AD-Ai73 662 SPECIES PROFILES LIFE HISTORIES AND ENVIRONMENTAL I/REQUIREMENTS OF COASTA (U) CALIFORNIA COOPERATIVEFISHERY RESEARCH UNIT ARCATA CA R A BARNHART JUN 86

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Biological Report 82 (11.60) TR EL-82-4June 1986

N Species Profiles: Life Histories and(0 Environmental Requirements of Coastal Fishes

and Invertebrates (Pacific Southwest)*~OCT 2 91986~,

STEELHEAD

IJ

Coastal EogyGroupF Fis h and Wildlife Service Waterways Experiment Station

U.S. Department of the Interior U.S. Army Corps of Engineers

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Biological Report 82(11.60)

TR EL-82-4June 1986

Species Profiles: Life Histories and Environmental Requirementsof Coastal Fishes and Invertebrates (Pacific Southwest)

STEELHEAD

by

Roger A. Barnhart "-7California Cooperative Fishery Research Unit

Humboldt State UniversityArcata, CA 95521

Project OfficerJohn Parsons

National Coastal Ecosystems TeamU.S. Fish and Wildlife Service

1010 Gause BoulevardSlidell, LA 70458

Performed for

Coastal Ecology GroupWaterways Experiment StationU.S. Army Corps of Engineers

Vicksburg, MS 39180

o %and

National Coastal Ecosystems Team

Research and DevelopmentFish and Wildlife Service

U.S. Department of the InteriorWashington, DC 20240

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PREFACE

This species profile is one of a series on coastal aquatic organisms,principally fish, of sport, commercial, or ecological importance. The profilesare designed to provide coastal managers, engineers, and biologists with a briefcomprehensive sketch of the biological characteristics and environmentalrequirements of the species and to describe how populations of the species may beexpected to react to environmental changes caused by coastal development. Eachprofile has sections on taxonomy, life history, ecological role, environmentalrequirements, and economic importance, if applicable. A three-ring binder isused 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 Engineersand the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directed to one ofthe following addresses.

Information Transfer Specialist

National Coastal Ecosystems TeamU.S. Fish and Wildlife ServiceNASA-Slidell Computer Complex

1010 Gause BoulevardSlidell, LA 70458

or

U.S. Army Engineer Waterways Experiment StationAttention: WESER-CPost Office Box 631Vicksburg, MS 39180

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By

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CONVERSION TABLE

Metric to U.S. Custonary

Multiply § To Obtain

millimeters (mm) 0.03937 inchescentimeters (cm) 0.3937 inchesmeters (W) 3.281 feetkilometers (kin) 0.6214 miles

square meters (m2 ) n2 10.76 square feet

square kilometers (km) 0.3861 square inileshectares (ha) 2.471 acres

liters (1) 0.2642 gallonscubic meters (m3) 35.31 cubic feetcubic meters 0.0008110 acre-feet

milligrams (mg) 0.00003527 ouncesgrams (g) 0.03527 ounceskilograms (kg) 2.205 poundsmetric tons (t) 2205.0 poundsmetric tons 1.102 short tonskilocalories (kcal) 3.968 British thermal units

Celsius degrees 1.8(-C) + 32 Fahrenheit degrees

U.S. Customary to Metric

inches 25.40 millimetersinches 2.54 centimetersfeet (ft) 0.3048 metersfathoms 1.829 metersmiles (mi) 1.609 kilometersnautical miles (nmi) 1.852 kilometers

:1 square feet (ft2) 0.0929 square metersacres 0.4047 hectares

-- square miles (mi') 2.590 square kilometers

gallons (gal) 3.785 literscubic feet (ft') 0.02831 cubic metersacre-feet 1233.0 cubic meters

ounces (oz) 28.35 grainspounds (lb) 0.4536 kilogransshort tons (ton) 0.9072 metric tonsBritish thermal units (Btu) 0.2520 kilocalories

Fahrenheit degrees 0.5556(0F - 32) Celsius degrees

iv

CONTENTS

Page

PREFACE ...................................................................... iiiCONVERSION TABLE .............................................................. ivACKNOWLEDGMENTS ............................................................... Vi

NOMENCLATURE/TAXONOMY/RANGE .................................................. 1MORPHOLOGY/IDENTIFICATION AIDS ............................................... 1REASON FOR INCLUSION IN SERIES ........................................... ... 4LIFE HISTORY ................................................................ 4THE FISHERY .............................................................. ... 6ECOLOGICAL ROLE .......................................................... ... 8ENVIRONMENTAL REQUIREMENTS ................................................... 9

Temperature ................................................................. 9Dissolved Oxygen ........................................................... 10Depth ................................................................... ..11Water Movement .......................................................... .. 11Sediment ................................................................... 12

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

v!

ACKNOWLEDGMENTS

I thank Dennis P. Lee, California Department of Fish and Game, and Terry D.Roelofs, Humboldt State University, for reviewinq the manuscript; Thomas H.Hassler, California Cooperative Fishery Research Unit,for acting as the liaisonwith the National Coastal Ecosystems Team; and Delores Neher for greatlyfacilitating the preparation of this report.

Ai

* vi

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Figure 1. Steelhead.

STEELHEAD

NOMENCLATURE/TAXONOMY/RANGE migration (Swift 1975). Early inthis century steelhead distribution

Scientific name .......... ... Salmo in the Pacific Southwest was moregairdneri Richardson widespread (Figure 3). Steelhead

Preferred common name . . . Steelhead run up the Santa Domingo River in(Figure 1) northern Baja California during high

Other common names . . . . steelhead runoff in winter (Needham and Gardtrout, steelie, half-pounder, iron- 1959).head

Class ............ .OsteichthyesOrder .... ......... Salmoniformes MORPHOLOGY/IDENTIFICATION AIDSFamily ............. .Salmonidae

The following descriptions wereGeographic range: At sea, from taken from McConnell and Snyder (1972)

northern Baja California to the and Fry (1973).Bering Sea and Japan. In thePacific Southwest, steelhead enter At sea, steelhead are steel bluecoastal streams from the northern above and bright silver on the sidesCalifornia border south to the and belly. Sharply defined blackVentura River (Figure 2). Adult spots are located on back, head,steelhead have been reported in the sides, and dorsal and caudal fins.Santa Clara and Santa Margarita The spots are small and vary inRivers in years when winter runoff number. After entering freshwater,was high enough to allow upstream steelhead gradually take on the

1240 1200

OREGONSmith 420-

kiamath

SAlmon

Mad rlnlt

E el.........NorthwestSouthwe g

M00tle Sacramento

Noyo FtathNavarro . Yuba

Garcia .*NEVADA

FRANCISCO:SAY

C ALIFOR NIA

PAC I FIC

OCEAN

N

10 IO ~ yeVn ura SantaClara

0 100 200km NEE

Santa

330 -. ,, Terminal DamMagrt

1150

Figure 2. Distribution of steelhead (shaded areas) in Pacific Southwest regionstreams in 1980's (California Department of Fish and Game, pers. comm.; Moore1980).

2

OCEAN

NN

0 50 WOmi

o 160 2;0km,

Figure 3.istribution of steelhead (shaded areas) in the Pacific Southwestregion in 1900 (National Council on Gene Resources 1982).

3

appearance of stream rainbow trout; LIFE HISTORYthe back becomes olive green and thesides and belly less silvery. Steelhead trout spend a portionMaturing adults usually develop a of their life in the ocean where mostbroad pink stripe along the lateral of their growth occurs and sexualline and pink coloration on the maturity is attained; then they enteroperculum. Steelhead lack the red freshwater to spawn. Spawning gener-streaks beneath the jaw which ally takes place from February to latecharacterize the cutthroat trout June. The eggs are laid in pits(Salmo clarki). Juvenile steelhead in (redds), dug in the gravel of thestreams cannot be distinguished from stream bottom by the female. Imme-juvenile resident rainbow trout. diately after the eggs are laid and

fertilized, they are covered withAnal rays 9-12, rarely 13; teeth gravel by the female; the length of

on tip and shaft of vomer and on their stay in the gravel depends upontongue; gill rakers on lower limb of water temperature, dissolved oxygenfirst arch range from 17-21; scales in concentration and substrate composi-first row above lateral line 115-180; tion. After the eggs hatch, the youngcaudal fin shallowly forked; maxillary steelhead gradually work their way todoes not reach past the posterior the surface of the stream bed. Juve-margin of eye; juvenile parr marks niles usually spend a year or longersmall and oval to nearly round. in freshwater (length of residence is

determined by environmental andGenerally adult steelhead in the genetic factors) and then descend to

Pacific Southwest region weigh less the ocean.than 4.5 kg but some exceed 11 kg.

The steelhead is a strain ofrainbow trout that has a strong urgeto migrate to the ocean; however, someindividuals may remain in a stream,

REASON FOR INCLUSION IN SERIES mature, and even spawn without goingto sea (Shapovalov and Taft 1954).

The steelhead is the most The life history of the steel-widespread anadromous sport fish in head trout varies more than that ofthe Pacific Southwest region, but its any other anadromous fish regardingabundance has declined since the the length of time spent at sea, the1950's because of increased sport length of time spent in freshwater,fishing intensity and damage to fish and the times of emigration from andhabitat through construction of dams, immigration to freshwater. Unlikewater diversion, road construction, salmon, steelhead do not usually dieand improper land management just after spawning.practices. In some rivers a steelheadfishery would be nonexistent without Steelhead are classified intohatchery stocks. two races (Withler 1966; Smith 1960,

1969; and Everest 1973): winterIn California, attempts are steelhead that enter streams between

being made to protect wild stocks of November 1 and April 30, and summersteelhead to maintain existing steelhead that enter streams betweenspawning and rearing habitat, to May 1 and October 30. Portions ofrestore or enhance degraded habitat, both groups may enter freshwater into use artificial propagation wisely, spring or fall and are then calledand to regulate the fishery to provide spring- or fall-run steelhead. Inquality angling (California Department large rivers, such as the Klamath andof Fish and Game 1975). Sacramento Rivers, steelhead may enter

4

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during most of the year. Winter-run Everest 1973). Summer steelhead spawnsteelhead usually enter freshwater as in January and February, whereasmaturing fish that spawn relatively winter steelhead spawn in April andsoon, whereas most summer steelhead May, and summer steelhead spawn inenter as immature fish and do not smaller streams or farther upstream.mature and spawn until several months The sex ratio of steelhead troutlater. In the Pacific Southwest, immigrants is about 1:1 (Shapovalovsummer steelhead are not abundant and and Taft 1954; Kesner and Barnhartthe runs in many streams consist of 1972). Female steelhead containless than 100 fish (Roelofs 1983). about 2,000 eggs per kilogram of bodyDegradation of habitat in inter- weight (Moyle 1976).mittent streams and susceptibility toangling and predation probably account Several researchers havefor the low numbers. The southernmost concluded that the incidence of steel-summer steelhead population is in the head spawning more than once increasesMiddle Fork Eel River. This river from north to south (Bali 1959;supports the largest run of these fish Withler 1966; Sheppard 1972). Widein the Pacific Southwest--up to 2,000 variations in the percent of repeatfish in a good year. spawning can be due to genetic

factors, habitat quality, fishingAn exception is the "half- intensity, and management practices.

pounder" summer steelhead (terminology Fish that have spawned twice make upof Snyder 1925) which has a unique 70% to 85% of repeat spawners, whereaslife history described by Kesner and those that have spawned three timesBarnhart (1972) and Everest (1973). make up 10% to 25% of all repeatHalf-pounder runs are confined to a spawners (Forsgren 1979). The fewsmall geographic range encompassing fish that have spawned four times areabout 120 miles of the southern Oregon likely to be females, which have aand northern California coasts, higher survival rate than males duringincluding the Rogue, Klamath, Mad and and following spawning. SpawningEel Rivers. These small immature males usually each serve more than onesteelhead (25-35 cm long) annually female, remain in the stream longerenter freshwater from late August to than the females (tagging studies byearly October, and are the basis of Jones (1974) indicated nearly twoimportant sport fisheries in the weeks longer), and are exposed to moreKlamath and Rogue systems. Half- prolonged physical exertion than thepounders spend only a few months at females (Meigs and Pautzke 1941).sea before they return to freshwaterand, in contrast to mature steelhead,feed extensively 4- freshwater. Half- Steelhead spawn in cool, clear,pounders that survive their first well-oxygenated streams with suitableupstream migration return to the ocean depth, current velocity, and gravelthe following spring and migrate back size (Reiser and Bjornn 1979). Mea-to freshwater as mature steelhead in surements made over steelhead reddsthe summer and fall. Everest (1973) showed that steelhead spawn at depthsreported that half-pounders annually of 0.10-1.5 m, current velocities ofcomprise about 65% of the summer-run 23-155 cm/sec, and in gravel of 0.64-of steelhead on the Rogue River and 12.70 cm in diameter (Smith 1973;that 97% of all adult summer steelhead Hunter 1973; Bovee 1978; Wesche andfrom there make their first upstream Rechard 1980). Intermittent streamsmigration as half-pounders. are often used by steelhead for

spawning (Everest 1973; Kralik andSummer and winter steelhead do Sowerwine 1977; Carroll 1984). Most

not interbreed; they are isolated of the fry produced emigrate to peren-temporally and spatially (Smith 1969; nial streams soon after hatching.

' *' .%

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The embryology of the steelhead (usually 2 years in the Pacificis similar to that of other salmonids Southwest) before becoming smolts and(Wales 1941). The number of days migrating to saltwater. Most of therequired for steelhead eggs to hatch migrants are 2 years old and 18.6 tovaries from about 19 at an average 21 cm long (fork length) in thewater temperature of 15 °C to about 80 Klamath River (Kesner and Barnhartdays at an average of 5 °C. Steelhead 1972), or 14 to 21 cm long in Waddellfry usually emerge from the gravel 2 Creek (Shapovalov and Taft 1954). Theto 3 weeks after hatching. larger the smolts, the better the

survival. Most steelhead smolts enterAfter emergence, steelhead fry the sea in March and April.

(often in small schools) usually livein shallow water along the stream Steelhead live 1 to 4 years inbanks. As the fry grow older the the ocean (usually 1 or 2 years in theschools break up and the individual Pacific Southwest). The length offish establish territories which they residence in both freshwater and salt-defend. Most steelhead in their first water increases from south to northyear of life tend to inhabit riffles (Withler 1966). Steelhead growbut some of the larger fish inhabit rapidly in the ocean and their sizepools or deeper faster runs. Mortal- upon reaching maturity depends pri-ity is high during the first few marily on how long they have lived inmonths after emergence and many the ocean. In California, the averageinvestigators have suggested that the length of adult steelhead after 2relative size of the year class is years in the ocean is 58 cm (Withler

k largely determined at that time 1966). Immature steelhead trout (half-(Chapman 1966; McFadden 1969; Burns pounders) increase about 30 mm in1971; Everest and Chapman 1972). length each month from the time they

enter the ocean until they return toIn recent years, habitat degra- freshwater (Kesner and Barnhart 1972).

dation has lowered the capacity ofmany itreams to rear steelhead to The distribution or migration ofsmolts . For example, excessive sedi- steelhead in the ocean is not wellmentation has reduced food production, defined, particularly in the Pacificpool depth, and cover--all important Southwest. As judged by tag returns,to juvenile steelhead survival, most steelhead tend to migrate north

and south along the Continental Shelf.Juvenile steelhead feed on a Steelhead stocks from the Klamath and

wide variety of aquatic and Rogue Rivers probably mix together interrestrial insects. Newly emerged a nearshore ocean staging area alongfry sometimes are preyed upon by older the northern California coast beforejuvenile steelhead. Young steelhead they migrate upriver (Everest 1973).moving about trying to find a suitableterritory are subject to the highestpredation (Shapovalov and Taft 1954; THE FISHERYChapman 1966).

Commercial fishing for steelheadJuvenile steelhead live in has been prohibited in the Pacific

freshwater for from 1 to 4 years Southwest since 1924. The species now___is managed exclusively as a sport

fishery. Most steelhead are caught in1Smolt: Term applied to an anadromous rivers rather than in the ocean. Thejuvenile salmonid that is physiologi- three most important steelhead troutcally prepared to adapt to a saltwater rivers are the Klamath and Eel Riversexistence; steelhead smolts lose their in northern California and theparr marks and become silvery. Sacramento River which empties into

6

R 6r orrr

San Francisco Bay. The California runs in the Pacific Southwest regionDepartment of Fish and Game (1965) in since the 1940's has required an in-its Fish and Wildlife Plan estimated crease in the stocking of hatcheryspawning runs of 221,000 steelhead in reared steelhead. A total of about 1.9the Klamath River and 82,000 steelhead million yearling steelhead for plant-in the Eel River. The Klamath River ing are supplied annually by Colemanstill has good runs of native National Fish Hatchery on Battlesteelhead, perhaps approaching 200,000 Creek, a tributary to the upper Sacra-fish in good years (Barnhart 1975). mento River, the Nimbus Fish HatcheryIn the upper Sacramento River from on the American River, the Feather1953 to 1959, the average run of adult River Hatchery on the Feather River,steelhead was 20,590 and consisted of and the Mokelumne River Hatchery on

' both natural and hatchery-produced the Mokelumne River. The goal is tofish (Hallock et al. 1961). mitigate the loss of steelhead trout

in those rivers that has been causedIn the California Fish and Wild- by dams and water diversion.

life Plan (1965), the State's annualspawning stock of steelhead was esti- The average weight of hatcherymated at 603,000 fish. The Plan also yearlings at the time of plantingreported that the State contained (January through May) is about 518,402 linear miles of steelhead habi- grams (9 fish per pound). At antat, 31% of which was available to expected return of about 2% annually,anglers. It was estimated that steel- planted steelhead contributed abouthead anglers fish 304,000 angler-days 38,000 adults to the run in theper year to harvest 122,000 fish Sacramento River (Hallock et al.(about 0.4 fish per angler-day). Many 1961). If steelhead were not stocked,California streams support a substan- the catch in the Sacramento Rivertial fishery for juvenile steelhead-- would be much smaller. The Californiafish that have not yet migrated to the Department of Fish and Game operatesocean. These 5- to 8-inch trout are four hatcheries on the coast thattaken in large numbers, mostly during produce about 1.5 million yearlingthe so-called summer trout season. steelhead annually. In addition,The fishing pressure for these trout trout rearing projects sponsored by(about 440,000 angler-days annually) the California Department of Fish andexceeds that on adult steelhead (Cali- Game, private groups, and countiesfornia Department of Fish and Game contribute an average of 114,0001965). yearlings annually (Boydstun 1977a).

On the Klamath system, where two ofAlthough adult steelhead do not the coast hatcheries are located,

commonly feed in freshwater, they are hatchery production contributes onlyreadily taken by angling. Bait fish- an estimated 8% of the run of adultsing is popular and effective. Salmon (Boydstun 1977b). California alsoeggs or clusters of roe or night releases nearly 3 million steelheadcrawlers are drifted through riffles, fingerlings annually (Jensen 1971).pools, and especially where a tribu-tary enters a large stream. Spinners Fishery regulations are used toand other artificial lures are also help protect declining steelheadused. When streams are low and clear, stocks from excessive fishing. Inartificial flies are sometimes effec- California, the daily bag limit istive. Most steelhead fishing is done three adult fish. All streams tribu-from the bank or by wading; drifting tary to major coastal rivers and mostin boats is popular in larger streams. small coastal streams are closed to

angling from mid-November to lateThe decline of wild, naturally April. Special regulations also are

.F produced steelhead in the spawning sometimes used. For example, a

.7

section of the upper Middle Fork Eel Corps also planted streamside vegeta-River has been permanently closed to tion and constructed fish passageangling to protect the adult summer weirs on some streams. In 1981-1983,steelhead trout that concentrate there an additional $1 million was spentin summer and fall. annually by California to enhance

spawning gravels on the upper Sacra-The use of hatchery reared trout mento River and the Shasta and Klamath

to boost populations is not entirely Rivers, and to construct salmonidbeneficial. Larger runs attract more rearing ponds (Rawstron and Hashagenfishermen, and more fishermen further 1984). Six Rivers National Forest inreduce the abundance of wild stocks. northern California during the past 5Also, the genetic mixing of hatchery years has carried out an extensiveand wild stocks could decrease the program to restore or enhance theadvantageous traits in wild stocks. spawning and rearing habitats ofThe steelhead trout policy of the anadromous fish (Overton et al. 1981;California Department of Fish and Game Overton 1984).limits artificial propagation andstocking to reduce such interferencewith natural salmonid stocks; such ECOLOGICAL ROLEmeasures are periodically reviewed toassess their effects on the wild In freshwater, young steelheadstocks. may be sympatric with sculpins, resi-

dent rainbow trout, coho salmonThe environmental quality of (Oncorhynchus kisutch), and in some

coastal rivers largely governs the instances other salmonids. Data are" level of steelhead production. In the lacking on the effects of interaction

Pacific Southwest, attempts have been of juvenile steelhead and residentmade to increase the protection of rainbow trout, largely due to theexisting spawning and rearing habitat difficulty in distinguishing betweenand to rehabilitate damaged habitat the fish. In the Pacific Southwest,where feasible. California's steel- resident rainbow trout are common inhead trout policy is directed toward streams above barriers to steelheadthree goals: (1) to protect and migration and fry sometimes driftimprove steelhead trout habitat; (2) downstream over the barriers. Bjornnto develop plans and programs for (1978) reported that steelhead fryassessment of habitat conditions and tended to displace juvenile residentadverse impacts, land use planning, trout in the Lemhi River, Idaho. Theand acquisition of interests in ecological requirements of the twostreams threatened with adverse devel- races are similar.opments; and (3) to study the effectsof habitat changes caused by over- Coho salmon and steelhead troutgrazing, gravel extraction, logging, are similar in geographical distribu-road construction, urbanization, and tion, systematic characteristics,water development, spawning locations, food habits, and

the length of time the young soend inIn 1982-84, California spent freshwater (Milne 1948), although

$900,000 annually on the restoration steelhead normally remain inof salmon and steelhead habitat. A freshwater longer than 1 year.Salmon Restoration Project was devel- Although both species live in similaroped within the California Conserva- habitat in their first year, Hartmantion Corps in 1980, and by the end of (1965) reported that in spring and1983, Corps enrollees removed 7,447 summer, most steelhead trout live incords of wood and debris from 233 riffles and most coho salmon live inmiles of streams used by anadromous pools. Several investigators havefish (Kreb 1984). The Conservation reported that spatial segregation is

,, ' ..

also vertically stratified; coho fish and marine mammals but the extentsalmon live near the stream surface and effect of predation are unknown.and steelhead near the bottom (Hartman1965; Peterson 1966; Edmundson et al. In freshwater, steelhead feed1968; Bustard and Narver 1975). primarily on immature aquatic stagesAlthough coho salmon hatch earlier and of insects and secondarily on matureconsequently are larger at first, terrestrial insects. Ephemeropterasteelhead fry grow so much faster that (mayflies), Diptera (true flies), andby late summer the size difference is Trichoptera (caddisflies) are usuallysmall. Interspecific competition is the most important taxa in the dietprobably not serious because of the (Shapovalov and Taft 1954; Royal 1972;initial difference in size, differ- Fite 1973; Hiss 1984). Juvenileences in habitat preference, and the steelhead are somewhat opportunistic,difference in age at first emigration feeding on almost any available insectto the sea (Fraser 1969). Much the (Fite 1973). The size of thesame relation was reported for juve- territory for a single juvenile isnile chinook salmon (Oncorhynchus determined largely by the availabilitytshawytscha) and juvenile steelhead by of food and the size of the fishChapman and Bjornn (1969). (Allen 1969). In the ocean, steelhead

feed on a variety of organismsincluding juvenile greenlings

Native and hatchery-reared (Hexaqrammos spp.), squids, and amphi-steelhead exhibit some competition. pods (LeBrasseur 1966; Manzer 1968).Steelhead trout planted at the wrongtime or at the wrong size tend to stayin the stream longer than usual and ENVIRONMENTAL REQUIREMENTSare more competitive with wild fish(Royal 1972). Heavy predation by Temperaturehatchery-raised steelhead smolts onchinook salmon fry below Coleman Water temperature affects allNational Fish Hatchery on the metabolic and reproductive activitiesSacramento River was reported by of fish, including such criticalMenchen (1981). Wagner (1967) stated functions as growth, swimming, and thethat ideally the stream is to serve ability to capture and assimilate foodonly as a highway to the sea and not (Tebo 1974). Optimum temperatureas a postliberation rearing area for requirements may vary, depending onhatchery products. Pollard and Bjornn the season and life stage. A(1973) reported that the scocking of productive steelhead stream shouldhatchery rainbow trout also caused a have summer temperatures in the rangelocalized temporary decrease in the of 10 to 15 0C and an upper limit ofabundance of juvenile steelhead. 20 °C. Steelhead have difficulty

extracting oxygen from water 8tIn coastal streams, steelhead temperatures much over 21 C

fry are preyed on by sculpins (Cottus regardless of the amount of oxygenspp.), larger steelhead, and rainbow present (Hooper 1973). Bell (1973)trout (Shapovalov and Taft 1954); by listed the preferred temperatures ofbirds such as the great blue heron young steelhead as 7.2 to 14.5 °C, the(Ardea herodias), belted kingfisher optimum as about 10 °C, and the upper(Ceryle alcyon), American dipper lethal limit as 23.9 °C. Bovee(n s mexicanus), and common (1978), who developed a probability-mergansers ergus merganser); by of-use temperature curve for rearinggarter snakes hThamno his spp.); and winter steelhead juveniles, wrote thatby various mamma s hapovalov and optimum temperatures ranged from 0 toTaft 1954; Sheppard 1972; Cross 1975). 24 °C and peaked at 15 °C. In studiesIn the ocean, steelhead are eaten by of the smolting of cultured steelhead

9

in the spi ing, Wagner (1974) and Dissolved OxygenKerstetter and Keeler (1976) reportedthat low temperatures extend the Stream-dwelling salmonids requiresmolting period and high temperatures high dissolved oxygen in both the"'shorten it. They reported that smolt- water column and intragravel waters.

ing ceased rather abruptly when water The swimming performance of juveniletemperatures increased to 14-18 0C. and adult salmon was impaired when the

dissolved oxygen concentration wasDuring the spawning season a reduced below the air-saturation level

sudden drop in water temperature may (Davis et al. 1963). A sharp decreasecause all salmonid spawning activity in performance was noted at 6.5-7.0to cease (Reiser and Bjornn 1979). mg/l. Reiser and Bjornn (1979) wroteReingold (1968) reported that water that the oxygen levels recommended fortemperatures of 2 to 10 °C impaired spawning anadromous fish (at least 80%the viability of eggs and delayed the of saturation, with temporary levelsripening of adult steelhead held for no lower that 5.0 mg/l) should meet51 days in a hatchery pond. Bovee the needs of migrating salmonids.(1978) reported a spawning temperatureranae for winter steelhead of 4 to13 C, and a peak of 8 °C. Bell Intragravel dissolved oxygen(1973) indicated that steelhead concentration is positively related tospawning temperatures are generally survival of salmonid embryos (Coblefrom 3.9 to 9.4 0C. The average 1961; Phillips and Campbell 1962;development time from fertilization to Silver et al. 1963). Silver et al.hatching lengthens considerably with (1963), in a controlled laboratorydecreasing temperature; f~r rainbow experiment conducted at 9.5 *C, foundtrout it is 19 days at 15 C, 31 days that steelhead embryos hatchedat 10 0C, and 80 days at 5 6C (Embody successfully at mean oxygen concen-1934). Bovee (1978) gave an trations as low as 2.6 mg/l but thatincubation temperature range for total mortality occurred at a meanwinter steelhead of 0 to 24 °C and an level of 1.6 mg/l. In a field experi-optimum temperature of about 10 °C. ment Phillips and Campbell (1962)

noted that total mortality ofsteelhead embryos occurred at mean

When water temperatures fall oxygen concentrations of 7.2 mg/l orbelow 4 °C in streams of the Pacific less. Although intragravel oxygenNorthwest, juvenile steelhead become levels may appear adequate, the amountinactive and hide in available cover of oxygen actually reaching theor in the substrate (Chapman and embryos also depends on the intra-Bjornn 1969; Bustard and Narver 1975). gravel water velocity (Wickett 1954).In California's coastal streams, The amount of intragravel oxygenjuvenile steelhead remain active year- available to developing embryos isround and in one small coastal stream, sometimes reduced by the biochemicalyoung steelhead grew throughout the oxygen demand of organic material inwinter (Reeves 1979). the gravel bed (Ponce 1974). Even if

embryos hatch at low or moderatelyThe virulence of many fish reduced oxygen levels, the incubation

diseases and the toxicity of most period is extended and the resultingchemicals increase with increasing fry are likely to be smaller andwater temperatures (Lantz 1971). weaker than those reared at oxygenRemoval of riparian vegetation can concentrations close to saturationresult in marked increases in summer (Silver et al. 1963; Shumway et al.stream temperatures and sometimes in 1964). Reiser and Bjornn (1979)reduced winter temperatures (Brown concluded that although dissolved1971). oxygen concentrations required for

10

.,. A 7U

successful incubation depend on both Steelhead tend to occupy thespecies and developmental stage, shallow riffle areas of streams,concentrations at or near saturation particularly during the first year ofwith no temporary reductions below 5.0 life (Hartman 1965). Bovee (1978)mg/l are generally required by anadro- presented probability of use curvesmous salmonids. showing that steelhead fry are most

commonly found in water 8 to 36 cmIn salmonid nursery and rearing deep, and steelhead juveniles are

streams dissolved oxygen concentra- usually located in water 25 to 50 cmtions of surface waters are normally deep. In the Southwest region steel-near saturation, except in small head streams are annually subjected totributaries with large amounts of low flow conditions due to thedebris from logging or other sources extended summer-fall dry season; thus,or in large, slow-moving streams pool frequency and depth are impor-receiving large amounts of municipal tant. In a 2-year study at Singleyor industrial waste (Reiser and Bjornn Creek, a small coastal stream just1979). Salmonids function normally at south of Cape Mendocino, Cross (1975)dissolved oxygen concentrations of found that 67%-96% of young-of-the-7.75 mg/l; exhibit various distress year steelhead resided in pools. Fromsymptoms at 6.00 mg/l; and are often June to September, the riffle surfacenegatively affected at 4.25 mg/l area was reduced 45% while the surface(Davis 1975). Low dissolved oxygen area of pools diminished only 26%.impairs metabolic rate, growth, swim- Excessive sediment inputs that fillming performance, and overall survival pools can greatly reduce a stream'sof young salmonids. capacity to rear steelhead to smolt

size.Depth

Water MovementWater depth usually does not

prevent migration because adult steel- Steelhead may encounter waterhead normally migrate when stream velocity barriers during upstreamflows are relatively high. Thompson migration, often at falls or culverts.(1972) wrote that 18 cm is the minimum Velocities of 3-4 m/s begin to greatlydepth required for successful hinder the swimming ability ofmigration of adult steelhead. The steelhead and may retard migrationmigration of adult salmonids is more (Reiser and Bjornn 1979). Thompsoncommonly hindered by excessive water (1972) outlined methods to calculatevelocity or obstacles that impede the minimum and maximum acceptable stream-swimming or jumping of the fish. flows for migrating adult steelhead,

for specific stream sections.

Water depth may be important in Bovee (1978) showed that steel-the selection of redd sites. head spawn in areas with waterShapovalov and Taft (1954) stated that velocities of 30-110 cm/s but that thesteelhead redds are rarely exposed by preferred velocity approximatedfalling stream levels. Bovee (1978) 60 cm/s. Smith (1973) found that theshowed that steelhead most commonly preferred water velocity for spawningspawn at depths averaging 36 cm steelhead in Oregon ranged from 40 to(range, 15-61 cm). The depths of 91 cm/s. Steelhead redds were inWashington winter steelhead redds areas where velocities were 15 to 54ranged from 12 to 70 cm (Hunter 1973). cm/s in a small tributary of theCarroll (1984) measured water depths Klamath River (Carroll 1984). Sinceof 12 to 29 cm over steelhead redds in swimming performance improves witha Klamath River tributary, size, large adult steelhead can

11

establish redds in faster current carrying capacity of the streamareas of the stream. directly by reducing available rearing

habitat and indirectly by reducing thePermeability is defined as the production of invertebrate food.

capacity of the gravel to transmit Bjornn et al. (1977) found significantwater. Different spawning gravels reductions in numbers of juveniletransmit water at different rates. steelhead in stream channels whereSeveral studies have demonstrated that boulders were imbedded in sediment.increased intragravel velocities Crouse et al. (1981), who devised aimprove the survival of steelhead eggs visual substrate scoring system basedand fry before emergence and also the on particle size and degree of cobblecondition of fry that emerge (Shuriway embedment, reported that fish1960; Silver 1960; Coble 1961; Silver production was significantlyet al. 1963; Shumway et al. 1964). correlated with substrate score. TheyMcNeil and Ahnell (1964) concluded reported significant decreases in fishthat the streambeds of highly produc- production in streams where cobblestive spawning streams had gravels with were embedded 80%-100% and wherehigh permeability (24,000 cm/h) and sediment (2.0 mm or less) composedhad less than 5% (by volume) sand and 26%-31% (by volume) of the totalsilt that passed through a sieve of substrate composition. The authors0.833 mmv mesh. concluded that rearing habitats of

juvenile salmonids in streams, as wellSediment as spawning gravels, require

protection from excessive quantitiesQuantitatively, sediment is the of fine sediments.

greatest single pollutant in thenation's water (Ritchie 1972). Ander-son (1971) reported that sediment The size of substrate materialproduction in northern California has been related by numerous investi-watersheds increased markedly as a gators to the standing crops of inver-result of poor land management tebrates (Sprules 1947; Kimble andpractices and floods. The steelhead's Wesche 1975). Pennak and VanGerpenenvironment can be impaired both by (1947) found that the number ofsediment in suspension and by par- benthic invertebrates decreased in theticles deposited as bedload sediment. progression from rubble to bedrock toAbout 28% of the total spawning area gravel to sand. Reiser and Bjornnin a once important 16-mile stretch of (1979) reported that aquatic insectthe upper Trinity River has been lost production was highest in substratedue to the deposition of sediment composed largely of coarse gravel(California Resources Agency 1970). (3.2-6.7 cm) and rubble (7.6-30.4 cm).Stream channels of northern Californiamarkedly aggraded after large flood Suspended sediment occasionallyevents during the 1960's and 1970's reaches concentrations high enough to(Lisle 1982). Channel widening and directly injure steelhead (3,000 ppmloss of pool-riffle sequence due to or greater) (Cordone and Kelley 1961).aggradation damaged spawning and Physiological damage includes therearing habitat of steelhead. The adhesion of silt particles to thepool-riffle sequence and pool quality chorion of salmonid ova (Cordone andin some northern California streams Kelley 1961) and the abrasion,still had not fully recovered by 1980 thickening, and fusion of gillfrom a 1964 regional flood (Lisle filaments (Herbert and Merkens 1961).1982). Sigler et al. (1984) reported that

chronic turbidity in streams duringFor rearing juvenile steelhead, emergence and rearing of steelhead

deposited sediment reduces the affects the numbers and quality of

12

-~~~~~~, J,* ~ n nr nt ~ a .. ~ ~- ~ ~ ~ * ~ ~ *

fish produced. Turbid water also took place when turbidities hadaffects recreational angling for decreased to 30 Jackson Turbiditysteelhead. A study on the Eel River Units or less; such low levels ofby the California Department of Fish turbidity occurred during onlyand Game during two winter steelhead one-third of the fishing season (Blakeseasons showed that 85% of all fishing and Goodson 1969).

13

V.4

LITERATURE CITED

Allen, K.R. 1969. Limitations on Bjornn, T.C., M.A. Brusven, M.P. Mol-production in salmonid populations nay, J.H. Milligan, R.A. Klamt, E.in streams. Pages 3-18 in T.G. Chaco, and C. Schaye. 1977. Trans-Northcote, ed. Symposium on Trout port of granitic sediment in streamsand Salmon in Streams. Univ. of and its effects on insects and fish.B. C., Inst. of Fish., Vancouver. Univ. Idaho Coop. Fish. Res. Unit

Bull. 17. Completion Rep. Proj.Anderson, H.W. 1971. Relative B-036-1DA. 43 pp.contributions of sediment fromsource areas and transport pro- Blake, R.B., and L.F. Goodson. 1969.cesses. Pages 55-63 in J. Morris, Fishing pressure and turbidity ined. Symposium, Forest Land Uses and the upper Eel River basin. Calif.Stream Environment. Oregon State Dep. Fish Game Rep., Region I. 6 pp.University Press, Corvallis. (Mimeo).

Bali, J.M. 1959. Scale analysis of Bovee, K.D. 1978. Probability of usesteelhead trout, Salmo gairdneri criteria for the family Salmonidae.gairdneri Richardson, from various Instream Flow Information Paper No.coastal watersheds of Oregon. M.S. 4. U.S. Fish Wildl. Serv. FWS/OBS-Thesis. Oregon State University, 78/07. 80 pp.Corvallis. 189 pp.

Boydstun, L.B. 1977a. California'sBarnhart, R.A. 1975. Pacific slope steelhead program. Pages 26-30 insteelhead trout management. Pages T.J. Hassler, R.R. Van Kirk, eds.7-11 in W. Kinq, ed. Wild Trout Proceedings, Genetic Implications ofManagement Symposium. Trout Steelhead Management Symposium.Unlimited, Denver, Colo. Calif. Coop. Fish. Res. Unit,

Humboldt State University, Arcata.Spec. Rep. 77-1.

Bell, M.C. 1973. Fisheries handbook

of engineering requirements and bio- Bovdstun, L.B. 1977b. Lower Klamathlogical criteria. U.S. Army Corps River tagging study. Calif. Dep.of Engineers, Portland, Oregon. Fish Game. Performance Rep.Contract No. DACW 57-68-C-0086. 425 AFS-20-3. 28 pp. (mimeo).pp.

Brown, G.W. 1971. Water temperatureBjornn, T.C. 1978. Survival, pro- in small streams as influenced by

duction and yield of trout and chi- environmental factors. Pages 175-nook salmon in the Lemhi River, 181 in J. Morris, ed. Symposium,Idaho. Coll. of Forestry, Wildl., ForesT Land Uses and Streamand Range Sciences, Univ. of Idaho, Environment. Oregon State Univer-Moscow. Bull. No. 27. 57 pp. sity Press, Corvallis.

Burns, J.W. 1971. The carrying Cordone, A.J., and D.W. Kelley.

capacity for juvenile salmonids in 1961. The influences of inorganicsome northern California streams. sediment on the aquatic life ofCalif. Fish Game 57:44-57. streams. Calif. Fish Game 42(2):

189-228.Bustard, D.R., and D.W. Narver. 1975.Aspects of the winter ecology of Cross, P.D. 1975. Early life historyjuvenile coho salmon (Oncorhxnchus of steelhead trout (Salmo gairdneri)kisutch) and steelhead trout (Salmo in a small coastal stream. M.S.aai airi). J. Fish. Res. Boar-rd. Thesis. Humboldt State University,Can. M):667-680. Arcata, Calif. 44 pp.

California Department of Fish and Crouse, M.R., C.A. Callahan, K.W.Game. 1965. Pages 323-679 in Malveg, and S.E. Dominquez. 1981.California Fish and Wildlife Plan, Effects of fine sediments on growthVol. 3, Part B, The Resources of juvenile coho salmon inAgency, Sacramento. laboratory streams. Trans. Am.

Fish. Soc. 110(2):281-286.California Department of Fish andGame. 1975. Steelhead rainbow Davis, G.E., J. Foster, C.E. Warren,trout policy. Pages 407-409 in Fish and P. Doudoroff. 1963. Theand Game Code, 1983. Sacramento. influence of oxygen concentration on

the swimming performance of juvenileCalifornia Resources Agency. 1970. Pacific salmon at various

Sediment problems in the Trinity temperatures. Trans. Am. Fish. Soc.River near Lewiston. Task Force 92(2):111-124.Rep. to Sec. for Res. Calif. Res.Agency. 32 pp. Davis, J.C. 1975. Minimal dissolved

oxygen requirements of aquatic lifeCarroll, E.W. 1984. An evaluation of with emphasis on Canadian species:

steelhead trout and instream a review. J. Fish. Res. Board. Can.structures in a California 32(12):2295-2332.intermittent stream. M.S. Thesis.Humboldt State University, Arcata, Edmundson, E., F.H. Everest, and D.W.Calif. 51 pp. Chapman. 1968. Permanence of

station in juvenile chinook salmonand steelhead trout. J. Fish. Res.

Chapman, D.W. 1966. Food and space Board. Can. 25(7):1453-1464.as regulators of salmonidpopulations in streams. Am. Nat. Embody, G.C. 1934. Relation of100:345-357. temperature to the incubation

periods of eggs of four species ofChapman, D.W., and T.C. Bjornn. 1969. trout. Trans. Am. Fish. Soc.Distribution of salmonids in 64:281-292.streams, with special reference tofood and feeding. Pages 153-176 in Everest, F.H. 1973. Ecology andT.G. Northcote, ed. Symposium on management of summer steelhead inTrout and Salmon in Streams. Univ. the Rogue River. Oregon State Gameof B. C., Inst. of Fish., Vancouver. Comm. Fish. Res. Rep. No. 7,

Corvallis. 48 pp.Coble, D.W. 1961. Influence of waterexchange and dissolved oxygen in Everest, F.H., and D.W. Chapman.redds on survival of steelhead trout 1972. Habitat selection and spatialembryos. Trans. Am. Fish. Soc. interaction by juvenile chinook

90(4):469-474. salmon and steelhead trout in two

1...!.

Idaho streams. J. Fish. Res. Board. California. M.S. Thesis, HumboldtCan. 22:1035-1081. State University, Arcata, Calif. 48

PP.Fite, K.R. 1973. Feeding overlap

between roach and juvenile steelhead Hooper, D.R. 1973. Evaluation of thein the Eel River. M.S. Thesis, effects of flows on trout streamHumboldt State University, Arcata, ecology. Dep. of Eng. Res., PacificCalif. 38 pp. Gas and Electric Co., Emeryville,

Calif. 97 pp.Forsgren, H.L., II. 1979. Age,

growth, origin and repeat spawning Hunter, J.W. 1973. A discussion ofof winter steelhead (Salmo game fish in the State of Washingtonairdneri) in Mad River, California. as related to water requirements.A.S. Thesis. Humboldt State Univer- Rep. by Wash. State Dep. Game, Fish.sity, Arcata, Calif. 56 pp. Manage. Div., Wash. State Dep. Ecol.

66 pp.

Fraser, F.J. 1969. Population den- Jensen, P.T. 1971. Salmon andsity effects on survival and growth steelhead. Pages 1-13 in Report toof juvenile coho salmon and steel- the State Water ResouFces Controlhead trout in experimental stream Board. Calif. Fish Game Environ.channels. Pages 253-266 in T.G. Serv. Admin. Rep. No. 71-2.Northcote, ed. Symposium 6F Troutand Salmon in Streams. Univ. of Jones, D.E. 1974. The study ofB.C., Inst. of Fish., Vancouver. cutthroat-steelhead in Alaska.

Alaska Dep. Fish Game, Div. of SportFry, D.H., Jr. 1973. Anadromous Fish. Anadromous Fish Studies.fishes of California. Calif. Dep. Annu. Prog. Rep. 1973-1974, StudyFish Game, Sacramento. 111 pp. AFS-42-2. 31 pp.

Hallock, R.J., W.F. VanWoert, and L. Kerstetter, T.H., and M. Keeler.Shapovalov. 1961. An evaluation of 1976. Smolting in steelhead troutstocking hatchery-reared steelhead Salmo gairdneri: a comparativerainbow trout (Salmo gairdneri study of populations in twogairdneri) in the Sacramento River hatcheries and the Trinity River,system. Calif. Dep. Fish Game Fish northern California, using gill Na,Bull. No. 114. 74 pp. K, ATPase assays. Humboldt State

University Sea Grant Project,Hartman, G.F. 1965. The role of Arcata, Calif. HSU-SG9. 26 pp.behavior in the ecology andinteraction of underyearling coho Kesner, W.D., and R.A. Barnhart.salmon (Oncorhynchus kisutch) and 1972. Characteristics of the fall-steelhead trout (Salmo 9airdneri). run steelhead trout (Salmo gairdneriJ. Fish. Res. Board. Can. 22(4): gairdneri) of the Klamath River1035-1081. system with emphasis on the half-

pounder. Calif. Fish GameHerbert, D.W.M., and J.C. Merkens. 58(3):204-220.

1961. The effect of suspendedmineral solids on the survival of Kinble, L.A., and T.A. Wesche. 1975. Ntrout. Int. J. Air Water Pollut. Relationship between selected .%5(l):46-53. physical parameters and benthic

community structure in a smallHiss, J.M. 1984. Diet of age-O mountain stream. Univ. Wyo.,

steelhead trout and speckled dace in Laramie. Water Resour. Res. Inst.Willow Creek, Humboldt County, Series 55. 64 pp.

17

-:- .. .. " " " " , " I " " " . . . . . % "' ..1

Kralik, N.J., and J.E. Sowerwine. Northcote, ed. Symposium on Salmon1977. The role of two northern and Trout in Streams. Univ. B. C.,California intermittent streams in Inst. Fish., Vancouver.the life history of anadromoussalmonids. M.S. Thesis. Humboldt McNeil, W.J., and W.H. Ahnell. 1964.State University, Arcata, Calif. 68 Success of pink salmon spawningpp. relative to size of spawning bed

materials. U.S. Fish Wildl. Serv.Kreb, M. 1984. California Conser- Spec. Sci. Rep. Fish. No. 469. 15

vation Corps project 201. Pages 10- pp.11 in K. Hashagen, C. Toole, B.Wyatt, S. Sommarstrom, S. Taylor, Meigs, R.C., and C.P. Pautzke. 1941.eds. Report of the 2nd Calif. Additional studies on the life his-Salmon and Steelhead Restoration tory of the Puget Sound steelheadConference. Univ. Calif. Sea Grant, (Salmo gairdneri). State Wash. Dep.UCSG-MAP-21, Davis. 97 pp. Game, Biol. Bull. No. 5. 13 pp.

Lantz, R.L. 1971. Influence of water Menchen, R.S. 1981. Predation bytemperature on fish survival, growth yearling steelhead Salmo 9airdneri,and behavior. Pages 182-193 in J. released from Coleman National FishMorris, ed. Symposium, Forest-Land Hatchery, on naturally produced chi-Uses and Stream Environment. Oregon nook salmon Oncorhynchus tshawytschaState University, Corvallis. 252 fry and eggs in Battle Creek, T975.PP. Calif. Dep. Fish Game, Anadromous

Fisheries Branch Office Report,LeBrasseur, R.J. 1966. Stomach Sacramento. 6 pp.

contents of salmon and steelheadtrout in the northeastern PacificOcean. J. Fish. Res. Board. Can. Milne, D.J. 1948. The growth,23(1):85-100. morphology and relationship of the

species of Pacific salmon and theLisle, T.E. 1982. The recovery of steelhead trout. Ph.D. Thesis.

stream channels in north coastal McGill University, Montreal. 101pp.California from recent large floods.Pages 31-41 in K. Hashagen, ed. Moore, M.R. 1980. Factors influ-Habitat DistuFance and Recovery encing the survival of juvenileProceedings. Calif. Trout, San steelhead rainbow trout, SalmoFrancisco. 90 pp. gairdneri in the Ventura River,

California. M.S. Thesis. HumboldtManzer, J.1. 1968. Food of Pacific State University, Arcata, Calif. 82

salmon and steelhead trout in the pp.Pacific Ocean. J. Fish. Res. Board.Can. 25(5):1085-1089. Moyle, P.B. 1976. Inland fishes of

California. University of Cali-McConnell, R.J., and G.R. Snyder. fornia Press, Berkeley. 405 pp.

1972. Key to field identificationof anadromous juvenile salmonids in National Council on Gene Resources.the Pacific northwest. NOAA (Natl. 1982. Anadromous salmonid geneticOcean. Atmos. Adm.) Tech. Rep. NMFS resources: an assessment and plan(Natl. Mar. Fish. Serv.) Circ. 366. for California. California Gene6 pp. Resource Program, Berkeley. 168 pp.

McFadden, J.T. 1969. Dynamics and Needham, P.R., and R. Gard. 1959.regulation of salmonid populations Rainbow trout in Mexico and Cali-in streams. Pages 313-329 in T.G. fornia with notes on the cutthroat .i'

18

!

series. Univ. Calif. Publ. Zool. Rawstron, R., and K. Hashagen. 1984.67: 1-124. California Department of Fish and

Game salmonid enhancement andOverton, K. 1984. U.S. Forest restoration activities. Page 12 in

Service Six Rivers National Forest. K. Hashagen, C. Toole, B. Wyatt, S.Page 37 in K. Hashagen, C. Toole, B. Sommarstrom, S. Taylor, eds. ReportWyatt, S. Sommarstrom, S. Taylor, of the 2nd Calif. Salmon andeds. Report of the 2nd Calif. Steelhead Restoration Conference.Salmon and Steelhead Restoration Univ. Calif. Sea Grant, USCG-MAP-21,Conference. Univ. Calif. Sea Grant, Davis. 97 pp.UCSG-MAP-21, Davis. 97 pp.

Reeves, G.H. 1979. Population dynam-Overton, K., W. Brock, J. Moreau, and ics of juvenile steelhead in rela-J. Boberg. 1981. Restoration and tion to density and habitat charac-enhancement program of anadromous teristics. M.S. Thesis. Humboldtfish habitat and populations on Six State University, Arcata, Calif. 67Rivers National Forest. Pages 158- pp.168 in T.J. Hassler, ed. Proceed-ings: Propagation, Enhancement, and Reingold, M. 1968. Water temperatureRehabilitation of Anadromous Sal- affects the ripening of adult fallmonid Populations and Habitat chinook salmon and steelhead. Prog.Symposium. Humboldt State Univer- Fish-Cult. 30(1):41-42.sity, Arcata, Calif. 168 pp.

Reiser, D.W., and T.C. Bjornn. 1979.Pennak, R.W., and E.D. VanGerpen. Habitat requirements of anadromous

1947. Bottom fauna production and salmonids. 54 pp. in W.R. Meehan,physical nature of the substrate in ed. Influence of F~Fest and Rangea northern Colorado trout stream. Management on Anadromous FishEcology 28:42-48. Habitat in Western North America.

Pacific N.W. Forest and Range Exp.Peterson, G.R. 1966. The relation- Sta. USDA For. Serv., Portland.

ship of invertebrate drift abundance Gen. Tech. Rep. PNW-96.to the standing crop of benthicorganisms in a small stream. M.S. Ritchie, J.C. 1972. Sediment, fishThesis. University of British and fish habitat. J. Soil WaterColumbia, Vancouver. 39 pp. Conser. 27(3):124-125.

Phillips, R.W., and H.J. Campbell. Roelofs, T.D. 1983. Current status1962. The embryonic survival of of California summer steelheadcoho salmon and steelhead trout as (Salmo gairdneri) stocks and habitatinfluenced by some environmental and recommendations for theirconditions in gravel beds. Pac. management. USDA For. Serv., RegionMar. Fish. Comm. 14th. Annu. Rep. 5, San Francisco, Calif. 77 pp.1961: 60-75.

Royal, L.A. 1972. An examination ofPollard, H.A., II, and T.C. Bjornn. the anadromous trout program of the

1973. The effects of angling and Washington State Game Department.hatchery trout on the abundance of Wash. State Dep. Game, Final Rep.juvenile steelhead trout. Trans. AFS-49, Olympia. 176 pp.Am. Fish. Soc. 102(4):745-752.

Shapovalov, L., and A.C. Taft. 1954.Ponce, S.L. 1974. The biochemical The life histories of the steelheadoxygen demand of finely divided rainbow trout (Salmo gairdnerilogging debris in stream water. gairdneri) and silver salmonWater Resour. Res. 10(5):983-988. 19 n-corhnchus kisutch) with special

19

k"'

reference to Waddell Creek, Call- rainbow trout, Salmo gairdnerifornia, and recommendations regard- Richardson. Ph.D. is. Univer-ing their management. Calif. Dep. sity of Alberta, Edmonton, Alberta.Fish Game Fish. Bull. 98. 375 pp.

Smith, S.B. 1969. ReproductiveSheppard, D. 1972. The present isolation in summer and winter races

status of the steelhead trout stocks of steelhead trout. Pages 21-38 inalong the Pacific coast. Pages 519- T.G. Northcote, ed. Symposium on556 in D.H. Rosenberg, ed. A Review Salmon and Trout in Streams. Univ.of the Oceanography and Renewable B. C., Inst. Fish. Vancouver.Resources of the Northern Gulf ofAlaska. IMS Rep. R72-23, Sea GrantRep. 73-3. Institute of Marine Snyder, J.0. 1925. The half-pounderScience, University of Alaska, of Eel River, a steelhead trout.Fairbanks. 690 pp. Calif. Fish Game 11(2):49-55.

Shumway, D.L. 1960. The influence of Sprules, W.M. 1947. An ecologicalwater velocity on the development of investigation of stream insects insalmonid embryos at low oxygen Algonquin Park, Ontario. Univ.levels. M.S. Thesis. Oregon State Toronto Stud. Biol. 56., Publ. Ont.University, Corvallis. 49 pp. Fish. Res. Lab. 69:1-81.

Shumway, D.L., C.E. Warren, and P. Swift, C.C. 1975. Survey of the

Doudoroff. 1964. Influence of freshwater fishes and their habitatsoxygen concentration and water in the coastal drainages of southernmovement on the growth of steelhead California. Natural History Museumtrout and coho salmon embryos. of Los Angeles County, California.Trans. Am. Fish. Soc. 93(4):342-356. 364 pp.

Sigler, J.W., T.C. Bjornn, and F.H.Everest. 1984. Effects of chronic Tebo, L.B., Jr. 1974. Review ofturbidity on density and growth of selected parameters of trout streamsteelheads and coho salmon. Trans. quality. Pages 20-32 in SymposiumAm. Fish. Soc. 113:142-150. on Trout Habitat Research and

Management, Proceedings. Appala-Silver, S.T. 1960. The influence of chian Consortium Press, Boone, N.C.water velocity and dissolved oxygenon the development of salmonid Thompson, K. 1972. Determiningembryos. M.S. Thesis. Oregon State stream flows for fish life. PagesUniversity, Corvallis. 50 pp. 31-50 in Proceedings, Instream Flow

Requirement Workshop. Pac. N.W.Silver, S.J., C.E. Warren, and P. River Basin Comm., Vancouver, Wash.Doudoroff. 1963. Dissolved oxygenrequirements of developing steelheadtrout and chinook salmon embryos at Wagner, H.H. 1967. A summary ofdifferent water velocities. Trans. investigations of the use ofAm. Fish. Soc. 92(4):327-343. hatchery-reared steelhead in the

management of a sport fishery.Smith, A.K. 1973. Development and Oregon State Game Comm. Fish. Rep.

application of spawning velocity and 5. 62 pp.depth criteria for Oregon salmonids.Trans. Am. Fish. Soc. 102:312-316. Wagner, H.H. 1974. Photoperiod and

temperature regulation of smoltingSmith, S.B. 1960. Racial charac- in steelhead trout (Salmo gaird-

teristics in stocks of anadromous neri). Can. J. Zool. 52 -82.12.

20

Wales, J.H. 1941. Development of Wickett, W.P. 1954. The oxygensteelhead trout eggs. Calif. Fish supply to salmon eggs in spawningGame 27(4):250-260. beds. J. Fish. Res. Board. Can.

11(6):933-953.

Wesche, T.A., and P.A. Rechard. 1980. Withler, I.L. 1966. Variability inA summary of instream flow methods life history characteristics offor fisheries and related research steelhead trout (Salmo gairdneri)needs. Univ. Wyo., Water Resour. along the Pacific coast of NorthRes. Inst., Eisenhower Consortium America. J. Fish. Res. Board. Can.Bull. 9. 122 pp. 23(3):365-393.

.

21

' .MWX - d7L P. ,'

REPORT DOCUMENTATION L EPORT NO. 3. Rec,.-t's AcCeso. No

PAGE iBiological Report 82(11.60)*" -Wn. at,., s.,.u. Species Profiles: Life Histories and Environmental S Re.o o.,.Requirements of Coastal Fishes and Invertebrates (Pacific South- June 1986west) -- Steelhead 6

7. Authets) L Perform-n Orgenization Root. No

Roger A. Barnhart9. ftfl*emw Og11.,et1-t Name aind Address 20. Proiectf/reshiWot Unit No.

California Cooperative Fishery Research UnitHumboldt State University 1. Contract=or Graf(G) No.Arcata, CA 95521 (C)

(G)I2. Spenalaeni. Ovulentoatn Nme end Address

National Coastal Ecosystems Team U.S. Army Corps of Engineers ILType oReot&Pon.dCoved

Fish and Wildlife Service Waterways Experiment StationU.S. Department of the Interior P.O. Box 631Washirgton, D.C. 20240 Vicksburg, MS 39180 z,.

IL Supoeaweey Not"

* U.S. Army Corps of Engineers, TR EL-82-4

I& Allitract1 CUmut MO eerg)*NSpecies profiles are summaries of the literature on the taxonomy, life history, and

*, environmental requirements of coastal fishes and aquatic invertebrates. They areprepared to assist with environmental impact assessment. The steelhead Salmo gairdneri,an anadromous rainbow trout, supports an important sport fishery in the PacificSouthwest. Although native populations of steelhead have declined, these fishannually enter coastal streams from northern to southern California in years whenwinter stream flow is high. Steelhead ascend coastal streams from the ocean tospawn in cool, well-oxygenated waters with suitable depth, current velocity, and-*gravel size. After hatching, steelhead fry emerge from the gravel and begin a fresh-water rearing phase that generally extends from 1 to 3 years. Rearing habitat withproper environmental conditions is extremely important to steelhead production.Excessive sedimentation reduces food production, pool depth, and cover--all importantto juvenile steelhead survival. Steelhead smolts migrate during spring to saltwater,where most of their growth and sexual maturity is attained in 1 or 2 years. Attemptshave increased to protect wild steelhead stocks, to maintain existing spawning andrearing habitat, to restore or enhance degraded habitat where feasible, to use

o. artificial propagation efficiently, and to establish fishing regulations that providequality angling for steelhead.

17. 0oue nne. Anely.,e a. Oeicnitton

Trout Salinity Feeding habits GrowthStreams Temperature Food chains OxygenFisheries Sediments Life cycles Animal migrations

SI. Identfier/Ogenclnded Terms

Steelhead troutSalmo gairdneriSpawning requirementsRearing requirements

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Eastern Energy antd Land Use TeamLeetown WVj

*Nahonal Coastal Ecosystems Team L -Virgin lilands

S OetI LA:0 Western Energy and Land Use TeamFt Col trns CO

Locations of Regional Offices

REGION I REGION 2 REGION 3Regional Director Regional Director Regional DirectorU.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife ServiceLloyd Five Hundred Building. Suite 169~2 P.O. Box 1306 Federal Building. Fort Snelling500 N.E. Multnomnah Street Albuquerque, New Mexico 87103 Tw.in Cities, Minnesota 55111Portland. Oregon 97232

REGION 4 REGION 5 REGION 6Regional Direct or Regional Director Regional DirectorU.S. Fish and Wildlife Service U.S. Fish and Wildlife Service U.S. Fish and Wildlife ServiceRichard B. Russell Building One Gateway ('enter P.O. Box 2548675 Spring Street. S.W. Newton Corner, Massachusetts 02158 Denver Federal CenterAtlanta. Georgia 30303 Denver, Colorado 8022 5

REGION 7Regional Direc.tot

AT%,,U.S. Fish and Wildlife Service

1011 E. Tudor RoadAnchorage, Alaska 950

TAKE PRIDEinAmerica

DEPARTMENT OF THE INTERIORU.S. FISH AND WILDLIFE SERVICE

As the Nation's principal conservation agency, the Department of the Interior has respon-sibility for most of our nationally owned public lands and natural resources. This includesfostering the wisest use of our land and water resources, protecting our fish and wildlife,preserving the, environmental and cultural values of our national parks and historical places,and providing for the enjoyment of life through outdoor recreation. The Department as-sesses our energy and mineral resources and works to assure that their development is inthe best interests of all our people. The Department also has a major responsibility forAmerican Indian reservation communities and for people who live in island territories underU.S. administration.

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