Food of adult razorback sucker in Lake Mohave, Arizona-Nevada. Transac- tions of the American Fisheries Society 116:1 17-119. Marsh, P. C., F. J. Abarca, M. E. Douglas, and W. L. Minckley. 1989. Spikedace (Meda fulgida) and loach minnow (Tiaroga cobitis) relative to introduced red shiner (Cyprinella lutrensis). Final Report Arizona Game and Fish Department Contract. Arizona State Uni- versity, Tempe. 116 p. McAda, C. W. 1977. Aspects of the life history of three catostomids native to the upper Colorado River basin. Unpublished Master's Thesis, Utah State University, Logan. 73 p. McAda, C. W., and R. W. Wydoski. 1980. The razorback sucker, Xyrauchen texanus, in the

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170 COLORADO RIVER ECOLOGY AND DAM MANAGEMENT upper Colorado River basin, 1974-1976. U.S. Fish Wildlife Service Technical Papers 99:1 -15. McAda, C. W., and R. W. Wydoski. 1983. Maturity and fecundity of the bluehead sucker, Catostomus discobolus (Catostomidae), in the upper Colorado River basin, 1975-76. The Southwestern Naturalist 28:120-123. McAda, C. W., and R. W. Wydoski. 1985. Growth and reproduction of the flannelmouth sucker, Catostomus latipinnis, in the upper Colorado River basin, 1975-76. Great Basin Natural ist 45 :281-286. McCarthy, M. S., and W. L. Minckley. 1987. Age estimation for razorback sucker (Pisces: Catostomidae) from Lake Mohave, Arizona and Nevada. Journal of the Arizona-Nevada Academy of Science 21:87-97. McDonald, D. B., and P. A. Dotson. 1960. Fishery investigations of the Glen Canyon and Flaming Gorge impoundment areas. Utah State Department of Fish and Game Information Bulletin 60-3: 1-70. McKee, E. D. 1930. Briefs. Grand Canyon Nature Notes 4(4):37. Mcaseles, E. B. 1981. A Crossing on the Colorado, Lees Ferry. Pruett Publishing Co., Boulder, Colorado. 13 8 p. Meffe, G. K. 1983. Ecology of species replacement in the Sonoran topminnow (Poeciliopsis occidentalis) and the mosquitofish (Gambusia affinis) in Arizona. Unpublished Doctoral Dissertation, Arizona State University, Tempe. 143 p. Mcffe, G. K. 1984. Effects of abiotic disturbance on coexistence of predatory-prey fish spe- cies. Ecology 65:1525-1534. Mcffe, G. K. 1985. Predation and species replacement in Southwestem fishes: A case study. The Southwestern Naturalist 30:173-187. Meffe, G. K., and W. L. Minckley. 1987. Persistence and stability of fish and invertebrate assemblages in a repeatedly disturbed Sonoran Desert stream. American Midland Naturalist 1 17:177-191. Miller, R. R. 1943. The status of Cyrinodon macularius and Cyprinodon nevadensis, two desert fishes of western Nonh America. Occasional Papers of the University of Michigan Museum of Zoology 473:1-25. Miller, R. R. 1946a. Gila cyp ha, a remarkable new species of fish from the Colorado River in Grand Canyon, Arizona. Journal of the Washington Academy of Science 35:403-415. Miller, R. R. 1946b. The need for ichthyological surveys of the major rivers of western North America. Science 104:517-519. Miller, R. R. 1952. Bait fishes of the lower Colorado River, from Lalce Mead, Nevada, to Yuma, Arizona, with a key for their identification. California Fish and Game 38:7-42. Miller, R. R. 1955. Fish remains from archaeological sites in the lower Colorado River basin, Arizona. Papers of the Michigan Academy Science, Arts, and Letters 40:125-136. Miller, R. R. 1959. Origin and affinities of the freshwater fish fauna of western North America. Pages 187-222, in C.L. Hubbs, ea., Zoogeography. American Association for the Advance- ment of Science Publication 51(1958), Washington, D.C. Miller, R. R. 1961. Man and the changing fish fauna of the American Southwest. Papers of the Michigan Academy of Science, Arts, and Letters 46:365-404. Miller, R. R. 1963a. Is our native underwater life worth saving? Nationa Parks Magazine 37 :4-9. Miller, R. R. 1963b. Distribution, variation and ecology of Lepidomeda vittata, a rare cyprinid fish endemic to eastem Arizona. Copeia 1963:1-5. Miller, R. R. 1964. Fishes of Dinosaur. Naturalist 15:24-29. Miller, R. R. 1968. Unpublished field notes: 1968 Arizona collecting expedition. On file in the Fish Division, University of Michigan Museum of Zoology, Ann Arbor, Mich. Miller, R. R., and C. L. Hubbs. 1960. The spiny-rayed cyprinid fishes (Plagopterini) of the

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NATIVE FISIlES... 171 Colorado River system. Miscellaneous Publications of the University of Michigan Museum of Zoology 1 1 5:1 -39. Miller, R. R., and G. R. Smith. 1984. Fish remains from Stanton's Cave, Grand Canyon of the Colorado, Arizona, with notes on the taxonomy of Gila cypha. P. 61-65, in R. C. Euler, ea., The Archaeology, Geology, and Paleabiology of Stanton's Cave, Grand Canyon National Park, Arizona. Grand Canyon Natural History Association Monograph 6. Miller, R. R., and P. W. Webb. 1986. Tactics used by Rhinichthys and some other fresh water fishes in response to strong currents. Abstracts of the Sixty-fourth Annual Meeting, Amen- can Society of Ichthyologists and Herpetologists, Victoria, British Columbia. Miller, W. H., H. M. Tyus, and C. A. Carlson, eds. 1982a. Fishes of the Upper Colorado River System: Present and Future. Western Division, American Fisheries Society, Bethesda, Md. 131 p. Miller, W. H., J. J. Valentine, D. L. Archer, H. M. Tyus, R. A. Valdez, and L. R. Kacding, eds. 1982b. Colorado River Fisheries Project, Part 1. Summary Report. Final Report, U.S. Bu- reau of Reclamation Contract 9-070-L-1016, and U.S. Bureau of Land Management Memo- randum of Understanding CO-910-MU9-933. U.S. and Fish Wildlife Service, Salt Lake City. 42 p. Miller, W. H., J. J. Valentine, D. L. Archer, H. M. Tyus, R. A. Valdez, and L. R. Kaeding, eds. 1982c. Ibid. Part 2. Field investigations. Ibid. 365 p. Miller, W. H., J. J. Valentine, D. L. Archer, H. M. Tyus, R. A. Valdez, and L. R. Kacding, eds. 1982d. Ibid. Part 3. Contracted Studies. Ibid. 324 p. Minckley, C. O. 1978. A report on aquatic investigations conducted during 1976, 1977 on Bright Angel, Phantom and Pipe creeks, Grand Canyon National Park, Coconino County, Arizona. Annual Investigation Report, Grand Canyon National Park, Grand Canyon, Ariz. Northem Arizona University, Flagstaff. 112 p. Minckley, C. O. 1980. River resource monitoring project, Gila spp. studies, Grand Canyon National Park: Part 2. Repon National Park Service Contract CX-82-1070012. Museum of Northem Anzona, Flagstaff. 23 p. Minckley, C. O. 1988. Final report on research conducted on the Little Colorado River popula- tion of the humpback chub, during May 1987, 1988. Repon to Non-game Branch, Arizona Game and Fish Department, Phoenix. Nonhem Arizona University, Flagstaff. 131 p. Minckley, C. O. 1989. Final report on research conducted on the Little Colorado River popula- tion of the humpback chub during May 1989. Ibid. 36 p. Minckley, C. O., and D. W. Blinn, 1976. Summer distribution and reproductive status of fish of the Colorado River in Grand Canyon National Park and vicinity, 1975. Final Report National Park Service Contract CX-82-1060008. Northern Arizona University, Flagstaff. 17 p. Minckley, C. O., and S. W. Carothers. 1980. Recent collections of the Colorado squawfish and razorback sucker from the San Juan and Colorado rivers in New Mexico and Arizona. The Southwestern Naturalist 24:686-687. Minckley, C. O., S. W. Carothers, J. W. Jordan, and H. D. Usher. 1981. Observations on the humpback chub, Gila cyp ha, within the Colorado and Little Colorado rivers, Grand Canyon National Park, Arizona. P. 176-183, in Proceedings of the Second Annual Conference on Scientific Research in the National Parks. U.S. National Park Service Transactions and Proceedings Series 8. Minckley, W. L. 1965. Native fishes as natural resources. P. 48-60, in J. L. Gardner, ea., Native Plants and Animals as Resources in Arid Lands of the southwestern United States. Southwest-Rocky Mountain Division, American Association for the Advancement of Sci- ence, Contribution 8, Committee on Desert and Arid Lands Research. Minckley, W. L. 1973. Fishes of Arizona. Arizona Game and Fish Department, Phoenix. 293 p. Minckley, W. L. 1979. Aquatic habitats and fishes of the lower Colorado River, southwestern

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172 COLORADO RIVER ECOLOGY AND DAM MANAGEMENT United States. Final Report U.S. Bureau of Reclamation Contract 14-06-300-2529. Lower Colorado River Region, Boulder City, Nevada. Anzona State University, Tempe. 478 p. Minckley, W. L. 1980. Morphological variation in catostomid fishes of the Grand Canyon Region, middle Colorado River basin. Final Report U.S. National Park Service Contract, Grand Canyon National Park, Grand Canyon, Arizona. Arizona State University, Tempe. 27 p. Minckley, W. L. 1981. Ecological studies of Aravaipa Creek and environs, central Arizona, relative to past, present, and future uses. Final Report, U.S. Bureau of Land Management Contract YA-512 OCR for page 124
NATIVE FISHES... 173 Montgomery, W. L, W. Liebfried, and K. Gooby. 1986. Feeding by rainbow trout on Cladophora glomerata at Lee's Ferry, Colorado River, Arizona; the roles of Cladophora and epiphytic diatoms in trout nutrition. Final Report, U.S. Bureau of Reclamation Contract, Lower Colo- rado River Region, Boulder City, Nev. Northem Arizona University, Flagstaff, et al. 20 p. Moyle, P. B. 1976. Inland Fishes of California. University of California Press, Berkeley. 405 p. Mueller, G. 1989. Obsecvations of spawning razorback sucker (Xyrauchen texanus) utilizing river habitat in the lower Colorado River, Arizona-Nevada. The Southwestern Naturalist 34:147-149. National Research Council. 1987. River and Dam Management: A Review of the Bureau of Reclamation's Glen Canyon Environmental Studies. National Academy Press, Washington, D.C. 203 p. Nicola, S. J. 1979. Fisheries problems associated with the development of the lower Colorado River. California Department of Fish and Game, Inland Fisheries Endangered Species Program Special Publication 79-4:1-18. Ohmart, R. D., B. W. Anderson, and W. C. Hunter. 1988. The ecology of the lower Colorado River from Davis Dam to the Mexico-United States International Boundary: A community profile. U.S. Fish and Wildlife Service Biological Report 85(7.19):1-243, 2 appendices. Ono, R. D., J. D. Williams, and A. Wagner. 1983. Vanishing Fishes of North America. Stone Wall Press, Washington, D.C. 257 p. Pendergast, D. M., and C. C. Stout, editors. 1961. Ecological studies of the flora and fauna of Navajo Reservoir basin, Colorado and New Mexico, University of Utah Anthropological Papers 55:1-122. Persons, W. B., K. McCormack, and T. McCall. 1985. Fishery investigations of the Colorado River from Glen Canyon Dam to the confluence of the Pana River: Assessment of the impact of fluctuating flows on the Lee's Ferry fishery. Arizona Garne and Fish Department Publication 86-6:1-93. Pelts, G. E. 1984. Impounded Rivers Perspectives for Ecological Management. John Wiley and Sons, New York. 326 p. Pillsbury, A. F. 1981. The salinity of rivers. Scientific American 245:55-65. Pirnental, R., R. V. Bulkley, and H. M. Tyus. 1985. Choking of Colorado squawfish, Ptychocheilus lucius (Cyprinidae), on channel catfish, Ictalurus punctatus (Ictaluridae), as a cause of mortality. The Southwestern Naturalist 30:154-158. Platania, S. P., K. R. Bestgen, M. A. Morretti, D. L. Propst, and J. E. Brooks. In press. Status of Colorado squswfish and razorback sucker in the San Juan River, Colorado, New Mexico, and Utah, The Southwestern Naturalist 36. Powell, J. W. 1875, Exploration of the Colorado River of the West and Its Tributaries, Ex- plored in 1869, 1870, 1871, and 1872. U.S. Government Printing Office, Washington, D.C. Reisner, M. 1986. Cadillac Desert: The American West and Its Disappearing Water. Viking Press, New York. 582 p. Richards, M. T., and D. B. Wood. 1985. The economic value of sportfishing at Lees Ferry, Arizona. U.S. Department of Agriculture. Forest Service General Technical Report RM- 120:219-222. Rinne, J. N., J. E. Johnson, B. L. Jensen, A. W. Ruger, and R. Sorenson. 1986. The role of hatcheries in the management and recovery of threatened and endangered fishes. Pages 271-285, in R. H. Stroud, ea., Fish Culture in Fisheries Managment. American Fisheries Society, Bethesda, Md. Rinne, J. N., and P. R. Turner. 1991. Reclamation and alteration as management techniques, and a review of methodology in stream renovation. In press, in W.L. Minckley and J. E. Deacon, eds., Battle Against Extinction: Native Fish Management in the American West. University of Arizona Press, Tucson.

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174 COLORADO RIVER ECOLOGY AND DAM MANAGEMENT Scoppettone, G. G. 1988. Growth and longevity of the cui-ui (Chasmistes cujus) and suspected longevity in other catostomids and cyprinids. Transactions of the American Fisheries Soci- ety 117:301-307. Seethaler, K. H. 1978. Life history and ecology of the Colorado squawfish (Ptychocheilus lucius) in the Upper Colorado River Basin. Unpublished Master's Thesis, Utah State Uni- versity, Logan. 156 p. Siebert, D. J. 1980. Movements of fishes in Aravaipa Creek, Arizona. Unpublished NIaster's Thesis, Arizona State University, Tempe. 56 p. Sigler, W. F., and R. R. Miller. 1963. Fishes of Utah. Utah Department of Fish and Game, Salt Lake City. 203 p. Sigler, W. F., and J. W. Sigler. 1987. Fishes of the Great Basin: A Natural History. University of Nevada Press, Reno. 425 p. Smith, D. L., and C. G. Crampton, eds. 1987. The Colorado River Survey: Robert B. Stanton and the Denver, Colorado Canyon & Pacific Railroad. Howe Brothers Publishers, Salt Lake City. 305 p. Smith, G. R. 1959. Annotated checklist of fishes of Glen Canyon. P. 195-199, in A. M. Woodbury, ea., Ecological Studies of the Flora and Fauna in Glen Canyon. University of Utah Anthropological Papers 40. Smith, G. R. 1966. Distribution and evolution of the North American catostomid fishes of the subgenus Pantosteus, genus Catostomus. Miscellaneous Publications of the University of Michigan Museum of Zoology 29:1-132. Smith, G. R. 1981. Effects of habitat size on species richness and adult body size of desert fishes. P. 125-172, in R. J. Naiman and D. L. Soltz, eds., Fishes in North American Deserts. John Wiley and Sons, New York. Smith, G. R., G. G. Musser, and D. B. McDonald. 1959. Appendi~c A. Aquatic survey tabula- tion. P. 177-194, in A. hI. Woodbury, ea., Ecological studies of the flora and fauna in Glen Canyon. University of Utah Anthropological Papers 40. Smith, R., R. R. Miller, and W. D. Sable. 1979. Species relationships among fishes of the genus Gila of the upper Colorado River Basin. P. 613-623, in Proceedings of the First Annual Conference on Scientific Research in the National Parks. U.S. National Park Ser- vice Transactions and Proceedings Series 5. Smith, M. L 1981. Late Cenozoic fishes ~n the warm deserts of Nonh America: A reinterpretation of desen adaptations. P. 11-38, in R. J. Naiman and D. L. Soltz, eds., Fishes in Nor~h American Deserts. John Wiley and Sons, New York. Snyder, J. O. 1915. Notes on a collection of fishes made by Dr. Edgar A. Mearns from rivers tributa~y to the Gulf of Califomia. Proceedings of the U.S. National Museum 49: 573-586. Spofford, W. O., A. L. Parker, and A. V. Kneese, eds. 1980. Energy Development in the Southwest~'roblems of Water, Fish and Wildlife in the Upper Colorado River Basin, 2 Volumes. Resources for the Future, Washington, D.C. Stanford, J. A., and J. V. Ward. 1986a. The Colorado River system. P. 353-374, in B. R. Davies and K. F. Walker, eds., The Ecology of River Systems. Dr. SV. Junk, Dordrecht, The Netherlands. Stanford, J. A., and J. V. Ward. 1986b. Reservoirs of the Colorado system. P. 375-383, Ibid. Stanford, J. A., and J. V. Ward. 1986c. Fishes of the Colorado system. P. 385-402, Ibid. Stegner, W. 1954. Beyond the Hundredth Meridian~ohn Wesley Powell and the Second Opening of the West. Houghton-MifRin Company, Boston. 438 p. Stone, J. L. 1965. Tailwater fisheries investigations-creel census of the Colorado River below Glen Canyon Dam, July 1, 1964-June 30, 1965. Arizona Game and Fish Department, Colorado River Storage Project (Public Law 485, Section 8) Report. 13 p. Stone, J. L. 1966. Ibid. July 1, 1965-June 30, 1966. Ibid. 13 p. Stone, J. L. 1967. Ibid. July 1, 1966-June 30, 1967. Ibid. 33 p.

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NATIVE FISIlES... 175 Stone, J. L. 1968, Ibid. July 1, 1967-June 30, 1968. Ibid. 35 p. Stone, J. L. 1969. Ibid. July 1, 1968-June 30, 1969. Ibid. 47 p. Stone, J. L. 1971. Ibid. July 1, 1970-June 30, 1971. Ibid. 47 p. Stone, J. L. 1972. Ibid. July 1, 1971 -June 30, 1972. Ibid. 23 p. Stone, J. L., and N. L. Rathbun. 1967. Tailwater fisheries investigations, creel census and limnological study of the Colorado River below Glen Canyon Dam, July 1, 1966-Junc 30, 1967. Arizona Game and Fish Department, Phoenix. 54 p. Stone, J. L., and N. L. Rathbun. 1968. Ibid. July 1, 1967-June 30, 1968. Ibid. 56 p. Stone, J. L., and N. L. Rathbun. 1969. Ibid. July 1, 1968-June 30, 1969. Ibid. 60 p. Suttkus, R. D., and G. H. Clemmer. 1977. The humpback chub, Cila cypha, in the Grand Canyon area of the Colorado River, Occasional Papers of the Tulane University Museum of Natural History 1: 1-30. Suttkus, R. D., and G. H. Clemmer. 1979. Fishes of the Colorado River in Grand Canyon National Park. Pages 599-604, in Proceedings of the First Annual Conference on Scientific Research in the National Parks. U.S. National Park Service Transactions and Proceedings Series 5. Suttkus, R. D., G. H. Clcmmer, C. Jones, and C. R. Shoop, 1976. Survey of fishes, mammals and herpetofauna of the Colorado River in Grand Canyon. Colorado River Research Series Contribution (Grand Canyon National Park, Grand Canyon, Ariz.) 34:1-48. Swales. S. 1989. The use of instream habitat improvement methodology in mitigating the adverse effects of river regulation on fisheries. P. 185-208, in J. A. Gore and G. E. I'ctts, eds., Alternatives in Regulated River Management. Chemical Rubber Company Press, Boca Raton, Fla. Sykes, G. 1937. The Colorado Delta. Publications of the Carnegie Institution of Washington 460. 193 p. Taubert, B. Do 1985. Changes in the trout fisheries of the lower Colorado Roar in Arizona 1'. 175-180, in F. Richardson and R.H. Hamre, tech. coords., Wild Trout Symposium, III. U.S. Government Printing Office, Washington, D.C. Tyus, H. M. 1985. Homing behavior noted for Colorado squawfish. Copeia 1985:213-215. Tyus, H. M. 1986. Life strategies in the evolution of the Colorado squawfish (Ptychocheil~s lucius). Great Basin Naturalist 46:656-661. Tyus, H. M. 1987. Distribution, reproduction and habitat use of the razorback sucker in the Green River, Utah, 1979-86. Transactions of the American Fisheries Society 116: 111-116. Tyus, H. M. 1988. Long-term retention of implanted transmitters in Colorado squawfish and razorback suckers. North American Journal of Fisheries Management 8:264-267. Tyus, H. M. 1991. Management of the Colorado squawfish. In press, in W. L. NIincklcy and J. E. Deacon, eds., Battle Against Extinction: Native Fish Management in the American West. University of Arizona Press, Tucson. Tyus, H. M., R. L. Jones, and L. M. Tnnca. 1987. Green River rare and cndangcrcd fish studies, 1982-1985. Final Report, Colorado River Fishes Monitoring Project. U.S. Fish and Wildlife Service, Vernal, Utah. 117 p. Tyus, H. M., and C. A. Karp. 1989. Habitat use and streamflow needs of rare and endangered fishes, Yampa River, Colorado. U.S. Fish Wildlife Service Biological Report 89(14):1-27. Tyus, H. M., and C. W. McAda. 1984. Migration, movements, and habitat prcfcrcnccs of Colorado squawfish, Ptychocheilus Lucius, in the Green, White, and Yampa rivers, Colo- rado and Utah. The Southwestern Naturalist 29:289-299. Tyus, H. M., and W. L. Minckley. 1988. Migrating Mormon crickets, Anabrus simplex (Or- thoptera: Tettigoniidae), as food for stream fishes. Great Basin Naturalist 48:24-30. ., . , .. r ~ ---a ~~ Glen U.S. U.S. Fish and Wildlife Service. 1978. Memorandum (Biological opinion of the effects of Canyon Dam on the Colorado River as it affects endangered species) to Regional Director, Bureau of Reclamation, Salt Lake City. U.S. Fish and Wildlife Service, Albuquerque. 7 p.

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176 COLORADO RIVER ECOLOGY AND DAM MANAGEMENT U.S. Fish and Wildlife Service. 1980. Aquatic study Colorado River from Lee's Ferry to the southern International Boundary, and selected tributaries, Arizona, Califomia, Nevada. Special report on distribution and abundance of fishes of the lower Colorado River. Final Report, U.S. Water and Power Resources Service (Bureau of Reclamation) Contract 97-03-X0066, Lower Colorado River Region, Boulder City, Nevada. U.S. Fish Wildlife Service, Phoenix. 157p. U.S. Fish and Wildlife Service. 1981. Aquatic study of the lower Colorado River. Ibid. 278 p. U.S. Fish and Wildlife Service. 1987. Final recovery implementation program for endan- gered fish species in the upper Colorado River basin. U.S. Fish and Wildlife Service, Denver. 74 p. U.S. Fish and Wildlife Service. 1989a. Title Wildlife and Fisheries. Part 17 Endangered and threatened wildlife and plants. Subpart B lists. Endangered and threatened wildlife (CFR 17.11 and 17.12). U.S. Government Printing Office, Washington, D.C. U.S. Fish and Wildlife Service. 1989b. Humpback chub, Gila cypha, recovery plan (technical review draft). Region 6, U.S. Fish and Wildlife Service, Denver. 56 p. U.S. Fish and Wildlife Service. 1989c. Bonytall chub, Gila elegans, recovery plan (technical review draft). Ibid. 49 p. U.S. Fish and Wildlife Service. 1989d. Colorado squawfish Ptychocheilus lucius, recovery plan (technical review draft). Ibid. 78 p. U.S. Fish and Wildlife Service. 1990. Endangered and threatened wildlife and plants; Proposal to determine the razorback sucker ~yrauchen texanus) to be an endangered species. Fed- eral Register 55:21154-21161. Usher, H. D., S. W. Carothers, C. O. Minckley, and J. W. Jordan. 1980. Age and growth rate of the flannelmouth sucker, Catostomus latipinnis and bluehead mountain sucker, Pantosteus discobolus, in the Colorado River, Grand Canyon National Parlc. P. 163-175, in Proceed- ings of the Second Annual Conference on Scientific Research in the National Parks. U.S. National Park Service Transactions and Proceedings Series 8. U.S. National Park Service. 1977. Grand Canyon National Park, Arizona. Natural Resource Management Plan and Environmental Assessment. Grand Canyon National Park, Grand Canyon, Arizona 127 p. Valdez, R. A. 1990. The endangered fish of Cataract Canyon. Report U.S. Bureau of Reclama- tion Contract 6-CS-40-03980. BID/WEST, Incorporated, Logan, Utah. 94 p., 8 appendices. Valdez, R. A., and G. C. Clemmer. 1982. Life history and prospects for recovery of the humpback and bonytail chubs. P. 109-119, in W. H. Miller, H. M. Tyus, and C. A. Carlson, eds., Fishes of the upper Colorado River system: Present and future. Westen~ Division, American Fisheries Society, Bethesda, Md. Valdez, R. A., P. G. Magnan, R. Smith, and B. Nilson. 1982a. Upper Colorado River investiga- tions (Rifle, Colorado, to Lake Powell, Utah), P. 101-279, in W. H. Miller, J. J. Valentine, D. L. Archer, H. M. Tyus, R. A. Valdez, and L. R. Kaeding, 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. Valdez, R. A., P. G. Maqnan, M. McInery, and R. P. Smith. 1982b. Tributary report: Fishery investigations of the Gunnison and Dolores rivers. P. 321-362, Ibid. Vanicek, C. D. 1967. Ecological studies of native Green River fishes below Flaming Gorge Dam, 1964-1966. Unpublished Doctoral Dissertation. Utah State University, Logan, Utah. 124p. Vanicek, C. D., and R. H. Kramer. 1969. Life history of the Colorado squawfish, Ptychocheilus lucius, and the Colorado chub, Gila robusta, in the Green River in Dinosaur National Monument. 1964-1966. Transactions of the American Fisheries Society 98:193-208. Vanicek, C. D., R. H. Kramer, and D. R. Franklin. 1970. Distribution of Green River fishes in

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NATIVE FISHES... 177 Utah and Colorado following closure of Flaming Gorge Dam. The Southwestern Naturalist 14:297-315. Wallen, I. E. 1951. The direct effect of turbidity on fishes. Bulletin of the Oklahoma Agricul- tural and Mechanical College 48:1-27. Wallis, O. L. 1951. The status of the fish fauna of the Lake Mead National Recreation Area, Arizona-Nevada. Transactions of the American Fisheries Society 80:84-92. Welcomme, R. L. 1989. Floodplain fisheries management. P. 209-233, in J. A. Gore and G. E. Pelts, eds., Alternatives in Regulated River Management. Chemical Rubber Company Press, Boca Raton, Fla. Weatherford, G. D., and F. L. Brown, eds. 1986. New Courses for the Colorado River: Major Issues for the Next Century. University of New Mexico Press, Albuquerque. 253 p. Williams, C. D., and J. E. Deacon. 1991. Ethics, federal legislation,, and litigation in the battle against extinction. In press, in W. L. Minckley and J. E. Deacon, eds., Battle Against Extinction: Native Fish Management in the American West. University of Arizona Press, Tucson. Williams, J. E., D. B. Bowman, J. E. Brooks, A. A. Echelle, R. J. Edwards, D. A. Hendrickson, and J. J. Landye. 1985. Endangered aquatic ecosystems in North American deserts with a list of vanishing fishes of the region. Journal of the Arizona-Nevada Academy of Science 20:1 -62. Williamson, R. R., and C. F. Tyler. 1932. Trout propagation in Grand Canyon National Park. Grand Canyon Nature Notes 7(2):11-16. Wiltzius, W. J. 1976. Some historic influences of resewairs and irrigation diversions on flows, temperatures and fish distributions in the Gunnison River. Colorado Division Wildlife Resources, Fort Collins. 100 p. Winn, H. E., and R. R. Miller. 1954. Native postlarval fishes of the lower Colorado River basin, with a key to their identification. California Fish and Game 40:273-285. Woodbury, A. M., ed. 1959. Ecological studies of the flora and fauna in Glen Canyon. Univer- sity of Utah Anthropological Papers 40:1-229. Woodbury, A. M., ed. 1963. Studies of biota in Dinosaur National Monument, Utah and Colorado. University of Utah, Division of Biological Science Miscellaneous Papers 1: 1-77. Worster, D. 1985. Rivers of Empire: Water, Aridity, and Growth of the American West. Pan- theon Press, New York. 402 p. Wydoski, R. S., K. Gilbert, K. H. Seethaler, C. W. McAda, and J. A. Wydoski. 1980. Anno- tated bibliography for aquatic resource management of the upper Colorado River ecosys- tem. U.S. Fish and Wildlife Service Research Publication 135:1-186. Wydoski, R. S., and J. Hamill. 1991. Evolution of a cooperative recovery program for endan- gered fishes in the upper Colorado River basin. In press, in W. L. Minckley and J. E. Deacon, eds., Battle Against Extinction: Native Fish Management in the American West. University of Arizona Press, Tucson.