lzz
<br />Ft. Thomas on the mainstream of the Gila (Kirsch, 1889), and farther up-
<br />stream."
<br />The comparable decline in abundance of squawfish in the Colorado
<br />River mainstream, below Lake Mead, as indicated in the quotation by
<br />Miller (19616)'given above, has appeared to continue to essential ex-
<br />tinction. Minckley & Deacon (1968) heard of only two specimens from that
<br />region between 1962 and 1967 (one of which was seen and was from
<br />Lake Mead itself), and none has been reported subsequently.
<br />There is no published information on the life history of this fish in
<br />Arizona waters. Vanicek & Kramer (1969) have provided data, •~owever,
<br />from the Green River in Dinosaur National Monument, and some informa-
<br />tion may be gleaned from a number of general publications (Simon, 1951;
<br />Beckman, 1963; Sigler & Miller, 1963; and others). Prior to dams, squaw-
<br />fish moved upstream in "runs" just before the spawning period. This, along
<br />with their great size, presumably result in the name "salmon." Other
<br />squawfishes, P. grandis (Ayres) of the Sacramento River basin (Taft &
<br />Murphy, 1950) and P, oregonensis (Richardson) (citations below), also
<br />seem to "run" into streams to spawn, at least in the past. Reproduction
<br />by Colorado River squawfish in the Green River appeared to occur in July
<br />and August, attuned to increasing water temperatures and receding water
<br />levels. In the southern part of its range, and perhaps in the Green River,
<br />Utah, before moderation of temperatures induced by damming, spawning
<br />must have occurred earlier in the year. Young, 32 mm long, were taken
<br />in mid-May in Arizona (Miller, in Sigler & Miller, 1963). Spawning by
<br />other species of Ptychocheilus occurs in early to late spring.
<br />Actual spawning sites or breeding behavior of P. lucius has yet to be
<br />observed. The northern squawfish (P. oregonensis) spawns in lakes or
<br />streams of the Columbia River basin, in shallow water over gravel or rubble
<br />(Jeppson & Platts, 1959), or in streams over larger rubble in riffles or at
<br />the lower ends of pools in tributaries (Keating, in Vanicek & Kramer, 1969).
<br />Spawning by P. grandis seems to occur at the lower ends of pools, just
<br />above riffles, or in riffles (Taft & Murphy, 1950). Spawning behavior of
<br />P. oregonensis in Merwin Reservoir, Washington, has recently been de-
<br />scribed in detail by Patten & Rodman (1969). Fish congregated in in-
<br />creasingly greater numbers from June, to a peak of reproductive activities
<br />in the first week of July. They were over a specific area of rocky bottom,
<br />deeper than three meters, but above the thermocline. Swarms of males
<br />moved slowly and irregularly about 30 cm above the bottom, with indi-
<br />viduals darting abruptly upward, then returning to the swarm. The
<br />11F. W. Chamberlain's 1904 field notes report as follows: "Several years ago fish
<br />were abundant (near Safford, Graham County, in the Gila River). Then pools of
<br />sufficient depth for men to swim in existed. Salmon reached a weight of 35 Ib.,
<br />humpback and other suckers were abundant. None of these has been taken in the
<br />last two years. It is believed that minerals and concentrate-wash from the mines
<br />and works at Morenci and Clifton have killed the fish."
<br />123
<br />boulders beneath the swarm were cleaned, presumably by current created
<br />by the moving bodies, while those in other places were covered by sedi-
<br />ment. Females swam alone, peripheral to the swarm of males. When a
<br />female moved near, 2 to 3 males usually chased her as she accelerated
<br />to an open area, attempting to maintain position with their snout near
<br />the female's vent. A female followed by males dipped near the bottom,
<br />usually over a crevice, recurved her back, and spawned with vibratory,
<br />lateral movements. Males moved over and about her in a "solid mass,"
<br />presumably emitting sperm. Only a small number of eggs was released at
<br />a time, so a female must have performed the act repeatedly to deplete her
<br />ovaries. The eggs adhered to the substrate (as has been noted in other spe-
<br />cies); however, none was found on the upper surfaces of stones. Unpro-
<br />tected eggs were likely eaten by predators.
<br />Young of P, lucius appeared in August in all three years that Vanicek
<br />& Kramer (1969) studied the Green River population, 3 to 6 weeks after
<br />the estimated time of spawning. Growth of the young fish continued
<br />throughout the year, and by the end of their third summer juveniles were
<br />averaging about 160 mm in total length. The largest specimen taken by
<br />them was 600 mm total {ength, about 1.85 kg in weight, and was 11
<br />years of age.
<br />Foods of Colorado River squawflsh in the Green River consisted of
<br />crustaceans and aquatic dipteran larvae when the fish were shorter than
<br />50 mm total length. An increasing number of aquatic insects, plus terrestrial
<br />insects, was taken when the fish were between 50 and 100 mm, and other
<br />fishes predominated the diet of specimens longer than 101 mm. There
<br />was an increasing incidence of empty stomachs in larger fish, reflecting
<br />perhaps the sporadic nature of feeding o#ten found in piscivores. Large
<br />adults of P. lucius have been known to take carcasses of small animals
<br />and birds (Beckman, 1963), and they also have been caught on artificial
<br />lures (as has P. grandis; Taft & Murphy, 1950). Relatively complete studies
<br />of foods of other squawfish are those of Chapman & Quistorff (1938) and
<br />Thompson (1959) for P, oregonensis, and notes of Taft & Murphy (1950)
<br />for P. grandis. The last two species, and P. umpquae Snyder, are often
<br />thought to be severe competitors of, and predators upon, trouts and sal-
<br />mons, and have been subject to intensive research for eradication and
<br />control in areas where they are abundant (references already cited, and
<br />MacPhee & Ruelle, 1969). It is notable that the other three species of
<br />squawfishes appear to be maintaining populations in their native ranges,
<br />whereas P. lucius is declining; the reasons for this are unknown.
<br />As has been repeatedly implied, Colorado River squawfish, and
<br />most other species of this genus, are big-river fishes, characteristic of
<br />waters deeper than a meter and generally with strong to moderate cur-
<br />rents. Young frequent backwaters over almost any bottom type, where
<br />currents are slight or absent. -
<br />Carhart (1950), Miller (in Minckley, 1965a), Joseph Stone, Arizona
<br />Game and Fish Department (pers. comm.), and a number of the "old
<br />
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