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Colorado squawfish. Service data from 1981 to 1988 <br />indicated that spawning occurs with declining flows <br />(following spring runoff) and increasing temperatures <br />(Fig. 10) -about 26 days (range: 17-33 days) after <br />migration. Regression analyses and multiple regression <br />techniques were used to investigate possible <br />cause-effect relations between spawning period, <br />discharge, and temperature for 1981-88. Peak discharge <br />preceding spawn, and mean minimum temperatures <br />during spawn, were highly correlated with the spawning <br />period (r = 0.84 and r = 0.88, respectively; P < 0.05), <br />ostensibly because discharge, temperature, and <br />spawning period are correlates. We do not presume the <br />above relations represent all conditions necessary for <br />successful spawn-rather, spawning of Colorado <br />squawfish is a result of complex environmental and <br />biological influences and is not triggered by a single flow <br />or temperature event. For example, Nesler et al. (1988) <br />hypothesized that flow spikes from rainstorms during <br />spring runoff may be important influences on ovulation <br />and spawning in Colorado squawfish. <br />Colorado squawfish spawn in the lower 51.2 km of the <br />Yampa River, particularly in a reach extending from <br />Warm Springs Rapid (RK6.56) to the vicinity of Harding <br />Hole (RK 32; Tyus et al. 1982b, 1987; Wick et al. 1983; <br />Haynes et al. 1984; McAda and Tyus 1984). Numerous <br />captures of ripe fish in a 6.4 km reach near Mathers Hole <br />(RK 28.3) suggest that egg deposition and fertilization <br />may be concentrated in this area (Fig. 4), where large, <br />deep pools and eddies are intermingled with runs and <br />cobble bars of gravel, cobble, and boulder substrates. <br />Spawning behavior of Colorado squawfish was <br />divided into two phases: (1) cresting-staging phase in <br />pools or large shoreline eddies, where the fish may find <br />suitable resting and feeding habitat between spawning <br />forays or where males aggregate until females are ripe; <br />and (2) adeposition-fertilization phase on cobble bars, <br />where actual spawning occurs (Archer and Tyus 1984). <br />Breeding adults occupied pools or eddies having an <br />average depth and velocity of 2 m and 0.2 m/s, <br />respectively, and cobble bars with an average depth and <br />velocity of 1 m and 0.5 m/s (Table 4). <br />Breeding adults were most often concentrated in <br />river reaches containing deep pools, eddies, and cobble <br />(rubble) bars. Radio-tagged fish moved from pools or <br />eddies to cobble-gravel bars (where they presumably <br />spawned). This behavior is similar to that of spawning <br />northern squawfish (Beamesderfer and Congleton <br />1981). Turbid conditions in the Yampa River have <br />precluded direct observations of egg deposition; <br />however, cobbles removed from the substrate during <br />that time of year are clean of sediment and algae <br />Table 4. Depths and velocities taken at location of radio-tagged Colorado squawfish o~i Yampa River spawning <br />grounds, 1981, 1983, 1984, 1985, 1987, and 1988. <br />Depth (m) Velocity (m/s) <br />Number Number of <br />Yeara of fish contactsb Mean Range Mean Range <br />Resting-Stagingc <br />1981 7 74 1.80 (0.76-2.68) 0.39 (0.03-1.19) <br />1983 7 46 2.36 (1.07-5.79) 0.47 (0.09-1.46) <br />1984 5 261 1.70 (0.76-3.05) 0.17 (0.00-0.40) <br />1985 3 77 3.30 (0.76.27) 0.22 (0.00-0.30) <br />1987 4 22 1.97 (1.28-2.74) 0.14 (0.06-0.18) <br />1988 2 8 2.86 (2.74-3.05) 0.06 (0.06) <br />Weighted Mean 2.06 0.24 <br />Deposition-Fertilizationd <br />1981 7 65 1.16 (0.61-1.68) 0.59 (0.36-1.20) <br />1983 4 21 1.02 (0.55-1.22) 0.79 (0.27-1.04) <br />1984 6 82 0.89 (0.61-1.52) 0.40 (0.09-1.01) <br />1985 1 1 1.52 - 0.15 - <br />1988 3 12 0.87 (0.46-1.22) 0.48 (0.15-0.92) <br />Weighted Mean 1.01 0.52 <br />a No fish located in deposition-fertilization habitat in 1987. <br />b One contact =one 15-min period of observation. <br />c Includes eddy and pool habitat. <br />d Includes riffle and shallow run habitat over cobble and boulder substrate. <br />12 <br />