REFERENCE COPY 

Do Not Remove from the Library 

U. S. Fish and W~lril~fo $ryicz . . 

Biological Report 82 (I 1.60) National Wetlands Research Center TR EL-82-4 

June 1986 700 Cajun Dome Boulevard 

Lo fayette, Louisiana 70506 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (Pacific Southwest) 

STEELHEAD 

Fish and Wildlife Service 

Coastal Ecology Group 

Waterways Ex~eriment Station 

U.S. Department of the Interior U.S. Army Corps of Engineers 
Biological Report 82(11.60) 

TR EL-82-4 

June 1986 

Species Profiles: Life Histories and Environmental Requirements 

of Coastal Fishes and Invertebrates (Pacific Southwest 

STEELHEAD 

Roger A. Barnhart 

California Cooperative Fishery Research Unit 

Humboldt State University 

Arcata, CA 95521 

Project Officer 

John Parsons 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

1010 Gause Boulevard 

Slidell, LA 70458 

Performed for 

Coastal Ecology Group 

Waterways Experiment Station 

U.S. Army Corps of Engineers 

Vicksburg, MS 39180 

and 

National Coastal Ecosystems Team 

Research and Development 

Fish and Wildlife Service 

U.S. Department of the Interior 

Washington, DC 20240 
This series should be referenced as follows: 

U.S. Fish and Wildlife Service. 1983-19-. Species profiles: li,fe histories 

and environmental requirements of coastal fishes and invertebrates. U.S. Fish 

Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4. 

This profile should be cited as follows: 

Barnhart, R.A. 1986. Species profiles: 1 ife histories and environmental 

requirements of coastal fishes and invertebrates (Pacific Southwest -- 

steelhead. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.60). U.S. Army Corps of 

Engineers, TR EL-82-4. 21 pp. 
PREFACE 

This species profile is one of a series on coastal aquatic organisms, 

principal ly fish, of sport, commercial , or ecological importance. The profiles 

are designed to provide coastal managers, engineers, and biologists with a brief 

comprehensive sketch of the biological characteristics and environmental 

requirements of the species and to describe how populations of the species may be 

expected to react to environmental changes caused by coastal development. Each 

profile has sections on taxonomy, 1 ife history, ecological role, environmental 

requirements, and economic importance, if appl icabl e. A three-ri ng binder is 

used for this series so that new profiles can be added as they are prepared. 

This project is jointly planned and financed by the U. S. Army Corps of Engineers 

and the U.S. Fish and Wildlife Service. 

Suggestions or questions regarding this report should be directed to one of 

the foll owing addresses. 

Information Transfer Specialist 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

NASA-Slidell Computer Complex 

1010 Gause Boulevard 

Slidell, LA 70458 

U. S. Army Engineer Waterways Experiment Station 

Attention: WESER-C 

Post Office Box 631 

Vicksburg, MS 39180 
CONVERSION TABLE 

Metric to U.S. Customary 

Mu1 tiply 

mi 11 imeters (mn) 

centimeters (cm) 

meters (m) 

ki 1 ometers ( km) 

2 

square meters (m ) 10.76 

square ki 1 meters ( km2) 0.3861 

hectares (ha) 2.471 

liters (1) 

cubic meters (m3) 

cubic rneters 

To Obtain 

inches 

inches 

feet 

mil es 

square feet 

square (nil es 

acres 

gal 1 ons 

cubic feet 

acre-feet 

mil 1 igra~ns (mg) 0.00003527 ounces 

gram (g) 0.03527 ounces 

kil ogra~ns (kg) 2.205 pounds 

metric tons ( t) 2205.0 pounds 

~netric tons 1.102 short tons 

ki 1 ocal ories ( kcal ) 3.968 British thennal units 

Cel sius degrees 1.8("C) + 32 Fahrenheit degrees 

U.S. Customary to Metric 

inches 25.40 

inches 2.54 

feet (f t) 0.3048 

fathoms 1.829 

miles (mi) 1.609 

nautical miles (mi) 1.852 

square feet (ft2) 

acres 2 

square miles (mi ) 

gal 1 ons ( gal ) 

cubic feet (ft3) 

acre- feet 

ounces (oz) 28.35 

pounds (lb) 0.4536 

short tons (ton) 0.9072 

British thermal units (Btu) 0.2520 

mil 1 imeters 

centimeters 

meters 

meters 

kilometers 

ki 1 orneters 

square meters 

hectares 

square kilometers 

1 i ters 

cubic meters 

cubic meters 

grains 

ki 1 og rarns 

metric tons 

ki 1 ocal ori es 

Fahrenheit degrees 0.5556("F - 32) Cel sius degrees 
CONTENTS 

Page 

PREFACE ...................................................................... iii 

CONVERSION TABLE .............................................................. iv 

ACKNOWLEDGMENTS ............................................................... vi 

NOMENCLATURE/TAXONOMY/RANGE .................................................. 1 

MORpHOLOGY/IDENTIFICATION AID'S ............................................... 1 

REASON FOR INCLUSION IN SERIES ............................................... 4 

LIFE HISTORY ................................................................. 4 

THE FISHERY .................................................................. 6 

ECOLOGICAL ROLE .............................................................. 8 

ENVIRONMENTAL REQUIREMENTS ................................................... 9 

Temperature ................................................................. 9 

Dissolved Oxygen ........................................................... 10 

Depth ...................................................................... 11 

Water Movement ............................................................. 11 

Sediment ................................................................... 12 

LITERATURE CITED ............................................................ 15 
ACKNOWLEDGMENTS 

I thank Dennis P. Lee, California Department of Fish and Game, and Terry D. 

Roelofs, Humboldt State University, for reviewing the manuscript; Thomas H. 

Hassler, California Cooperative Fishery Research Unit,for acting as the liaison 

with the National Coastal Ecosystems Team; and Delores Neher for greatly 

facilitating the preparation of this report. 
Figure 1. Steelhead. 

STE ELHEAD 

Scientific name ........ Salmo 

gairdneri Richardson 

Preferred common name ... Steelhead 

(Figure 1) 

Other common names .... steelhead 

trout, steelie, half-pounder, ironhead 

Class .......... Osteichthyes 

Order ......... Salmoniformes 

Family .......... Salmonidae 

Geographic range: At sea, from 

northern Baja California to the 

Bering Sea and Japan. In the 

Pacific Southwest, steelhead enter 

coastal streams from the northern 

California border south to the 

Ventura River (Figure 2). Adult 

steelhead have been reported in the 

Santa Clara and Santa Margarita 

Rivers in years when winter runoff 

was high enough to allow upstream 

migration (Swift 1975). Early in 

this century steelhead distribution 

in the Pacific Southwest was more 

widespread (Figure 3). Steel head 

run up the Santa Domingo River in 

northern Baja Cal ifornia during high 

runoff in winter (Needham and Gard 

1959 1. 

MORPHOLOGY/IDENTIFICATION AIDS 

The fol lowing descriptions were 

taken from McConnell and Snyder (1972) 

and Fry (1973). 

At sea, steelhead are steel blue 

above and bright silver on the sides 

and belly. Sharply defined black 

spots are located on back, head, 

sides, and dorsal and caudal fins. 

The spots are small and vary in 

number. After entering freshwater, 

steelhead gradually take on the 
I 

I 

124O I 

I zoo 

OREGON 

CALIFORNIA 

0 SO lOOmi 

0 100 200km 

- 330 & Terminal Dam 

l IS0 -- - - -- ~~-p 

PACIFIC 

OCEAN 

Figure 2. Distribution of steelhead (shaded areas) in Pacific Southwest region 

streams in 1980's (California Department of Fish and Game, pers. comm. ; Moore 

1980). 
CALIFORNIA 

0 50 l0Om1 

Figure 3.Distribution of steelhead. (shaded areas) in the Pacific Southwest 

region in 1900 (National Council on Gene Resources 1982). 
appearance of stream rainbow trout; 

the back becomes olive green and the 

sides and belly less silvery. 

Maturing adults usually develop a 

broad pink stripe along the lateral 

line and pink coloration on the 

operculum. Steel head lack the red 

streaks beneath the jaw which 

characterize the cutthroat trout 

(Salmo -- clarki). Juvenile steel head in 

streams cannot be di sti naui shed from 

juvenile resident rainlbow trout. 

Anal rays 9-12, rarely 13; teeth 

on tip and shaft of vomer and on 

tongue; gill rakers on lower limb of 

first arch range from 17-21; scales in 

first row above lateral line 115-180; 

caudal fin shallowly forked; maxi1 lary 

does not reach past the posterior 

margin of eye; juvenile parr marks 

small and oval to nearly round. 

Generally adult steelhead in the 

Pacific Southwest region weigh less 

than 4.5 kg but some exceed 11 kg. 

REASON FOR INCLUSION IN SERIES 

The steelhead is the most 

widespread anadromous sport fish in 

the Pacific Southwest region, but its 

abundance has declined since the 

1950's because of increased sport 

fishing intensity and damage to fish 

habitat through construction of dams, 

water diversion, road construction, 

and improper 1 and management 

practices. In some rivers a steelhead 

fishery would be nonexistent without 

hatchery stocks. 

In Cal ifornia, attempts are 

being made to protect wild stocks of 

steelhead to maintain existing 

spawning and rearing habitat, to 

restore or enhance degraded habitat, 

to use artificial propagation wisely, 

and to regulate the fishery to provide 

qua1 ity angling (California Department 

of Fish and Game 1975). 

LIFE HISTORY 

Steelhead trout spend a portion 

of their life in the ocean where most 

of their growth occurs and sexual 

maturity is attained; then they enter 

freshwater to spawn. Spawning generally takes place from February to late 

June. The eggs are laid in pits 

(redds), dug in the gravel of the 

stream bottom by the female. Immediately after the eggs are laid and 

fertilized, they are covered with 

gravel by the female; the length of 

their stay in the gravel depends upon 

water temperature, dissolved oxygen 

concentration and substrate composition. After the eggs hatch, the young 

steelhead gradually work their way to 

the surface of the stream bed. Juveniles usually spend a year or longer 

in freshwater (length of residence is 

determined by environmental and 

genetic factors) and then descend to 

the ocean. 

The steelhead is a strain of 

rainbow trout that has a strong urge 

to migrate to the ocean; however, some 

individuals may remain in a stream, 

mature, and even spawn without going 

to sea (Shapovalov and Taft 1954). 

The life history of the steelhead trout varies more than that of 

any other anadromous fish regarding 

the length of time spent at sea, the 

length of time spent in freshwater, 

and the times of emigration from and 

immigration to freshwater. Unlike 

salmon, steelhead do not usually die 

just after spawning. 

Steelhead are classified into 

two races (Withler 1966; Smith 1960, 

1969; and Everest 1973): winter 

steelhead that enter streams between 

November 1 and April 30, and sumner 

steelhead that enter streams between 

May 1 and October 30. Portions of 

both groups may enter freshwater in 

spring or fa1 1 and are then called 

spring- or fall-run steelhead. In 

large rivers, such as the Klamath and 

Sacramento Rivers, steelhead may enter 
during most of the year. Winter-run 

steelhead usual ly enter freshwater as 

maturing fish that spawn relatively 

soon, whereas most summer steelhead 

enter as immature fish and do not 

mature and spawn until several months 

later. In the Pacific Southwest, 

sumner steelhead are not abundant and 

the runs in many streams consist of 

less than 100 fish (Roelofs 1983). 

Degradation of habitat in intermi ttent streams and suscepti bi 1 ity to 

angling and predation probably account 

for the low numbers. The southernmost 

summer steelhead population i s i n the 

Middle Fork Eel River. This river 

supports the largest run of these fish 

in the Pacific Southwest--up to 2,000 

fish in a good year. 

An exception is the "halfpounder" sumner steelhead (terminology 

of Snyder 1925) which has a unique 

1 ife hi story described by Kesner and 

Barnhart (1972) and Everest (1973). 

Half-pounder runs are confined to a 

small geographic range encompassing 

about 120 miles of the southern Oregon 

and northern Cal i fornia coasts, 

including the Rogue, Klamath, Mad and 

Eel Rivers. These small immature 

steelhead (25-35 cm long) annually 

enter freshwater from late August to 

early October, and are the basis of 

important sport fisheries in the 

Klamath and Rogue systems. Halfpounders spend only a few months at 

sea before they return to freshwater 

and, in contrast to mature steelhead, 

feed extensively in freshwater. Halfpounders that survive their first 

upstream migration return to the ocean 

the following spring and migrate back 

to freshwater as mature steelhead in 

the sumner and fall. Everest (1973) 

reported that half-pounders annually 

comprise about 65% of the summer-run 

of steelhead on the Rogue River and 

that 97% of all adult summer steelhead 

from there make their first upstream 

migration as half-pounders. 

Sumner and winter steelhead do 

not interbreed; they are isolated 

temporally and spatially (Smith 1969; 

Everest 1973 1. Sumner steel head spawn 

in January and February, whereas 

winter steelhead spawn in April and 

May, and summer steel head spawn in 

smaller streams or farther upstream. 

The sex ratio of steelhead trout 

immigrants is about 1:l (Shapovalov 

and Taft 1954; Kesner and Barnhart 

1972). Female steelhead contain 

about 2,000 eggs per kilogram of body 

weight (Moyle 1976). 

Several researchers have 

concluded that the incidence of steelhead spawning more than once increases 

from north to south (Bali 1959; 

Withler 1966; Sheppard 1972). Wide 

variations in the percent of repeat 

spawning can be due to genetic 

factors, habitat qua1 ity, fishing 

intensity, and management practices. 

Fish that have spawned twice make up 

70% to 85% of repeat spawners, whereas 

those that have spawned three times 

make up 10% to 25% of a1 1 repeat 

spawners (Forsgren 1979). The few 

fish that have spawned four times are 

likely to be females, which have a 

higher survival rate than males during 

and fol lowing spawning. Spawning 

males usually each serve more than one 

female, remain in the stream longer 

than the females (tagging studies by 

Jones (1974) indicated nearly two 

weeks longer), and are exposed to more 

prolonged physical exertion than the 

females (Meigs and Pautzke 1941). 

Steelhead spawn in cool, clear, 

well-oxygenated streams with suitable 

depth, current velocity, and gravel 

size (Reiser and Bjornn 1979). Measurements made over steelhead redds 

showed that steelhead spawn at depths 

of 0.10-1.5 m, current velocities of 

23-155 cm/sec, and in gravel of 0.64- 

12.70 cm in diameter (Smith 1973; 

Hunter 1973; Bovee 1978; Wesche and 

Rechard 1980). Intermittent streams 

are often used by steelhead for 

spawning (Everest 1973; Kralik and 

Sowerwine 1977; Carroll 1984). Most 

of the fry produced emigrate to perennial streams soon after hatching. 
The embryology of the steelhead 

is similar to that of other salmonids 

(Wales 1941). The number of days 

required for steelhead eggs to hatch 

varies from about 19 at an average 

water temperature of 15 OC to about 80 

days at an average of 5 OC. Steelhead 

fry usually emerge from the gravel 2 

to 3 weeks after hatching. 

After emergence, steel head fry 

(often in small schools) usually live 

in shallow water along the stream 

banks. As the fry grow older the 

schools break up and the individual 

fish establish territories which they 

defend. Most steelhead in their first 

year of life tend to inhabit riffles 

but some of the larger fish inhabit 

pools or deeper faster runs. Mortality is high during the first few 

months after emergence and many 

investigators have suggested that the 

relative size of the year class is 

largely determined at that time 

(Chapman 1966; McFadden 1969; Burns 

1971; Everest and Chapman 1972). 

In recent years, habitat degradation has lowered the capacity of 

many ftreams to rear steelhead to 

smolts . For example, excessive sedimentation has reduced food production, 

pool depth, and cover--all important 

to juvenile steelhead survival. 

Juvenile steelhead feed on a 

wide variety of aquatic and 

terrestri a1 insects. Newly emerged 

fry sometimes are preyed upon by older 

juvenile steelhead. Young steelhead 

moving about trying to find a suitable 

territory are subject to the highest 

predation (Shapovalov and Taft 1954; 

Chapman 1966 1. 

Juvenile steelhead live in 

freshwater for from 1 to 4 years 

Ismolt: Term applied to an anadromous 

juvenile salmonid that is physiologically prepared to adapt to a saltwater 

existence; steelhead smolts lose their 

parr marks and become silvery. 

(usually 2 years in the Pacific 

Southwest) before becoming smolts and 

migrating to saltwater. Most of the 

migrants are 2 years old and 18.6 to 

21 cm long (fork length) in the 

Kl amath River (Kesner and Barnhart 

19721, or 14 to 21 cm long in Waddell 

Creek (Shapovalov and Taft 1954). The 

larger the smolts, the better the 

survival. Most steelhead smolts enter 

the sea in March and April. 

Steelhead live 1 to 4 years in 

the ocean (usually 1 or 2 years in the 

Pacific Southwest). The length of 

residence in both freshwater and saltwater increases from south to north 

(Withler 1966). Steel head grow 

rapidly in the ocean and their size 

upon reaching maturity depends primari ly on how long they have 1 ived in 

the ocean. In California, the average 

length of adult steelhead after 2 

years in the ocean is 58 cm (Withler 

1966). Immature steelhead trout (halfpounders) increase about 30 mm in 

length each month from the time they 

enter the ocean until they return to 

freshwater (Kesner and Barnhart 1972). 

The distribution or migration of 

steelhead in the ocean is not well 

defined, particularly in the Pacific 

Southwest. As judged by tag returns, 

most steelhead tend to migrate north 

and south along the Continental Shelf. 

Steelhead stocks from the Klamath and 

Rogue Rivers probably mix together in 

a nearshore ocean staging area along 

the northern California coast before 

they migrate upriver (Everest 1973). 

THE FISHERY 

Commercial fishing for steelhead 

has been prohibited in the Pacific 

Southwest since 1924. The species now 

is managed exclusively as a sport 

fishery. Most steelhead are caught in 

rivers rather than in the ocean. The 

three most important steelhead trout 

rivers are the Klamath and Eel Rivers 

in northern California and the 

Sacramento River which empties into 
San Francisco Bay. The California 

Department of Fish and Game (1965) in 

its Fish and Wildlife Plan estimated 

spawning runs of 221,000 steel head in 

the Klamath River and 82,000 steelhead 

in the Eel River. The Klamath River 

still has good runs of native 

steel head, perhaps approaching 200,000 

fish in good years (Barnhart 1975). 

In the upper Sacramento River from 

1953 to 1959, the average run of adult 

steelhead was 20,590 and consisted of 

both natural and hatchery-produced 

fish (Hallock et al. 1961). 

In the California Fish and Wildlife Plan (19651, the State's annual 

spawning stock of steel head was estimated at 603,000 fish. The Plan also 

reported that the State contained 

8,402 linear miles of steelhead habitat, 31% of which was available to 

anglers. It was estimated that steelhead anglers fish 304,000 angler-days 

per year to harvest 122,000 fish 

(about 0.4 fish per angler-day). Many 

Cal ifornia streams support a substantial fishery for juvenile steelhead-- 

fish that have not yet migrated to the 

ocean. These 5- to 8-inch trout are 

taken in large numbers, mostly during 

the so-called summer trout season. 

The fishing pressure for these trout 

(about 440,000 angl er-days annual ly) 

exceeds that on adult steelhead (California Department of Fish and Game 

1965 1. 

A1 though adult steelhead do not 

commonly feed in freshwater, they are 

readily taken by angling. Bait fishing is popular and effective. Salmon 

eggs or clusters of roe or night 

crawlers are drifted through riffles, 

pools, and especially where a tributary enters a large stream. Spinners 

and other artificial lures are also 

used. When streams are low and clear, 

artificial flies are sometimes effective. Most steelhead fishing is done 

from the bank or by wading; drifting 

in boats is popular in larger streams. 

The decl ine of wi ld, natural ly 

produced steelhead in the spawning 

runs in the Pacific Southwest region 

since the 1940's has required an increase in the stocking of hatchery 

reared steelhead. A total of about 1.9 

mi 1 lion yearl ing steelhead for planting are supplied annually by Coleman 

National Fish Hatchery on Battle 

Creek, a tributary to the upper Sacramento River, the Nimbus Fish Hatchery 

on the American River, the Feather 

River Hatchery on the Feather River, 

and the Mokelumne River Hatchery on 

the Mokelumne River. The goal is to 

mitigate the loss of steelhead trout 

in those rivers that has been caused 

by dams and water diversion. 

The average weight of hatchery 

yearlings at the time of planting 

(January through May) is about 51 

grams (9 fish per pound). At an 

expected return of about 2% annually, 

planted steel head contributed about 

38,000 adults to the run in the 

Sacramento River (Hallock et al. 

1961). If steelhead were not stocked, 

the catch in the Sacramento River 

would be much smaller. The California 

Department of Fish and Game operates 

four hatcheries on the coast that 

produce about 1.5 million yearling 

steel head annual ly. In addition, 

trout rearing projects sponsored by 

the Cal ifornia Department of Fish and 

Game, private groups, and counties 

contribute an average of 114,000 

yearl ings annual ly (Boydstun 1977a). 

On the Klamath system, where two of 

the coast hatcheries are located, 

hatchery production contributes only 

an estimated 8% of the run of adults 

(Boydstun 1977b). Cal ifornia also 

releases nearly 3 mi 11 ion steel head 

f ingerl ings annually (Jensen 1971). 

Fishery regulations are used to 

help protect decl ining steel head 

stocks from excessive fishing. In 

California, the daily bag limit is 

three adult fish. All streams tributary to major coastal rivers and most 

small coastal streams are closed to 

angling from mid-November to late 

April. Special regulations also are 

sometimes used. For example, a 
section of the upper Middle Fork Eel 

River has been permanently closed to 

angling to protect the adult summer 

steelhead trout that concentrate there 

in summer and fall. 

The use of hatchery reared trout 

to boost populations is not entirely 

beneficial . Larger runs attract more 

fishermen, and more fishermen further 

reduce the abundance of wild stocks. 

Also, the genetic mixing of hatchery 

and wild stocks could decrease the 

advantageous traits in wild stocks. 

The steelhead trout policy of the 

California Department of Fish and Game 

1 imits artificial propagation and 

stocking to reduce such interference 

with natural salmonid stocks; such 

measures are periodically reviewed to 

assess their effects on the wild 

stocks. 

The environmental qua1 ity of 

coastal rivers 1 argely governs the 

level of steelhead production. In the 

Pacific Southwest, attempts have been 

made to increase the protection of 

existing spawning and rearing habitat 

and to rehabi 1 i tate damaged habitat 

where feasible. California's steelhead trout policy is directed toward 

three goals: (1) to protect and 

improve steelhead trout habitat; (2) 

to develop plans and programs for 

assessment of habitat conditions and 

adverse impacts, land use planning, 

and acquisition of interests in 

streams threatened with adverse developments; and (3) to study the effects 

of habitat changes caused by overgrazing, gravel extraction, logging, 

road construction, urbanization, and 

water development. 

In 1982-84, Cal ifornia spent 

$900,000 annual ly on the restorat ion 

of salmon and steelhead habitat. A 

Salmon Restoration Project was developed within the California Conservation Corps in 1980, and by the end of 

1983, Corps enrol 1 ees removed 7,447 

cords of wood and debris from 233 

miles of streams used by anadromous 

fish (Kreb 1984). The Conservation 

Corps also planted streams ide vegetation and constructed fish passage 

weirs on some streams. In 1981-1983, 

an additional $1 million was spent 

annually by California to enhance 

spawning gravels on the upper Sacramento River and the Shasta and Klamath 

Rivers, and to construct salmonid 

rearing ponds (Rawstron and Hashagen 

1984). Six Rivers National Forest in 

northern Cal iforni a during the past 5 

years has carried out an extensive 

program to restore or enhance the 

spawning and rearing habitats of 

anadromous fish ( Overton et a1 . 1981; 

Overton 1984). 

ECOLOGICAL ROLE 

In freshwater, young steel head 

may be sympatric with sculpins, resident rainbow trout, coho salmon 

(Oncorhynchus kisutch), and in some 

instances other salmonids. Data are 

lacking on the effects of interaction 

of juveni le steel head and resident 

rainbow trout, largely due to the 

difficulty in distinguishing between 

the fish. In the Pacific Southwest, 

resident rainbow trout are common in 

streams above barriers to steelhead 

migration and fry sometimes drift 

downstream over the barriers. Bjornn 

(1978) reported that steel head fry 

tended to displace juvenile resident 

trout in the Lemhi River, Idaho. The 

ecological requirements of the two 

races are similar. 

Coho salmon and steelhead trout 

are simi 1 ar in geographical distribution, systematic characteristics, 

spawning locations, food habits, and 

the length of time the young mend in 

freshwater (Mi 1 ne 1948), a1 though 

steel head normal ly remain in 

freshwater longer than 1 year. 

Although both species live in similar 

habitat in their first year, Hartman 

(1965) reported that in spring and 

summer, most steelhead trout live in 

riffles and most coho salmon live in 

pools. Several investigators have 

reported that spati a1 segregation is 
also vertically stratified; coho 

salmon live near the stream surface 

and steelhead near the bottom (Hartman 

1965; Peterson 1966; Edmundson et al. 

1968; Bustard and Narver 1975). 

Although coho salmon hatch earlier and 

consequently are larger at first, 

steelhead fry grow so much faster that 

by late summer the size difference is 

small. Interspecific competition is 

probably not serious because of the 

initial difference in size, differences in habitat preference, and the 

difference in age at first emigration 

to the sea (Fraser 1969). Much the 

same re1 ation was reported for juveni le chinook salmon (Oncorhynchus 

tshawytscha) and juvenile steel head by 

Chapman and Bjornn (1969). 

Native and hatchery-reared 

steelhead exhibit some competition. 

Steelhead trout planted at the wrong 

time or at the wrong size tend to stay 

in the stream longer than usual and 

are more competitive with wild fish 

(Royal 1972). Heavy predation by 

hatchery-raised steelhead smolts on 

chinook salmon fry below Coleman 

National Fish Hatchery on the 

Sacramento River was reported by 

Menchen (1981 1. Wagner (1967) stated 

that ideally the stream is to serve 

only as a highway to the sea and not 

as a postliberation rearing area for 

hatchery products. Pollard and Bjornn 

(1973) reported that the stocking of 

hatchery rainbow trout also caused a 

local i zed temporary decrease in the 

abundance of juvenile steel head. 

In coastal streams, steelhead 

fry are preyed on by sculpins (Cottus 

spp. ), larger steelhead, and rainbow 

trout (Shapovalov and Taft 1954); by 

birds such as the great blue heron 

(Ardea herodi as), be1 ted kingfisher 

(Cer le alcyon), American dipper 

(&s mex~canus), and common 

mergansers 

Taft 1954; Sheppard 1972; Cross 1975). 

In the ocean, steelhead are eaten by 

fish and marine mammals but the extent 

and effect of predation are unknown. 

In freshwater, steelhead feed 

primarily on immature aquatic stages 

of insects and secondarily on mature 

terrestrial insects. Ephemeroptera 

(mayflies), Diptera (true flies), and 

Trichoptera (caddisf 1 ies) are usual ly 

the most important taxa in the diet 

(Shapovalov and Taft 1954; Royal 1972; 

Fite 1973; Hiss 1984). Juven i 1 e 

steelhead are somewhat opportunistic, 

feeding on almost any available insect 

(Fite 1973). The size of the 

territory for a single juvenile is 

determined 1 argely by the avai labi 1 i ty 

of food and the size of the fish 

(Allen 1969). In the ocean, steelhead 

feed on a variety of organisms 

including juvenile greenlings 

(Hexa rammos spp. 1, squids, and amphipods -?- LeBrasseur 1966; Manzer 1968). 

ENVIRONMENTAL REQUIREMENTS 

Temperature 

Water temperature affects all 

metabolic and reproductive activities 

of fish, including such critical 

functions as growth, swimming, and the 

ability to capture and assimilate food 

(Tebo 1974). Optimum temperature 

requirements may vary, depending on 

the season and life stage. A 

productive steel head stream should 

have summer temperatures in the range 

of 10 to 15 OC and an upper limit of 

20 OC. Steelhead have difficulty 

extracting oxygen from water at 

temperatures much over 21 OC 

regardless of the amount of oxygen 

present (Hooper 1973). Be1 1 (1973) 

listed the preferred temperatures of 

young steelhead as 7.2 to 14.5 OC, the 

optimum as about 10 OC, and the upper 

lethal limit as 23.9 OC. Bov ee 

(19781, who developed a probabilityof-use temperature curve for rearing 

winter steelhead juveniles, wrote that 

optimum temperatures ranged from 0 to 

24 OC and peaked at 15 OC. In studies 

of the smolting of cultured steelhead 
in the spring, Wagner (1974) and 

Kerstetter and Keeler (1976) reported 

that low temperatures extend the 

smolting period and high temperatures 

shorten it. They reported that smolting ceased rather abruptly when water 

temperatures increased to 14-18 OC. 

During the spawning season a 

sudden drop in water temperature may 

cause all salmonid spawning activity 

to cease (Reiser and Bjornn 1979). 

Reingold (1968) reported dhat water 

temperatures of 2 to 10 C impaired 

the viability of eggs and delayed the 

ripening of adult steelhead held for 

51 days in a hatchery pond. Bovee 

(1978) reported a spawning temperature 

ranye for winter steelhead of 4 to 

13 C, and a peak of 8'~. Bell 

(1973) indicated that steel head 

spawning temperatures are generally 

from 3.9 to 9.4OC. The average 

development time from ferti 1 ization to 

hatching lengthens considerably with 

decreasing temperature; f8r rainbow 

trout it is 19 days at 15 C, 31 days 

at 10 OC, and 80 days at 5 OC (Embody 

1934 1. Bovee (1978) gave an 

incubat ion temperature range for 

winter steelhead of 0 to 24 OC and an 

optimum temperature of about 10 OC. 

When water temperatures fall 

below 4 OC in streams of the Pacific 

Northwest, juveni le steel head become 

inactive and hide in available cover 

or in the substrate (Chapman and 

Bjornn 1969; Bustard and Narver 1975). 

In California's coastal streams, 

juvenile steelhead remain active yearround and in one small coastal stream, 

young steelhead grew throughout the 

winter (Reeves 1979). 

The virulence of many fish 

diseases and the toxicity of most 

chemicals increase with increasing 

water temperatures (Lantz 1971 1. 

Removal of riparian vegetation can 

result in marked increases in sumner 

stream temperatures and sometimes in 

reduced winter temperatures (Brown 

1971 1. 

Dissolved Oxygen 

Stream-dwel 1 ing salmonids require 

high dissolved oxygen in both the 

water column and i ntragravel waters. 

The swimming performance of juveni 1 e 

and adult salmon was impaired when the 

dissolved oxygen concentration was 

reduced below the air-saturation level 

(Davis et al. 1963). A sharp decrease 

in performance was noted at 6.5-7.0 

mg/l. Reiser and Bjornn (1979) wrote 

that the oxygen levels recommended for 

spawning anadromous f i sh (at 1 east 80% 

of saturation, with temporary 1 evels 

no lower that 5.0 mg/l) should meet 

the needs of migrating salmonids. 

Intragravel dissolved oxygen 

concentration is positively related to 

survival of salmonid embryos (Coble 

1961; Phillips and Campbell 1962; 

Silver et al. 1963). Silver et al. 

(19631, in a controlled -laboratory 

experiment conducted at 9.5 OC, found 

that steelhead embryos hatched 

successfully at mean oxygen concentrations as low as 2.6 mg/l but that 

total mortality occurred at a mean 

level of 1.6 mg/l. In a field experiment Phillips and Campbell (1962) 

noted that total mortal i ty of 

steelhead embryos occurred at mean 

oxygen concentrations of 7.2 mg/l or 

less. Although intragravel oxygen 

levels may appear adequate, the amount 

of oxygen actually reaching the 

embryos also depends on the intragravel water velocity (Wickett 1954). 

The amount of intragravel oxygen 

available to developing embryos is 

sometimes reduced by the biochemical 

oxygen demand of organic material in 

the gravel bed (Ponce 1974). Even if 

embryos hatch at low or moderately 

reduced oxygen levels, the incubation 

period is extended and the resulting 

fry are likely to be smaller and 

weaker than those reared at oxygen 

concentrations close to saturation 

(Silver et al. 1963; Shurnway et al. 

1964). Reiser and Bjornn (1979) 

concluded that although dissolved 

oxygen concentrations required for 
successful incubation depend on both 

species and developmental stage, 

concentrations at or near saturation 

with no temporary reductions below 5.0 

mg/l are generally required by anadromous salmonids. 

In salmonid nursery and rearing 

streams dissolved oxygen concentrations of surface waters are normal ly 

near saturation, except in small 

tributaries with large amounts of 

debris from logging or other sources 

or in large, slow-moving streams 

receiving 1 arge amounts of municipal 

or industrial waste (Reiser and Bjornn 

1979). Salmonids function normally at 

dissolved oxygen concentrations of 

7.75 mg/l ; exhibit various distress 

symptoms at 6.00 mg/l; and are often 

negatively affected at 4.25 mg/l 

(Davis 1975). Low dissolved oxygen 

impairs metabol ic rate, growth, swimming performance, and overall survival 

of young salmonids. 

Water depth usually does not 

prevent migration because adult steelhead normally migrate when stream 

flows are relatively high. Thompson 

(1972) wrote that 18 cm is the minimum 

depth required for successful 

migration of adult steelhead. The 

migration of adult salmonids is more 

commonly hindered by excessive water 

velocity or obstacles that impede the 

swimming or jumping of the fish. 

Water depth may be important in 

the selection of redd sites. 

Shapovalov and Taft (1954) stated that 

steelhead redds are rarely exposed by 

fa1 1 ing stream levels. Bovee (1978) 

showed that steelhead most commonly 

spawn at depths averaging 36 cm 

(range, 15-61 cm). The depths of 

Washington winter steel head redds 

ranged from 12 to 70 cm (Hunter 1973). 

Carroll (1984) measured water depths 

of 22 to 29 cm over steelhead redds in 

a Klamath River tributary. 

Steelhead tend to occupy the 

shallow riffle areas of streams, 

particularly during the first year of 

1 ife (Hartman 1965). Bovee (1978) 

presented probability of use curves 

showing that steelhead fry are most 

commonly found in water 8 to 36 cm 

deep, and steelhead juveniles are 

usual ly located in water 25 to 50 cm 

deep. In the Southwest region steelhead streams are annually subjected to 

low flow conditions due to the 

extended summer-fall dry season; thus, 

pool frequency and depth are important. In a 2-year study at Singley 

Creek, a small coastal stream just 

south of Cape Mendocino, Cross (1975) 

found that 67%-96% of young-of-theyear steelhead resided in pools. From 

June to September, the riffle surface 

area was reduced 45% while the surface 

area of pools diminished only 26%. 

Excessive sediment inputs that f i 11 

pools can greatly reduce a stream's 

capacity to rear steelhead to smolt 

size. 

Water Movement 

Steel head may encounter water 

velocity barriers during upstream 

migration, often at falls or culverts. 

Velocities of 3-4 m/s begin to greatly 

hinder the swimming ability of 

steelhead and may retard migration 

(Rei ser and Bjornn 1979 1. Thompson 

(1972) out1 ined methods to calculate 

minimum and maximum acceptable streamflows for migrating adult steelhead, 

for specific stream sections. 

Bovee (1978) showed that steelhead spawn in areas with water 

velocities of 30-110 cm/s but that the 

preferred velocity approximated 

60 cm/s. Smith (1973) found that the 

preferred water velocity for spawning 

steelhead in Oregon ranged from 40 to 

91 cm/s. Steelhead redds were in 

areas where velocities were 15 to 54 

cm/s in a small tributary of the 

Klamath River (Carroll 1984). Since 

swimming performance improves with 

size, large adult steelhead can 
establish redds in faster current 

areas of the stream. 

Permeability is defined as the 

capacity of the gravel to transmit 

water. Different spawning gravels 

transmit water at different rates. 

Several studies have demonstrated that 

increased intragravel velocities 

improve the survival of steelhead eggs 

and fry before emergence and also the 

condition of fry that emerge (Shumway 

1960; Silver 1960; Coble 1961; Silver 

et a1 . 1963; Shumway et al. 1964). 

McNei 1 and Ahnel 1 (1964) concluded 

that the streambeds of highly productive spawning streams had gravels with 

high permeabi 1 i ty (24,000 cm/h) and 

had less than 5% (by volume) sand and 

silt that passed through a sieve of 

0.833 mm mesh. 

Sediment 

Quantitatively, sediment is the 

greatest single pollutant in the 

nation's water (Ritchie 1972). Anderson (1971) reported that sediment 

production in northern California 

watersheds increased markedly as a 

resu 1 t of poor 1 and management 

practices and f 1 oods. The steel head's 

environment can be impaired both by 

sediment in suspension and by particles deposited as bedload sediment. 

About 28% of the total spawning area 

in a once important 16-mile stretch of 

the upper Trinity River has been lost 

due to the deposition of sediment 

(California Resources Agency 1970). 

Stream channels of northern Cal iforni a 

markedly aggraded after 1 arge f 1 ood 

events during the 1960's and 1970's 

(Lisle 1982). Channel widening and 

loss of pool-riffle sequence due to 

aggradat ion damaged spawning and 

rearing habitat of steelhead. The 

pool -riffle sequence and pool qua1 i ty 

in some northern California streams 

sti 11 had not fully recovered by 1980 

from a 1964 regional flood (Lisle 

1982 ) . 

For rearing juvenile steel head, 

deposited sediment reduces the 

carrying capacity of the stream 

directly by reducing avai 1 able rearing 

habitat and indirectly by reducing the 

production of invertebrate food. 

Bjornn et al. (1977) found significant 

reductions in numbers of juvenile 

steelhead in stream channels where 

boulders were imbedded in sediment. 

Crouse et al. (1981), who devised a 

visual substrate scoring system based 

on particle size and degree of cobble 

embedment, reported that fish 

production was significantly 

correlated with substrate score. They 

reported significant decreases in fish 

production in streams where cobbles 

were embedded 80%-100% and where 

sediment (2.0 mm or less) composed 

26%-31% (by volume) of the total 

substrate composition. The authors 

concluded that rearing habitats of 

juvenile salmonids in streams, as well 

as spawni ng gravel s , require 

protection from excessive quantities 

of fine sediments. 

The size of substrate material 

has been related by numerous investigators to the standing crops of invertebrates (Sprules 1947; Kimble and 

Wesche 1975). Pennak and VanGerpen 

(1947) found that the number of 

benthic invertebrates decreased in the 

progression from rubble to bedrock to 

gravel to sand. Reiser and Bjornn 

(1979) reported that aquatic insect 

production was highest in substrate 

composed 1 argely of coarse gravel 

(3.2-6.7 cm) and rubble (7.6-30.4 cm). 

Suspended sediment occasionally 

reaches concentrations high enough to 

directly injure steelhead (3,000 ppm 

or greater) (Cordone and Kel ley 1961). 

Physiological damage includes the 

adhesion of silt particles to the 

chorion of salmonid ova (Cordone and 

Kelley 1961) and the abrasion, 

thickening, and fusion of gill 

f i 1 aments (Herbert and Merkens 1961 1. 

Sigler et al. (1984) reported that 

chronic turbidity in streams during 

emergence and rearing of steelhead 

affects the numbers and quality of 
fish produced. Turbid water also took place when turbidities had 

affects recreational angl i ng, for decreased to 30 Jackson Turbidity 

steelhead. A study on the Eel River Units or less; such low levels of 

by the California Department of Fish turbidity occurred during only 

and Game during two winter steelhead one-third of the fishing season (Blake 

seasons showed that 85% of a1 1 fishing and Goodson 1969). 

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ad--recommendations for their 

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gmLhus kisutch) with special 
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23(3):365-393. 
1017Y .I01 

17,. Sponsorin# Or&anlzmtlon Name and Addrass 

National Coastal Ecosystems Team U.S. Amy Corps of Engineers 

Fish and Wildlife Service LJaterways Ex per inen t Station 

U.S. Department of the Interior P.O. Box 631 

Uashicgton, D.C. 20240 Vicksburg, MS 39130 

REPORT DOCUMENTATION 1. '0. 2. 

PAGE i Biological Report 82(11.60)* 1 4. nt10 and subt~t~a Species Profiles: Life tli stories and Environmental 

Requirements of Coastal Fi shes and Invertebrates (Pacific Southwest) -- Steelhead 

7. Avlhofis) 

Roger A. Barnhart 9. ?ar(onnlng O~an~zrtlon Nam. and Addross 

California Cooperative Fishery Research Unit 

tlumboldt State University 

Arcata, CA 95521 

* U.S. Ar~y Corps of Engineers, TR EL-82-4 

la -net (Urnit: 200 mms) 

Species profiles are summaries of the literature on the taxonomy, life history, and 

environmental requirements of coastal fishes and aquatic invertebrates. They are 

prepared to assist with environmental impact assessnient. The steel head Salmo gairdneri , an anadrornous rainbow trout, supports an important sport fishery in the Pacific 

Southwest. Although native populations of steelhead have declined, these fish 

annually enter coastal streams from northern to southern California in years when 

winter stream flow is high. Steelhead ascend coastal streams from the ocean to 

spawn in cool, well-oxygenated waters with suitable depth, current velocity, and 

gravel size. After hatching, steelhead fry emerge fron the gravel and begin a freshwater rearing phase that generally extends fron 1 to 3 years. Rearing habitat with 

proper environmental conditions is extremely important to steel head production. 

Excessive sedimentation reduces food production, pool depth, and cover--a1 1 important 

to juvenile steelhead survival. Steelhead smol ts migrate during spring to saltwater, 

where most of their growth and sexual nlaturi ty is attained in 1 or 2 years. Attempts 

have increased to protect wild steelhead stocks, to maintain existing spawning and 

rearing habitat, to restore or enhance degraded habitat where feasible, to use 

artificial propagation efficiently, and to establ isli fishing regulations that provide 

qua1 i ty angl ing for steel head. 

1. R-cn~a.-t's Acc.swon NO 

r R~~QII oat. 

June 1986 

C. 

8. Parfomlng Organllallon Ra.1. NO 

10. ?ml~RRasklWorh Unat No. 

11. Cancract(o or trant(t) NO. 

(c1 

I?. Dscumnt 

Trout 

Analpis a. 0.scripton 

Salini'ty Feeding habits Growth 

rn 

Streams Temperature Food chains Oxygen 

Fisheries Sediments Life cycles Animal migrations 

I Steelhead trout 

Salnio gairdneri I Spawni no resuiremen ts 

! I unclassified - I 1 

(S.. ANSI-239.18) OPTIONAL FORM 272 (4-771 

War-rlv NTIS-35) 

c. COSATI Fiald/trouo 

lb Avatlab\lih Stalarnwnt 19. Secunly Class Ohm Rewn) 21. NO. Of Pages 

Unclassified Unl inii ted 21 --- 20. Suurltv Class nhls Pane) I 9. Pnce 
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