83 Summary of Technical Literature Examining the Physiological Effects of Temperature fall spawning salmonids and has been documented for chinook of the Pacific Northwest ( Groves and Chandler 1999, Lindsay et al. 1986).

Although spawning for bull trout may begin as early as mid August, spawning activity is reported to be initiated when water temperatures begin to fall to 48.2 ° F ( 9 ° C) or lower ( McPhail and Murray 1979, Shepard et al. 1982, Kraemer 1994, Brenkemen 1998). Both the coastal rainbow and the redband trout spawn in the spring, stimulated by rising water temperatures. Behnke ( 1992) suggested that along the Pacific coast a water temperature of about 37.4 42.8 ° F ( 3 6 ° C) may initiate some spawning activity, but spawning does not usually occur until temperatures reach 42.8 48.2 ° F ( 6 9 ° C). Although this spawning activity would typically occur from late December through April, in some very cold headwater streams local temperatures may delay spawning until July or August for some stocks. Beschta et al. ( 1987) suggested that rainbow trout spawn between 35.9 and 68 ° F ( 2.2 and 20 ° C), Bell ( 1986) set the range at 35.9 66 ° F ( 2.2 18.9 ° C), and Piper et al. ( 1982) concluded the range was 50 55 ° F ( 10 12.8 ° C).

Conclusions for spawning. Egg mortality, alevin development linked to thermal exposure of eggs in ripe females or newly deposited in gravel, and egg maturation are negatively affected by exposure to temperatures above approximately 54.5 57.2 ° F ( 12.5 14 ° C). Therefore, a spawning temperature range of 42 55 ° F ( 5.6 12.8 ° C) ( maximum) appears to be a reasonable recommendation for Pacific salmon, unless colder thermal regimes are natural in any tributary.

SUPPORTING DISCUSSION AND LITERATURE   LETHAL EFFECTS What is the utility of UILT data and how has it been applied?

Upper incipient lethal temperature data were tabulated in NAS ( 1972) for juveniles and adults of many fish species. UILT values for many salmonid species have since been added to the literature; a cross section is summarized in Table 4, extracted from McCullough ( 1999). The UILT values correspond to the highest acclimation temperatures, and consequently, are very similar to UUILT values. Given prior acclimation to temperatures lower than listed in the table, however, the UILT values would likely be lower. This means that in the field, mortality can be induced at temperatures significantly lower than UUILT levels.

Studies of the effect of elevated water temperature on survival of a wide variety of salmonids using transfer to high constant temperature ( UILT experiments) show much consistency among species. In those tests in which acclimation temperature was 68 ° F ( 20 ° C) and the UILT was approximately equal to the UUILT, UILT values found ranged from 73.4 to 80.6 ° F ( 23 27 ° C). Redband trout tend to be the most heat resistant of the salmonids; UILT values for all other species ranged from 73.4 to 78.8 ° F ( 23 26 ° C).

NAS ( 1972) recommended that for any acclimation temperature, short  term exposure be limited to UILT ( factor of safety, 3.6 ° F [ 2 ° C]). This assumes that at the UILT temperature, 50% of the population would die within the test period ( at least 1,000 min), but if the temperature is reduced by 3.6 ° F ( 2 ° C), no mortalities would occur in this time period. Although this assumption may generally be valid, it also relies on no incidence of disease or other sublethal effects. When 84 Summary of Technical Literature Examining the Physiological Effects of Temperature this method was proposed, cumulative effects of repeat exposure to high temperatures were not well known ( see DeHart 1975, Golden 1975, Golden 1976, Golden and Schreck 1978).

Although temperatures of 73.4 78.8 ° F ( 23 26 ° C) are generally considered the UUILT for most salmonids, UILT values are typically 1.8 3.6 ° F ( 1 2 ° C) less than UUILT. The UILT values are incipient lethal temperatures that correspond to acclimation temperatures lower than the UUILT. Because we can never assume that fish in the field will be acclimated to the highest acclimation temperature, the more appropriate lethal temperature in the field may be up t o 3.6 ° F ( 2 ° C) less than UUILT. The factor of safety would then have to be applied t o this value, and even then the additional sublethal or cumulative lethal concerns remain.

The 73.4 78.8 ° F ( 23 26 ° C) UUILT range for salmonids applies to the juvenile life stage. Although information on salmon adults is much more limited, it indicates that adults are far more sensitive than juveniles to high temperatures. Becker ( 1973) identified the thermal tolerance of chinook jacks to be 69.8 71.6 ° F ( 21 °  22 ° C) on the basis of a 168 h TLM test. Coutant ( 1970) identified the incipient lethal temperature for chinook jacks as 71.6 ° F ( 22 ° C) with prior acclimation to 66.2 ° F ( 19 ° C) ( estimated from ambient river temperatures). Columbia River steelhead, acclimated to a river temperature of 66.2 ° F ( 19 ° C) had a lethal threshold of 69.8 ° F ( 21 ° C) ( Coutant 1970). These lethal limits are 9.9 ° F ( 5.5 ° C) lower than for juvenile rainbow acclimated to 64.4 ° F ( 18 ° C) ( Alabaster and Welcomme 1962, as cited by Coutant 1972).

Servizi and Jensen ( 1977) found that the geometric mean survival times ( GMST) for adult sockeye were less than for juveniles. They also reported that the median survival times ( MST) for adult coho found by Coutant ( 1969) were similar to those of sockeye over the exposure range 80.6 86 ° F ( 27 30 ° C). The GMST for adult sockeye was 1,000 min at 75.2 ° F ( 24 ° C) with acclimation at 60.4 64.9 ° F ( 15.8 18.3 ° C). Survival time at 78.8 ° F ( 26 ° C) was only 100 min. Time to loss of equilibrium and survival time of adults were plotted vs. exposure temperature on the same graph. The curve for loss of equilibrium was considerably lower than the time to death curve. For this reason, Servizi and Jensen ( 1977) considered the loss of equilibrium temperature more ecologically significant. Furthermore, because sockeye exposed to temperatures of approximately 64.4 69.8 ° F ( 18 21 ° C) become highly susceptible to Flexibacter columnaris, researchers took this temperature range as a greater thermal threat to continued sto ck survival.

Although UUILT or UILT temperatures are well known and consistent for the various salmonids, they are probably not useful in setting temperature standards. Certainly they represent the upper limits to tolerance, but salmonids in a stream system tend to be restricted to maximum temperatures that are 3.6 7.2 ° F ( 2 4 ° C) lower than UILT values. In general, a maximum temperature of 71.6 75.2 ° F ( 22 24 ° C) represents the normal upper temperature limit in the field ( see McCullough 1999). As this limit is approached, juvenile density declines t o zero. Using the presence/ absence threshold as a temperature standard for salmon habitat can only be done when density is at or near zero.

Although the UILT has limited value in establishing the temperature standard itself, the a and b coefficients for a given acclimation temperature are useful in estimating exposure times that will result in 50% mortality. In addition, NAS ( 1972) recommended ( MWAT) as an index to tolerable prolonged exposures. This index was estimated as either: 85 Summary of Technical Literature Examining the Physiological Effects of Temperature [( opt. temp. + zng temp.)/ 2],

where the zng or zero net growth temperature is that temperature which results in zero net growth of a population ( i. e., subt racting tissue lost as population mortality from that added in growth)

or opt. temp. + ( UILT ! opt. temp)/ 3.

Each formula is based o n the assumption that adequate growth rates can be maintained if the weekly average maximum temperature falls between the optimum and the UILT or the zero net growth temperature. However, the decline in growth rate can be very steep if temperature is above the optimum. Consequently, limiting reductions in growth rates to, for example, 80% of maximum levels can lead to much greater reductions in growth, given errors estimating the relationship or managing temperature in a watershed.

Are there potential weaknesses in reliance on MWAT?

Hokanson et al. ( 1977) advised caution in using short term exposure experiments to calculate long term exposures, such as with MWAT. They reported for O. mykiss that, given a physiological optimum of 60.8 64.4 ° F ( 16 18 ° C) and a UILT of 78 ° F ( 25.6 ° C) ( at 60.8 ° F [ 16 ° C] acclimation), one would calculate an MWAT of 66.2 ° F ( 19 ° C) and a maximum temperature ( applying the 2 ° C safety factor of Coutant 1972) of 75.2 ° F ( 24 ° C) for short term exposure. Measurement of rainbow trout growth showed that at a fluct uating temperature of 71.6 ± 6.8 ° F ( 22 ± 3.8 ° C) specific growth rate was zero. Under this temperature regime mortality rate was 42.8%/ d during the first 7 d. For experiments within the optimum range ( 59.9 63.1 ° F [ 15.5 ° C 17.3 ° C] for a fluct uating regime), average specific mort ality was 0.36%/ d. Combining data on specific growth and mortality rates, the authors were able to predict yield for a hypothetical population under t he temperature regimes. A rainbow trout population would exhibit zero increase over a 40 d period ( maintenance) at a constant temperature of 73.4 ° F ( 23 ° C) and a fluctuating temperature with a mean of 69.8 ± 6.8 ° F ( 21 ± 3.8 ° C) because growth balances mortality. Several sources report temperatures of 69.8 73.4 ° ( 21 23 ° C) as the upper limit of rainbow trout distribution in the field ( Hokanson et al. 1977). Numerous authors have reported upper limits to salmonid distribution as approximately 71.6 75.2 ° F ( 22 24 ° C).

With this laboratory information and corroborating field information, Hokanson et al. ( 1977) recommended a mean weekly temperature of 62.6 ± 3.6 ° F ( 17 ± 2 ° C) for rainbow trout so that maximum yield is not reduced more than 27% under normal fluctuating temperature regimes. Production has been shown to be substantially reduced even just above the physiological optimum. This paper has great significance. It was published 5 years after the National Academy of Sciences recommended the use of MWAT to establish prolonged exposure temperatures. The NAS acknowledged that growth rate should be expressed as net biomass gain or net growth. Yield is that portion o f the population available to humans; t he remainder is lost as mortality, which can be substantial if temperatures are high. Also, if temperatures are high, much of the energy assimilated from food is lost as excessive metabolism. If the MWAT is 66.2 ° F ( 19 ° C), 86 Summary of Technical Literature Examining the Physiological Effects of Temperature and yield is reduced 27% from maximum, even at a mean weekly temperature of 62.6 ± 3.6 ° F ( 17 ± 2 ° C) it is obvious that MWAT is not protective.

In addition to concern for the inadequacy of the MWAT, this criterion also covered reproduction and development needs of salmonids. A quo te from NAS ( 1972) is useful:

Uniform elevations of temperature by a few degrees during the spawning period, while maintaining short term temperature cycles and seasonal thermal patterns, appear to have little overall effect on the reproductive cycle of resident aquatic species, other than to advance the timing for spring spawners or delay it for fall spawners. Such shifts are often seen in nature, although no quantitative measurements of reproductive success have been made in this connection.

However, significant recent research has shown that calculated MWAT temperatures ( e. g., 66.2 ° F ( 19 ° C) for rainbow trout) result in damage to gametes during reproductive stages. On the basis of these technical findings, any temperature criterion that can result in a 27% reduction in biomass and affect gamete viability must be questioned.

How can UILT data be evaluated against UUILT data?

Temperatures as low as 73.4 ° F ( 23 ° C) have been found to produce 50% mortality ( LT50) over the course of a week in rainbow trout acclimated to very cold ( 39.2 ° F [ 4 ° C]) waters ( Sonski 1982, Threader and Houston 1983 as cited in Taylor and Barton 1992), with the lethal temperature rising to 75.2 ° F ( 24 ° C) in moderately cold water acclimated 42.8 51.8 ° F ( 6 11 ° C) fish ( Black 1953, Stauffer et al. 1984, Bidgood 1980 as cited in Taylor and Barton 1992). However, at most acclimatio n temperatures likely to be encountered during the spring through fall seasons ( 53.6 68 ° F [ 12 20 ° C]), lethal levels are consistently in the range of 77 78.8 ° F ( 25 26 ° C) ( Bidgood and Berst 1969, Hokanson et al. 1977). With cautious acclimation to temperatures in the range of 73.4 75.2 ° F ( 23 24 ° C), rainbow trout may not experience LT50 effects until a week at 78.8 ° F ( 26 ° C) ( Charlon et al. 1970 as cited in Grande and Anderson 1991). Even with careful acclimatio n, 77 ° F ( 27 ° C) results in high or complete mortality in less than 24 hours ( Charlon et al. 1970), and temperatures of 84.2 86 ° F ( 29 30 ° C) result in 50% mortality in 1 2 hours ( Kaya 1978, Craigie 1963, Alabaster and Welcomme 1962 as cited in Taylor and Barton 1992).

How can prolonged exposure to cyclic temperatures be evaluated?

Under fluctuating temperature test conditions, rainbow trout have experienced 50% mortality in a week of daily cycles from 69.8 to 77 ° F ( 21 27 ° C) ( Lee 1980). Sonski ( 1983), however, was able to culture rainbow trout in ponds that reached 84 ° F ( 28.9 ° C), and Chandrasekaran and Subb Rao ( 1979) reported that rainbow trout were largely able to survive in rearing ponds with months of daily maximum temperatures of 78.8 84.2 ° F ( 26 29 ° C).

It seems important, given the low lethal levels reported in the literature, to evaluate whether individual research results would unduly influence the temperature recommendations. In Figures 3 and 4, lethality data for salmon and char species ( extracted from the summary by Hicks 2000) are combined and examined in two different ways to develop a stronger basis for regional daily 87 Summary of Technical Literature Examining the Physiological Effects of Temperature maximum criteria.

In Figure 3, all of the available 7 d LT50 data ( 50% of test organisms die over a 7 d constant exposure test) for char and salmonids are presented by acclimation temperature. This distribution is then used to make criteria recommendations for acclimation temperatures. At low acclimation temperatures, constant exposure to just above 72.5 ° F ( 22.5 ° C) would be expected to result in 50% mortality over a week. Reducing this value to a level where no lethality would be expected to any adults or juveniles would result in a daily maximum not to exceed 68.9 ° F ( 20.5 ° C).

Figure 4 considers resistance against short exposures to high temperatures, as might occur in a natural fluctuating stream environment. Resistance time is very important to estimating potential lethality; it is the time spent above a lethal threshold that determines whether short term lethal effects will occur. Different peak temperatures ( e. g., 71.6, 75.2, 80.6, and 86 ° F [ 22, 24, 27, and 30 ° C]), may be lethal to an organism, but the organism can likely withstand these temperatures for variable lengths of time. A populat ion of fish may be able to withstand 69.8 ° F ( 21 ° C) for 7 d of constant exposure without any mortality, but have 50% of the population die after 2 d at 75.2 ° F ( 24 ° C). At 80.6 ° F ( 27 ° C), 50% mortality may occur after less than 2 h of exposure, and at 86 ° F ( 30 ° C) complete mortality may occur in just a few minutes.

In considering the effect o f repeat ed hot days, it is important to incorporate cumulative effects. DeHart ( 1975) found that lethal effects depend on the area of the temperature time curve that is above a fish's UILT. Thermal effects accumulate over several days when the daily temperature cycle fluctuates above the UILT, and the time above the UILT influences the thermal resistance time regardless of any lower test temperatures. In other words, the ability of a fish to resist a single day's exposure to a lethal temperature may not be sufficient, and 15 minutes spent at 7.2 ° F ( 4 ° C) over the UILT is of more consequence than the same time spent at 3.6 ° F ( 2 ° C) over the UILT.

In Figure 3, LT50 results are plotted for durations of 1 hour or less. At acclimation levels less than 53.6 ° F ( 12 ° C), 50% mortality can be expected at 77 ° F ( 25 ° C) with a 1 h exposure, or at 75.5 ° F ( 24.2 ° C) with a 2 h exposure. Reducing these values to levels where no lethality would be expected would result in temperatures not exceeding 73.4 or 71.6 ° F ( 23 or 22 ° C), respectively. Because adults are considered more sensitive than juveniles ( all of the 1 h or less data were for juvenile fish), and the effects of lethal exposures are cumulative, we can assume that death may occur with repeated exposure to daily maximum temperatures greater than 69.8 71.6 ° F ( 21 22 ° C). This estimate is very similar to the results ( 68.9 70 ° F [ 20.5 21.1 ° C]) at low acclimation temperatures in the approach shown in Figure 2.

Acclimation Temperature ° F ( ° C) Combined LT50 for all Salmonids Esti mated LT1 with NAS Adjustment 41 ( 5) 72.46 ( 22.48) 68.9 ( 20.5)

50 ( 10) 73.56 ( 23.09) 70 ( 21.1)

39 ( 15) 74.66 ( 23.7) 71.06 ( 21.7) 88 Summary of Technical Literature Examining the Physiological Effects of Temperature 68 ( 20) 75.75 ( 24.31) 72.14 ( 22.3)

Figure 3. Combined lethality data for all salmonid species ( based on 7 d LT50 constant temperature exposure test results).

SUPPORTING DISCUSSION AND LITERATURE   SUBLETHAL AND LETHAL EFFECTS FOR NATIVE CHAR, REDBAND TROUT, AND CUTTHROAT TROUT SPECIES What are the thermal requirements of bull trout and Dolly Varden?

Incubation. For bull trout, McPhail and Murray ( 1979) compared egg survival and water temperature and reported the highest egg survival to hatching ( 80 95%) in water temperatures of 35.6 39.2 ° F ( 2 4 ° C). Shortest hatch, largest alevins, and largest hatching fry were also associated with these low temperatures 35.6 39.2 ° F ( 2 4 ° C).

Research suggesting that spawning does not peak until temperatures fall to below 44.6 ° F ( 7 ° C) is consistent with the results o f studies determining temperatures necessary for the 89 Summary of Technical Literature Examining the Physiological Effects of Temperature successful incubation of char eggs. These studies show that char require temperatures below 42.8 ° F ( 6 ° C) to achieve optimal egg survival ( Buchanan and Gregory 1997). It is generally agreed that poor survival occurs at temperatures above 44.6 46.4 ° F ( 7 8 ° C). Under t est conditions where temperatures were held constant, 46.4 50 ° F ( 8 10 ° C) resulted in very poor survival of eggs ( 0% 20%) in tests by McPhail and Murray ( 1979), and test temperatures in the range of 44.6 51.8 ° F ( 7 11 ° C) were reported Time to LT50 Temperature ( C) 1 sec 34.32 30 sec 30.42 1 min 29.62 60 min 24.99 120 min 24.2 Figure 4. Instantaneous mortality for char and salmon combined ( based on LT50 data for exposures of less than 1 hour and acclimation to < 12C).

to result in poor survival in hatchery culture by Brown ( 1985). McPhail and Murray ( 1979) found a temperature of 42.8 ° F ( 6 ° C) to produce variable survival rates ( 60% 90%), and the range of 90 Summary of Technical Literature Examining the Physiological Effects of Temperature 35.6 39.2 ° F ( 2 4 ° C) produced the best survival ( 80% 95%). In studies on the related species of Arctic char, Humpesch ( 1985) reported optimal incubation to occur at 41 ° F ( 5 ° C).

In conclusion, for bull trout, temperatures falling to 48.2 ° F ( 9 ° C) and below may initiate spawning, but colder temperatures during incubation and yolk absorption produce the largest size and greatest number of fry ( McPhail and Murray 1979). Although spawning tends to peak at 44.6 ° F ( 7 ° C), water temperatures continue to decline as the spawning season progresses and drop toward the optimum incubation temperatures of 35.6 42.8 ° F ( 2 6 ° C). Bull trout tend to select redds directly adjacent to or below areas of groundwater upwelling, resulting in relatively constant cold water temperatures for egg incubation with little diel fluctuation ( Baxter and Hauer 2000).

Growth. In a laboratory study by McMahon et al. ( 1999), limited rations lowered the optimal temperature for growth. For satiation fed and 66% of satiation fed juvenile bull trout, optimum growth occurred at a temperature range of 53.6 60.8 ° F ( 12 16 ° C). When energy availability was low ( one third satiation fed fish), maximum growth occurred at lower temperatures ( 46.4 53.6 ° F [ 8 12 ° C]). In a related species, Arctic char, the upper thermal limits to both feeding and growth were 70.7 71.2 ° F ( 21.5 21.8 ° C) ( Thyrel et al. 1999).

In a study analyzing the temperature effects on bull trout distribution in 581 sites, Rieman and Chandler ( 1999) found that juvenile/ small bull trout appeared most likely to occur at summer mean temperatures of 42.8 48.2 ° F ( 6 9 ° C) or single maximums of 51.8 57.2 ° F ( 11 14 ° C). When given a choice of temperatures from 46.4 to 59 ° F ( 8 15 ° C) in a large plunge pool, juvenile bull trout showed a clear preference for the coldest water available ( 6.4 48.2 ° F [ 8 9 ° C]) ( Bonneau and Scarnecchia 1996).

Migration. Upstream spring migration of adult bull trout may be related to water temperatures and flows. In Rapid River, Idaho, a review of trap counts and temperature for 1985 through 1992 reported a general trend of increasing upstream bull trout counts as water temperatures reached 50 ° F ( 10 ° C) ( Elle et al. 1994). McPhail and Murray ( 1979) found peak upstream movement coincided with water temperatures of 50 53.6 ° F ( 10 12 ° C).

Spawning. Bull trout spawning areas are often associated with cold water springs, groundwater infiltration, and the co ldest st reams in a given watershed ( Pratt 1992, Rieman and McIntyre 1993, Rieman et al. 1997). Bull trout spawning is initiated as temperatures drop to 48.2 ° F ( 9 ° C) or lower, and egg mortality is lowest and alevin development is strongest at colder temperatures ( McPhail and Murray 1979). In Indian Creek, tributary to the Yakima River, bull trout spawning activity peaked when stream temperatures were 42.8 46.4 ° F ( 6 8 ° C) ( James and Sexauer 1994). In the North Fork Skokomish River, bull trout spawned in October after water temperatures dropped below 43.7 ° F ( 6.5 ° C) ( Brenkman 1998). Mean daily river temperatures ranged from 38.3 to 45.5 ° F ( 3.5 7.5 ° C) during the remainder of the spawning period. This does not differ significantly from descriptions of temperatures initiat ing bull trout spawning in Montana ( Shepard et al. 1982), Oregon ( Ratliffe 1992), or Washington ( Kraemer 1994).

Although spawning for bull trout may begin as early as mid August, spawning activity is reported to be initiated when water temperatures begin to decrease and fall to 48.2 ° F ( 9 ° C) or 91 Summary of Technical Literature Examining the Physiological Effects of Temperature lower and does not peak until temperatures fall below 44.6 ° F ( 7 ° C) ( McPhail and Murray 1979, Shepard et al. 1982, Kraemer 1994, Brenkemen 1998). Kraemer ( 1994) noted that when stream temperatures rise to above 46.4 ° F ( 8 ° C) once spawning has been initiated, spawning usually stops or slows.

Bull trout require a narrow range of cold temperatures to rear and reproduce and may thrive in waters too cold for other salmonid species ( Balon 1980). McPhail and Murray ( 1979) reported that 0% 20%, 60% 90%, and 80% 95% of bull trout eggs from a British Columbia river survived to hatching at water temperatures of 46.4 50, 42.8, and 35.6 39.2 ° F ( 8 10, 6, and 2 4 ° C), respectively. In Montana, Weaver and White ( 1985) found that 39.2 42.8 ° F ( 4 6 ° C) was needed for bull trout egg development. Buchanan and Gregory ( 1997) defined a range o f 33.8 42.8 ° F ( 1 6 ° C) that would meet bull trout egg incubation requirements in Oregon.

Although data are not shown directly for char species, other salmonids are known to undergo some conditioning in the early stage of incubation that allows excellent survival at very low temperatures. Where conditioning does not occur, and t he eggs are incubated at an early stage at very low temperatures, significant reductions in survival have been noted ( Murray and Beacham 1986, Seymour 1956). Thus even if 35.6 ° F ( 2 ° C) is suboptimal at a constant incubation temperature, natural seasonal declines in temperature to 35.6 ° F ( 2 ° C) in the incubation period may not decrease survival. This assumption is supported by work showing that newly hatched Arctic char ( Salvelinus alpinus) alevins are tolerant of temperatures near 32 ° F ( 0 ° C) ( Baroudy and Elliott 1994) and that the lower limit for hatching in Arctic char is less than 33.8 ° F ( 1 ° C).

On the basis of the information examined, the initiation of spawning behavior and in vivo egg development will be fully supported by keeping maximum temperatures in the spawning areas below 44.6 46.4 ° F ( 7 8 ° C) during the spawning season. Given that excellent survival has been noted in test s at 42.8 and 43.7 ° F ( 6 and 6.5 ° C), that some increased problems with disease may be initiated at the higher end of this range, and that a variety of impacts to spawning have been noted above 44.6 ° F ( 7 ° C), it appears that constant or acclimation temperatures in the range of 37.4 42.8 ° F ( 3 6 ° C) are optimal for the incubation of char. Because char are highly resistant to low temperatures and low temperatures discourage disease organisms, water temperatures that swiftly decline to 35.6 39.2 ° F ( 2 4 ° C) as the incubation season progresses appear highly favorable.

What are the therm al requirem ents for Lahontan cutthroat tro ut?

Growth. Laboratory studies of growth conducted at constant temperatures showed that growth remained the same at temperatures of 55.4, 68, and 71.6 ° F ( 13, 20, and 22 ° C) ( Dickerson et al. 1999, as cited in Dunham 1999). Growth was significantly reduced at 75.2 ° F ( 24 ° C). Tests done under a fluctuating temperature regime of 68 78.8 ° F ( 20 26 ° C) with a daily mean of 73.4 ° F ( 23 ° C) for 1 wk showed growth rates were lower under this temperature regime than for fish exposed to constant temperatures of 55 and 68 ° F ( 13 and 20 ° C). The growth rates under the fluctuating regime were similar to growth rates of fish held at a constant 73.4 ° F ( 23 ° C).

Thermal stress   heat shock proteins. Lahontan cutthroat trout begin to produce heat shock proteins immediately when exposed to 78.8 ° F ( 26 ° C) water temperature. Fish exposed to 75.2 ° F ( 24 ° C) water temperature began to produce heat shock proteins within 24 h. Fish exposed 92 Summary of Technical Literature Examining the Physiological Effects of Temperature to 71.6 ° F ( 22 ° C) water temperature did not produce heat shock proteins, even after 5 d of exposure.

Occurrence. Dunham ( 1999) found that the distribution limit of most Lahontan populations corresponded closely to a maximum summer water temperature of 78.8 ° F ( 26 ° C). The Willow Creek population in Oregon occurred in water with daily maximum temperatures up to 83 ° F ( 28.4 ° C).

Lethal effects. It has often been assumed that Lahontan cutthroat trout have a greater tolerance for warm water than other salmonids because of their geographic distribution in warm climates. Although there are not abundant studies of Lahontan cutthroat thermal requirements, good evidence indicates that this species is comparable to other salmon and trout in its response to warm water.

Critical thermal maximum ( CTM) tests of thermal resistance were conducted on two strains of Lahontan cutthroat t rout by Vigg and Koch ( 1980). The two strains tested were Marble Bluff and the Summit Lake strain found in Pyramid Lake. The test was designed to determine the effect of alkalinity on CTM values. In both strains it was found that as alkalinity increased from 69 to 1,487 mg/ L, the CTM decreased. Average CTM values determined for the death ( D) temperature endpoint were approximately 72.3 67.2 ° F ( 22.4 19.6 ° C) over this alkalinity range ( average values for the two strains). This CTM study employed a stepped temperature increase equal to 1.8 ° F ( 1 ° C)/ d up to an exposure temperature of 68 ° F ( 20 ° C), starting from an acclimation temperature of 60.8 ° F ( 16 ° C). After 68 ° F ( 20 ° C) was reached, the increments were 1.8 ° F ( 1 ° C)/ 4d. Because of the stepped increases and the two different rates of heating, the study methodology is not completely analogous to conventional CTM technique. The initial period of increase to 68 ° F ( 20 ° C), however, could be considered to provide nearly full acclimation ( given a 4 d acclimation at each step) before the subsequent heating schedule. The temperature increase rates for the two exposure periods averaged approximately 0.04 ° C/ h and 0.01 ° C/ h, respectively ( averaging the thermal increase over the step time interval). CTM test s provide results that have a different meaning from UILT test results. Comparison of CTM values for other salmonids that have corresponding UILT values is useful to understand the relative thermal tolerance of Lahontan cutt hroat . For example, appropriate comparisons of CTM values among species can be made by contrasting the Lahontan results with CTM values for salmonids whose temperature increase rates are 0.018 0.14 ° F ( 0.01 0.08 ° C)/ h and starting from acclimation temperatures of 60.8 68 ° F ( 16 20 ° C) ( McCullough 1999). Grande and Anderson ( 1991) measured a CTM of 79.3 ° F ( 26.3 ° C) for 2 to 3 month old rainbow trout, 81 ° F ( 27.2 ° C) for 3 to 4 month old brook trout, and 78.6 ° F ( 25.9 ° C) for 2 t o 4 month old lake trout. Elliott and Elliott ( 1995) reported CTM of 81.9 ° F ( 27.74 ° C) for Atlantic salmon and 76.6 ° F ( 24.8 ° C) for brown trout. The Lahontan cutthroat studies reported CTM for a death endpoint; the Grande and Anderson studies also used a death endpoint. If a loss of equilibrium ( LE) endpoint is used to record CTM, the crit ical temperature is generally slightly lower than if a death endpoint is used. For example, at a heating rate of 1.8 ° F ( 1 ° C)/ h, Becker and Genoway ( 1979) measured a CTM for coho salmon as 81.8 and 81.7 ° F ( 27.7 and 27.6 ° C) for the LE and death temperature endpoints, respectively. However, at a 64.4 ° F ( 18 ° C)/ h heating rate, these values were 83.6 and 85.4 ° F ( 28.7 and 29.7 ° C), respectively. In conclusion, the CTM values for Lahontan cutthroat trout, compared with other salmonids tested in a similar manner ( i. e., rainbow trout, brook trout, lake trout, Atlantic salmon, brown trout), are 93 Summary of Technical Literature Examining the Physiological Effects of Temperature much lower. UILT values determined in other studies ( see Table 4) for all the species that were contrasted above with Lahontan ranged from 73.4 to 79.5 ° F ( 23 26.4 ° C). Using CTM data as a guide, these results appear t o indicate that Lahontan cutthroat would likely have lower UILT values than these other salmonids.

Dickerson and coworkers ( 1999, and unpublished data, in Dunham 1999) found in laboratory tests at constant temperatures that survival was 100% at 75.2 ° F ( 24 ° C) but dropped to 35% at 78.8 ° F ( 26 ° C). Tests done under a fluctuating temperature regime of 68 78.8 ° F ( 20 26 ° C) with a daily mean of 73.4 ° F ( 23 ° C) for 1 wk showed no mortality even though this regime provided 1 h/ d exposure to 78.8 ° F ( 26 ° C) for 7 consecutive days, a temperature that produced mortality during longer exposures ( Dickerson and Vinyard 1999). These data indicate that the UILT is probably between 77 and 78.8 ° F ( 25 and 26 ° C). However, temperature adjustment rates, starting from an initial temperature of 55.4 ° F ( 13 ° C), were 7.2 ° F/ d ( 4 ° C/ d) up to the test temperature. That is, a conventional time period for acclimation was not allowed prior to the final exposure temperature. This could result in a slight underprediction of UUILT.

What are the therm al requirem ents for w estslope cu tthroat trout?

Incubation and egg survival. In a study by Shepard et al. ( 1982), westslope cutthroat trout in the Flathead River basin, Montana, emerged in July and August following incubation temperatures ranging from 35.6 to 50 ° F ( 2 10 ° C). Fry were approximately 20 mm long at emergence. Adult westslope cutthroat trout held in cool 35.6 39.2 ° F ( 2 4 ° C) water temperatures produced more viable eggs than those held in constant water temperatures of 50 ° F ( 10 ° C) ( Smith et al. 1983, in Shepard et al. 1982).

Growth. Westslope cutthroat streams are typically cold, nutrient poor waters in which conditions for growth tend to be less than optimal ( Liknes and Graham 1988).

Spawning. Initiation and timing of spawning activity is related to water temperatures. Adults move into tributaries during high streamflows and spawn in the spring when water temperatures are near 50 ° F ( 10 ° C) ( Scott and Crossman 1973).

Occurrence. Westslope cutthroat trout and bull trout have similar life history patterns, often occupy the same headwater streams, and restrict themselves to the coldest sections of streams ( Jakober et al. 1998, Behnke 1992). Westslope trout prefer cooler water temperatures than do both brook trout and Yellowstone cutthroat trout ( B. Shepard, personal communication).

Westslope cutthroat trout and redband trout may occur in the same system. They can be allopatric or sympatric, but the redband generally inhabit lower reaches and cutthroat trout ( often with bull trout) dominate the upper, higher gradient sections where annual temperature units are considerably less ( Mullan et al. 1992).

What are the therm al requirem ents for redband trou t?

Growth and feeding. Dwyer et al. ( 1986) conducted temperature experiments at 39.2, 44.6, 50, 60.8, and 66.2 ° F ( 4, 7, 10, 13, 16, and 19 ° C) on rainbow trout and redband trout 94 Summary of Technical Literature Examining the Physiological Effects of Temperature collected from Threemile Creek, Catlow Basin. Redband trout from Threemile Creek showed the best growth at 66.2 ° ( 19 ° C) ( no higher temperatures were tested), whereas rainbow trout exhibited the best growth at 59 60.8 ° F ( 15 16 ° C).

In a study by Sonski ( 1982, as cited by Sonski 1983b), redband trout reached their maximum growth rate at 68 ° F ( 20 ° C). Growth rates were less at both 59 ° F ( 15 ° C) and 73 ° F ( 22.8 ° C).

Redband trout are thought to exhibit the upper limit for feeding response for all salmonids of the Pacific Northwest. No feeding was observed by Sonski ( 1982 as cited by Sonski 1984) for juvenile redband at temperatures above 77.9 80.6 ° F ( 25.5 27 ° C). In a comparison of thermal tolerance by three rainbow trout species, Sonski ( 1984) found that no redband trout or Wytheville rainbow fed at temperatures higher than 78.8 ° F ( 26 ° C), and the Firehole River rainbow would not feed beyond 80 ° F ( 26.7 ° C).

Metabolic activity and swimming speed. Gamperl ( in litt.) conducted temperature studies in Bridge Creek (" warm" stream) and Little Blitzen River (" cold" stream) in the Harney basin. Gamperl's studies found that despite the two streams' different thermal histories, redband trout from each stream exhibited a similar preferred temperature of 55 ° F ( 12.8 ° C). Bridge Creek trout had greater metabolic power and improved swimming efficiency at 75.2 ° F ( 24 ° C) than at 53.6 57.2 ° F ( 12 14 ° C) compared with the Little Blitzen River redband, which had similar values for metabolic power and swimming performance at 53.6 57.2 ° F ( 12 14 ° C) and 75.2 ° F ( 24 ° C). Gamperl concluded that some populations of redband trout can tolerate, and may have adapted to, warm environmental temperatures. However, these studies should be taken as preliminary because they were not replicated.

Occurrence. There are observations of redband trout feeding and surviving at relatively high temperatures for a salmonid ( 82.4 ° F [ 28 ° C], Behnke 1992; 81.3 ° F [ 27.4 ° C] Sonski 1986; 80.6 ° F [ 27 ° C] Bowers et al. 1979), although it is unclear whether temperatures were measured in the vicinity of the stream that the fish actually inhabited. These trout may rely on microhabitats or thermal refuges to maintain populations in desert environments ( see Ebersole et al. in press).

Lethal effects. In a comparison of thermal resistance among redband trout, Firehole River rainbow, and Wytheville rainbow, Sonski ( 1984) found very little difference. He measured UILT values for subyearling trout acclimated at 73.4 ° F ( 23 ° C) of 79.1, 79.3, and 80.6 ° F ( 26.2, 26.3, and 27.0 ° C), respectively. These values are probably equivalent to UUILT values because it appears that resistance was not improved by acclimation beyond 59 ° F ( 15 ° C). It is interesting that redband trout were not significantly different in their thermal tolerance from other rainbow stocks, despite their reputation as being tolerant of higher water temperatures. 95 Summary of Technical Literature Examining the Physiological Effects of Temperature LITERATURE CITED Adams BL, Zaugg WS, McLain LR. 1975. Inhibition of salt water survival and Na K ATPase elevation in steelhead trout ( Salmo gairdneri) by moderate water temperatures. Trans Am Fish Soc 104( 4): 766 769.

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