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BIOLOGICAL REPORT 82(10.108)

AUGUST 1985

HABITAT SUITABILITY INDEX MODELS

AND INSTREAM FLOW SUITABILITY

CURVES: CHUM SALMON

_ t:ic tl and Wildlife Service

SK ~. Department of the Interior

361

. U54

n o . 82-

10 . 108
This is one of the first reports to be published in the new "Biological

Report" series. This technical report series, published by the Research

and Development branch of the U.S. Fish and Wildlife Service, replaces

the "FWS/OBS" series published from 1976 to September 1984. The Biological Report series is designed for the rapid publication of reports with

an application orientation, and it continues the focus of the FWS/OBS

series on resource management issues and fish and wildlife needs.
MODEL EVALUATION FORM

Habitat models are designed for a wide variety of planning applications where habitat information is an important consideration in the

decision process. However, it is impossible to develop a model that

performs equally well in all situations. Assistance from users and

researchers is an important part of the model improvement process. Each

model is published individually to facilitate updating and reprinting as

new information becomes available. User feedback on model performance

will assist in improving habitat models for future applications. Please

complete this form following application or review of the model. Feel

free to include additional information that may be of use to either a

model developer or model user. We also would appreciate information on

model testing, modification, and application, as well as copies of modified

models or test results. Please return this form to:

Habitat Evaluation Procedures Group

or

Instream Flow Group

U.S. Fish and Wildlife Service

2627 Redwing Road, Creekside One

Fort Collins, CO 80526-2899

Thank you for your assistance.

Species

Habitat or Cover Type(s)

Geographic

Location

Type of Application: Impact Analysis Management Action Analysis

Baseline Other -------------------------

Variables Measured or Evaluated

Was the species information useful and accurate? Yes No

If not, what corrections or improvements are needed?------------
Were the variables and curves clearly defined and useful? Yes No

If not, how were or could they be improved?

Were the techniques suggested for collection of field data:

Appropriate? Yes No

Clearly defined? Yes No

Easily applied? Yes No

If not, what other data collection techniques are needed?

Were the model equations logical? Yes No

Appropriate? Yes No

How were or could they be improved?

Other suggestions for modification or improvement (attach curves,

equations, graphs, or other appropriate information)

Additional references or information that should be included in the model:

Model Evaluator or Reviewer Date ------------

Agency _

Address

Telephone Number Comm:----------- FTS _
Biological Report 82(10.108)

August 1985

HABITAT SUITABILITY INDEX MODELS AND INSTREAM FLOW

SUITABILITY CURVES: CHUM SALMON

by

Stephen S. Hale

Susitna River Aquatic Studies Program

Alaska Department of Fish and Game

333 Raspberry Road

Anchorage, AK 99502-2392

Thomas E. McMahon 1

Habitat Evaluation Procedures Group

Western Energy and Land Use Team

U.S. Fish and Wildlife Service

2627 Redwing Road

Ft. Collins, CO 80526-2899

and

Patrick C. Nelson

Instream Flow and Aquatic Systems Group

Western Energy and Land Use Team

U.S. Fish and Wildlife Service

2627 Redwing Road

Ft. Collins, CO 80526-2899

Western Energy and Land Use Team

Division of Biological Services

Research and Development

Fish and Wildlife Service

U.S. Department of the Interior

Washington, DC 20240

lPresent address: Department of Fisheries and Oceans, Pacific Biological

Station, Nanaimo, B.C. Canada V9R-5K6.
This report should be cited as:

Hale, S. S., T. E. McMahon, and P. C. Nelson. 1985. Habitat suitability

index models and instream flow suitability curves: Chum salmon. U.S.

Fish Wildl. Servo Biol. Rep. 82(10.108). 48 pp.
PREFACE

The Habitat Suitability Index (HSI) models and Instream Flow Suitability

Index (SI) curves presented in this publication aid in identifying important

variables that determine the quality of chum salmon habitat. Facts, concepts,

field data, and opinions obtained from published and unpublished reports are

synthesi zed and presented ina format that can be used for habi tat impact

assessment and development of management alternatives.

This report contains four major parts: (1) a Habitat Use Information

section which provides a general summary of the life history of chum salmon;

(2) a Specific Habitat Requirements section which provides a detailed discussion of habitat needs for each 1ife stage in freshwater; (3) a HSI Model

section which presents hypothesized relationships between various environmental

variables and habitat quality for chum salmon; and (4) an Instream Flow

Incrementa 1 Methodo logy (I FIM) section whi ch di scusses use of Sui tabi 1ity

Index Graphs from the HSI model with IFIM and provides additional SI graphs

developed specifically for analysis of chum salmon habitat via the IFIM.

Use of the HSI models with the Habitat Evaluation Procedures (HEP) or the

SI curves with the Instream Flow Incremental Methodology (IFIM) requires

project scoping, including the setting of clear study objectives. Armour

et al. (1984)1 present comparisons of the uses of HEP and IFIM for impact

assessment and helpful recommendations for selecting the method most appropriate for ach i evl nq study objectives. If HEP is to be used, HSI model

building techniques presented by the U.S. Fish' and Wildlife Service (1981)1

and the general guidel ines for modifying HSI model s presented by Terrell

et al. (1982)1 may be useful for simplifying and applying the HSI models to

specific chum salmon habitat assessment problems. Users of the SI curves for

IFIM analyses should be familiar with the guide to stream habitat analysis

(Bovee 1982)1 and the user's guide to the physical habitat simulation system

(Milhous et al. 1984).1

The HSI models and SI curves are hypotheses of species-habitat relationships, and users should recognize that the degree of veracity of the models,

curves, and assumptions will likely vary according to geographical area and to

the extent of the data base for each of the individual variables. The models

and curves have not been tested against field data. Therefore, the U.S. Fish

1Citations listed in REFERENCES section.

iii
and Wildlife Service encourages users of the models or curves to send comments,

suggestions, and field results that may help increase the utility and effectiveness of this habitat-based approach to impact assessment. Please send

comments to:

Habitat Evaluation Procedures Group or

Instream Flow and Aquatic Systems Group

Western Energy and Land Use Team

U.S. Fish and Wildlife Service

2627 Redwing Road

Fort Collins, CO 80526-2899

;v
CONTENTS

PREFACE iii

ACKNOWLEDGMENTS vi

HABITAT USE INFORMATION 1

Genera 1 1

Age, Growth, and Food 1

Reproduct ion 2

Rearing and Downstream Migration of Fry........................... 2

Specific Habitat Requirements..................................... 3

Upstream migration........................................... 3

Spawning..................................................... 5

Embryos and alevins 6

Rearing and downstream migration............................. 11

HABITAT SUITABILITY INDEX (HSI) MODELS................................. 13

Model Applicability.... 13

Geographi c a rea 13

Season 14

Verification level........................................... 14

Model Description................................................. 14

Overvi ew 14

Upstream migrant component 14

Spawning, embryos, and alevins component 15

Rearing and downstream migrant component..................... 16

Suitability Index (S1) Graphs for Model Variables.. 16

Use of the Mode 1 25

HSI determi nat ion ;.................... 25

Field application of the model............................... 25

Model Interpretation and Limitations.............................. 25

ADDITIONAL HABITAT MODELS 26

INSTREAM FLOW INCREMENTAL METHODOLOGY 26

Suitabil ity Index Curves as Used in I FIM 26

Avai 1abi 1ity of SI Curves for Use in IFIM 27

Spawning migration........................................... 27

Spawn i ng and egg i ncubat i on 27

Effecti ve spawn i ng habitat................................... 34

Fry 36

REFERENCES 37

II
ACKNOWLEDGMENTS

We thank C. Burger, U.S. Fish and Wildlife Service and J. Helle, U.S.

National Marine Fisheries Service, for review comments which improved the

quality and accuracy of this report. Helpful comments were also provided by

various staff of the Susitna River Aquatic Studies Program of the Alaska

Department of Fish and Game. The reader should not construe reviewer

assi stance as an endorsement of the Suitabi 1i ty Index curves and habitat

models, which are an independent synthesis and interpretation of the literature. T. Rosenthal provided editorial assistance and J. Terrell coordinated

manuscript production. Word processing was by C. Gulzow, D. Ibarra, and

E. Graf. The cover figure was drawn by S. Donovan, Alaska Department of Fish

and Game, based on a photograph by J. Helle.

vi
CHUM SALMON (Oncorhynchus keta)

HABITAT USE INFORMATION

Genera 1

Chum salmon have the widest distribution of the Pacific salmons, occurring

in the northern North Pacific Ocean, the Bering Sea, the Chukchi Sea, and

along the Arctic Ocean coasts of Siberia, Alaska, and northwest Canada (Bakkala

1970). Spawning occurs in North America from the Sacramento River in

California to the MacKenzie and Anderson Rivers on the Arctic coast of Canada

(Hart 1973), and in Asia, from the Nakdong River in Korea and the Tone River

in Japan to the Lena River in Siberia (Bakkala 1970). Spawning has been

recorded in at least 1,270 streams in the United States (Atkinson et al.

1967). Along the North American coast, chum salmon populations are relatively

small south of central Oregon or north of Kotzebue Sound, Alaska (Helle 1979).

Detailed descriptions of their distribution in Alaska were given by Alaska

Department of Fish and Game (1978) and Hale (1981).

Summaries of the life history and freshwater habitat requirements of chum

salmon were presented in Bakkala (1970) and Hale (1981). The freshwater

habitat requirements for Pacific salmon in general have been summarized by

Nicola et al. (1966) and Reiser and Bjornn (1979).

Age, Growth, and Food

Chum salmon typically mature at ages III or IV in Oregon and Washington

and III, IV, or V in British Columbia and Alaska (Bakkala 1970; Fulton 1970;

Helle 1979). Mature fish in North America generally range from 53 to 92 cm

long and weigh 0.8 to 13.4 kg (Bakkala 1970).

Some chum sa 1mon fry mi grate seaward soon a fter emergence from redds

(Hoar 1953; Neave 1966), whereas others remain in freshwater or intertidal

areas for as long as several weeks to feed (Sparrow 1968; Bakkala 1970; Scott

and Crossman 1973; Mason 1974). Fry remaining in freshwater feed on drifting

terrestrial insects or aquatic forms such as immature chironomids, blackflies,

caddisflies, mayflies, and stoneflies (Disler 1953; Sparrow 1968; Bakkala

1970). Those that stay in intertidal reaches feed on amphipods, copepods, and

insects (Mason 1974).

Growth in freshwater is negligible in fry that migrate to sea soon after

emergence, but may be considerable for those that remain in freshwater for

several weeks. Chum fry in the Tokachi River of Japan showed up to a 30%

1
increase in length and up to a 310% increase in weight during April and May

(Kaeriyama et al. 1978). Freshwater feeding and growth of chum salmon fry

have also been demonstrated in the Satsunai River (Kaeriyama and Sato 1979)

and in the Yurappu River (Kobayashi and Abe 1977). In the Susitna River,

Alaska, emerging chum salmon fry that were marked with a coded wire tag and

subsequently recaptured, showed a steady increase in length from a mean of

40 mm to a mean of 49 mm, 22 to 28 days later, an increase in length of 22.5%,

which would correspond to an even larger percentage increase in weight (Roth,

pers. comm.). Many chum fry from the upper reaches of the spawni ng areas in

the Susitna River move downstream at lengths not much longer than their

emergence length « 40 mm), but there are also many that spend several weeks

in freshwater and attain lengths of > 60 mm, an increase of over 20 mm.

Reproduction

The timing of the return of adult chum salmon to natal streams to spawn

varies with location. Summer runs are characteristic in Alaska and other

northern parts of the range, spawning in these areas occurring in August and

September (Alaska Department of Fi sh and Game 1978; Morrow 1980). In lower

southeast Alaska and southward, fall spawni nq runs of chum salmon are more

common (Hunter 1959; Neave 1966; Ful ton 1970). Chum salmon may spawn over a

prolonged period; both summer and fall runs occur in many streams, particularly large rivers. In the Yukon and other large river systems, fish in the

summer run spawn in lower sections of the river and those in the fall run

spawn in areas much farther upstream (Scott and Crossman 1973).

Spawning occurs primarily in short coastal streams and in the lower

200 km of large rivers (Neave 1966; Fulton 1970), but chum salmon migrate

upstream over 1900 km in the Amur River (Bakkala 1970) and over 2,500 km in

the Yukon River (Bakkala 1970; Morrow 1980). Like pink salmon (Q. gorbuscha),

chum salmon frequently spawn in intertidal areas. In general, they are less

likely to surmount waterfalls and other barriers than other salmon (Neave

1953, 1966; Bakkala 1970; Thorsteinson et al. 1971).

Eggs incubate in gravel redds 50 to 130 days (depending on temperature)

before hatching. Newly hatched fry (alevins) remain in the gravel until

spring, when their yolk sacs are nearly or completely absorbed (Bakkala 1970;

Scott and Crossman 1973). Mortality during incubation is high; survival from

egg deposition to fry emergence typically averages < 10% (Hunter 1959).

Rearing and Downstream Migration of Fry

Fry emerge from the gravel in March to May (Sano 1966; Bakkala 1970).

Timing of downstream migration of chum fry in Carnation Creek, British Columbia

over a 13-year period (1971-1983) is presented in Andersen (1983, 1984).

After leaving the gravel, many fry soon migrate downstream to the ocean (Hoar

1956; Neave 1966) but, in some systems at least, a considerable number remain

in streams [Scott and Crossman 1973; Kobayashi and Abe 1977; Roth et al. 1984;

personal observation (McMahon) in Carnation Creek, British Columbia] and

intertidal areas (Mason 1974) for at least several weeks. In the Fraser River

2
system, Beacham and Starr (1982) found that 80~6 of the downstream migrants

were < 40 mm FL, but that the proportion of fry> 40 mm increased with t-ime,

suggesting that later migrants had spent more time feeding in freshwater.

Mason (1974) found that the fry have complex behavioral responses that delay

seaward movement and allow the fish to feed in freshwater and intertidal areas

before entering the ocean. It is unknown if those fry that rear in freshwater

for longer periods have higher survival rates than those that migrate to the

ocean soon after emergence, as found for coho salmon (0. kisutch) fry (Crone

and Bond 1976). -

Chum salmon fry migrate downstream primarily at night (Andersen 1983), and

hide in the streambed (Neave 1966) or feed (McMahon personal observation)

during the day. In large, turbid rivers at more northern latitudes where

there is 1ittle darkness in June, such as the Susitna River in south-central

Alaska, migration occurs during the day as well (Roth et al. 1984). Preference

for higher salinities increases as the fry grow (Houston 1961; McInerney

1964), and migration to seawater within the first summer is considered

mandatory for survival (Houston 1961). Environmental factors that control

downstream migration include temperature, photoperiod, light intensity, size

of fish, and level of river discharge (Brannon and Salo 1982).

Specific Habitat Requirements

Tolerances and requirements of chum salmon change with season and age.

Although most developmental changes and movements to different habitats are

gradual, factors assumed to affect habitat qual ity can be specified most

effectively if the freshwater life cycle is divided into three distinct stages:

(1) upstream migrants - sexually mature chum salmon migrating from the ocean

upstream to spawn; (2) spawning, embryo, and alevins (yo l k-s ac fry) - period

from egg deposition to hatching and emergence of fry from redds; and (3) downstream migrant - young fish from the time of emergence from gravel to entry

into the ocean.

Upstream migration. Adult chum salmon must arrive at the spawning grounds

within a relatively restricted seasonal period and in good health if spawning

is to be successful. The habitat needs are relatively broad during this

phase, which lasts for only a few weeks or less for stocks that spawn in short

coastal streams or within a few kilometers from the stream mouth. This phase

lasts for a month or more for stocks that spawn upstream in larger rivers such

as the Yukon.

Temperature can adversely affect migrating salmon by altering the timing

of migration and rate of maturation, by increasing the susceptibility to

diseases, and by causing direct mortality (Holt et al. 1975; Reiser and Bjornn

1979). Chum salmon usually enter streams as temperatures fall below 15° C

(Bakkala 1970), and most upstream migration occurs in the range of 8 to 14° C

(Hunter 1959; Sano 1966; Barrett et al. 1984). In Alaska, the observed

temperature range during upstream migration was 5.0 to 12.8° C in the Kuskokwim

River (Alaska Department of Fish and Game 1980), 10.0 to 16.7° C in the Anvik

River (Trasky 1974), and 8.9 to 14.4° C (during peak migration) in the Traitors

River (Mattson and Hobart 1962). Bell (1973) suggested that the temperature

3
range of 8.3 to 15.6° C would allow successful upstream migration of chum

salmon, and that temperatures near 10° C were optimum. Temperatures above

about 15° C are likely to result in increased susceptibility to disease

(Wedemeyer 1970; Fryer and Pilcher 1974; Holt et al. 1975; Groberg et al .

1978) and to delay upstream migration (Bell 1973). Temperatures 25.5° Care

lethal to migrating Pacific salmon (Bell 1973). The effects of low

temperatures are less clear. Although most migration occurs above 7° C,

Mattson and Hobart (1962) observed upstream migration at temperatures as low

as 4.40 C; it is not known if upstream migration occurs at even lower

temperatures.

Dissolved oxygen (DO) levels> 6.3 mg/l are recommended for successful

upstream migration of anadromous salmonids (Davis 1975). Lower DO concentrations are likely to inhibit upstream movement by reducing swimming ability and

eliciting avoidance responses. Davis et al. (1963) found that the maximum

sustained swimming speed of coho salmon was sharply reduced at DO levels

< 6.5 mg/l. Whitmore et al. (1960) reported that juvenile coho salmon avoided

waters where DO was < 4.5 mg/l. About 300 pre-spawn pink and chum salmon died

in Porcupine Creek, southeastern Alaska, in 1981 when their upstream migration

was blocked by low flows, crowding them into a pool where the DO level dropped

to < 2.0 mg/l (Murphy 1985). Edgington (pers. comm. in Krueger 1981) reported

high mortalities of ripe pink salmon in southeast Alaska streams where DO had

dropped below 4 mg/l.

Chum salmon are less inclined to surmount rapids, waterfalls, and other

barriers than other species of Pacific salmon; thus, accessibility of the

spawning stream is probably a very important variable determining the success

of upstream migration. The maximum height of surmountable barriers for chum

salmon has not been determined.

Low flow can also be a barrier. Adult chum salmon travel upstream in

shallow riffles with the upper part of their bodies above water (personal

observation, Hale), but the distance that can be traversed in such shallow

water is unknown. In southeast Alaska, where the streams without a snow pack

generally have low reservoir capacity and the flow depends heavily on rainfall,

migrating chum salmon often have difficulty moving upstream during dry periods.

A similar situation has been reported in British Columbia (Wickett 1958).

High mortality of pre-spawn salmon caused by high water temperatures and low

DO levels during low flow conditions is not uncommon in southeastern Alaska;

up to 30,000 pink and chum salmon died in one stream (Murphy 1985). Thompson

(1972) suggested that the maximum water velocity for successful upstream

migration of adult chum salmon was 2.44 m/sec and that the minimum water depth

required was 18 cm. Blakely et al. (1985), who observed successful and

unsuccessful attempts by chum adults trying to reach spawning grounds at 85

passage reaches in the Susitna River, stated that the threshold depth for

successful passage was 15.2 cm. This depth increases as the passage reach

becomes longer than one or two meters.

The timing of upstream migrations is often correlated with flow conditions. Lister and Walker (1966) observed chum salmon delaying at the mouth of

the Big Qualicum River, British Columbia, during extremely low discharges and

then moving upriver when discharge increased. Similar observations have been

4
made in Carnation Creek, British Columbia (Scrivener, pers. comm.) and in the

Chitose River, Japan (Mayama 1978). High levels of discharge can also slow

upstream migration. Barrett et al. (1984) observed chum salmon milling in the

Susitna River during discharge peaks when the catch rate in fish wheel s

declined. High suspended sediment loads may also inhibit upstream migration.

In Prince William Sound, chum salmon did not spawn in a glacier-fed stream

until after the flow and silt load had decreased (Helle 1960). In the Delta

River, Alaska, chum salmon did not enter the stream until after freeze-up when

glacial run-off had ended and the only source of water was upwelling groundwater (Geiger, pers. comm. in Hale 1981).

Spawning. Temperature, water velocity, substrate composition, and

salinity appear to be the major factors determining the suitability of an area

for spawning. Chum salmon spawn at temperatures of 5.1 to 6.7° C in the

tributaries of the Columbia River; at 4.0 to 16.0° C for British Columbia in

general (Neave 1966); at a mean temperature of 12° C in Hooknose Creek, British

Columbia (Hunter 1959); at s 13° C in Sashin Creek, southeast Alaska (McNeil

1964); at 6.5 to 9.0° C in the Kizhuyak River and 9.0 to 12.5° C in the Terror

River on Kodiak Island (Wilson et al. 1981); and at about 4.5 to 13.0° C in

the Susitna River (Vincent-Lang et al. 1984). Schroder (1973) noted that

there was some inhibition of· spawning by chum salmon in Washington at temperatures < 2.5° C. Bell (1973) recommended 7.2 to 12.8° C as optimum for

successful spawning.

Chum salmon spawn over a wide range of water velocities; surface water

velocity may be less important than the presence of upwelling groundwater

(Kogl 1965; Sano 1967; He 11 e 1979; Vi ncent-Lang et a 1. 1984). Surface water

velocities (em/sec) measured at spawning sites in different areas were: 0 to

60 in the Chena River, Alaska (Kogl 1965); 0 to 118.9 in the Terror and

Kizhuyak Rivers (Wilson et al. 1981); a range of 46 to 97 (Thompson 1972) and

a mean of 73 em/sec over 214 redds in Oregon (Smith 1973); and 21 to 101

(12 em above chum salmon redds) in Washington (Collings 1974). The range of

water velocities at 333 chum salmon redds in the Susitna River, Alaska,

examined by Vincent-Lang et al. (1984) was from 0 to about 130 em/sec; the

velocity at most redds ranged from 0 to 30 em/sec. In a spawning channel in

Washington, velocities ranging from 0 (at high tide) to 22.5 em/sec had no

apparent effect on spawning behavior (Schroder 1973). In the Memu River,

Japan, redds were most abundant in areas where the velocity was 15 to 20 em/sec

(Sano and Nagasawa 1958). Sano (1967) stated that chum salmon spawn in water

with surface velocities of less than 15 em/sec provided there is groundwater

seepage through the gravel. Smith (1973) recommended velocities of 46 to

101 em/sec for successful spawning of chum salmon in Oregon. A chum salmon

spawning channel at Jones Creek, British Columbia, was designed for velocities

of 31 to 76 em/sec (MacKinnon et al. 1961).

Tautz and Groot (1975) suggested that chum salmon prefer to spawn in

areas where rocks protruding above the substrate create an upwelling, accelerating current. The current at the boundary between pools and riffles is also

favorable. Such irregularities are conducive to a good flow of intragravel

water (Reiser and Bjornn 1979). Streamflow during spawning, relative to the

flow during the incubation period, can be an important factor affecting

5
survival. Eggs deposited in shallow water during high flow stages are subject

to desiccation or freezing during low flow stages. Kogl (1965) found that

chum salmon dug redds in the Chena River at water depths of 5 to 120 cm; the

fish spawned at the greater depths (> 100 cm) in areas where upwelling spring

water or groundwater seepage was lacking. Eggs deposited in areas where the

water was < 100 em deep and where there was no upwelling were subject to

freezing in winter. Chum salmon spawned at depths of 61 to 92 cm in the Delta

River (Francisco 1976) and 7.6 to 106.7 cm in the Terror and Kizhuyak Rivers

(Wilson et al. 1981). In the Susitna River, water depths at chum salmon redds

ranged from about 5 to 90 cm; most were in the range of 10 to 60 cm

(Vincent-Lang et al. 1984). Water depths over chum salmon redds were 5.1 to

76.2 cm (mean 25 cm) in tributaries of the Columbia River (Burner 1951);

averaged 30 cm in five Oregon streams (Smith 1973); and ranged from 15 to

53 em in Washington streams (Collings 1974). A minimum water depth of 18 cm

was recommended by Thompson (1972) and Smith (1973).

Chum salmon excavate redds in gravel beds with particles of 2 to 3 cm in

diameter, but they also construct redds in substrates with larger rocks,

including bedrock covered with small boulders (Scott and Crossman 1973; Morrow

1980). Duker (1977) reported that spawning females given a choice of

substrates preferred particle sizes 0.7 to 7.6 cm in diameter over larger or

smaller ones.

Salinity may interfere with fertilization of eggs spawned in the intertidal zone. Rockwell (1956) found that salinities of 18 ppt inhibit fertilization of chum salmon eggs, although some fertilization occurred in salinities

as high as 30 ppt.

The outlet of a tidal lagoon in Prince William Sound, Alaska, drained

saline water after the 1964 earthquake that uplifted

Salinity of the outlet stream ranged from 4 to 8 ppt.

spawned in this stream and redd excavations yielded no

had not imbibed water to separate the membranes, i.e.,

har-den" (Helle, pers. comm.).

the a rea about 6 ft.

Chum and pink salmon

live eggs. The eggs

they di d not "waterEmbryos and alevins. Temperature, streamflow, dissolved oxygen, salinity,

and especially substrate composition are important factors affecting survival

of chum salmon embryos and alevins. Stream water temperatures from 0° C to as

high as 15° C have been observed during incubation (McNeil 1966; Koski 1975).

Intragravel temperatures in chum salmon redds in the Susitna River, Alaska,

ranged from 12.5° C in September to 0.0° C in December (Vining et al. 1985).

Chum salmon prefer to spawn in upwelling areas of the Susitna, where intragravel temperatures remain about 3 to" 4° C all winter. Embryos in redds where

the temperature was 1ess than 3° C hatched two to three months 1ater than

those in the upwelling areas (Vining et al. 1985). In northern Japan, Sano

(1966) found that fall chum salmon preferred to spawn in spring areas where

water temperatures did not decrease below 4° C. In Alaska, observed intragravel temperatures during incubation were 0.5 to 4.5° C in the Chena River

(Kogl 1965); 6.1 to 10.0° C in early October to 0.2 to 3.3° C in March in the

Noatak River (Merritt and Raymond 1982); 0.6 to 6.7° C in the Delta River

(Francisco 1977); and 4° C in October, near 0° C in December-January, and 1° C

6
in mid-Ma rch before fry emergence in 01 sen Creek (Ba i 1ey 1964). Long-term

exposure to temperatures < 4.40 C reduces survival of chum salmon embryos

(Schroder 1973; Koski 1975; Wangaard and Burger 1983; Alaska Department of

Fish and Game 1983). Schroder et al. (1974) reported that the percent survival

to emergence of chum salmon in Washington was decreased if the embryos were

exposed to water temperatures < 1.50 C during the early stages of embryonic

development. In a hatchery in Alaska, Raymond (1981) found that mortality

(fertilization to emergence) was higher (38%) when incubation temperatures

were 2.0 to 4.20 C than when they were at 3.6 to 4.50 C (18%). Wangaard and

Burger (1983), in a laboratory experiment, observed an increase in mortalities

and abnormalities in Susitna River chum embryos and alevins when the average

incubation temperature was < 3.40 C. Kirkwood (1962) found that survival of

Prince William Sound chum salmon embryos was higher in the upper intertidal

zone than farther upstream, due to the higher intertidal temperatures.

Similarly, Hartman and Leahy (1983) found that intragravel temperatures in

chum spawning areas in British Columbia during winter were up to 20 C higher

than stream temperatures and that these di fferences were most pronounced in

the upper intertidal zone. Koski (1975) found that exposure of Washington

chum salmon embryos to water temperatures < 4.40 C before gastrulation caused

high mortalities. McNeil and Bailey (1975) suggested that temperatures in

Pacific salmon hatcheries should be maintained above 4.40 C for at least 10

days, and preferably for 20 to 30 days, after fertilization. After this

period, Bailey and Evans (1971) found that pink salmon embryos can tolerate

temperatures as low as 00 C, provided that freezing does not occur.

Water temperatures can al so cause significant changes in the time of

emergence of fry from the substrate. Koski (1975) found that annual water

temperature variations in a Washington stream caused the time required to

reach 50% emergence of chum fry to vary as much as 30 days duri ng a 3-year

period. The time to hatching of fry is about 50 days at 7.0 to 15.00 C, but

130 days at 0.0 to 5.00 C (Bakkala 1970). Raymond (1981) reported that chum

salmon eggs taken from the Delta River and incubated at 2.0 to 4.20 C in a

hatchery emerged from the gravel 7 weeks later than fry from the same stock

exposed to a higher mean temperature (3.90 C) in the natural spawning area.

Incubation of chum salmon embryos was studied at three sloughs with upwelling

ground water (3.5 to 4.00 C) in the Susitna River (Alaska Department of Fish

and Game 1983). One of these sloughs was flooded with cold (- 00 C) mainstem

water when an ice jam caused the head of the slough to be overtopped. Chum

embryos and alevins in this slough experienced a higher mortality rate and a

delayed emergence as compared to the other sloughs which remained at 3.5 to

4.00 C. In a controlled laboratory experiment, Wangaard and Burger (1983)

found that complete yolk sac absorption of Susitna River chum fry was delayed

by up to 2 months when the average incubation temperature was 2.1 0 C, as

compared to 3.9 and 4.00 C. Kirkwood (1962) stated that chum salmon embryos

in Pri nce Wi 11 i am Sound that had not hatched before January or February di d

not survive. Clear-cut logging in the Carnation Creek, B.C., watershed

resulted in a 20 C rise in winter stream temperatures (Holtby and Newcombe

1982; Hartman et al. 1982; Holtby, pers. comm.) and a shift in chum fry migration of up to 6 weeks earlier (Andersen 1983, 1984). Since ongoing work

indicates that emergence timing in chum salmon is genetically controlled, it

is possible that earlier emergence of Carnation Creek chum salmon could have

7
significant, deleterious effects on early fry survival in the ocean and may be

a factor in the declines in adult returns to Carnation Creek that occurred

after logging (Holtby, pers. comm.).

Because chum salmon frequently spawn in streams where runoff is highly

variable, embryo mortality during freshetting may be severe (Hunter 1959;

McNeil 1966; Neave 1966). High flows cause increased mortality by directly

scouring redds or by depositing fine sediments over the redds which, in turn,

reduces substrate permeability and inhibits emergence of fry (Neave 1953;

Wickett 1958; McNeil 1966, 1969; Scrivener and Brownlee 1982, in prep.).

Conversely, low flows can lead to desiccation of eggs, low DO, high temperatures, or freezing (Neave 1953; Levanidov 1954; Wickett 1958; McNeil 1966;

Sano 1966; Vining et al. 1985). McNeil (1966, 1969), who observed that

survival to emergence varied in relation to stream discharge during the winter,

found that mortality of chum and pink salmon eggs and alevins was four to five

times greater in a stream with a 500-fold difference in average daily stream

discharge (0.11 to 55.0 m3/sec) than in a nearby stream where this difference

was only 80-fold (0.34 to 28.0 m3/sec). Cederholm and Koski (1977) reported

that 55% of chum salmon redds in a channelized section of a Washington stream

were scoured out or covered by sediment due to streambed shifting during

frequent and severe freshets. Also, Lister and Walker (1966) found an inverse

relation between percent survival of chum salmon eggs to the fry stage and the

peak daily discharge of the Big Qualicum River, British Columbia. On the

Noatak River, low adult returns appear to be correlated with high water levels

(Bird, pers. comm. in Hale 1981). Gallagher (1981) found that low flows

during the incubation period were correlated with good adult returns for

streams in the Puget Sound area. Beacham and Starr (1982) determined that

freshwater survival (based on fry escapement) from the Fraser River system was

due to interactions of temperature, rainfall (= flow), and egg abundance.

Survival was highest in years when winters were dry and warm and lowest in

years when winters were cold and wet. On the other hand, Wickett (1958)

showed a direct relationship between an index of production and discharge for

Hooknose Creek in November. In summary, the effect of varying discharge is

not constant between streams and may depend on other conditions in the selected

stream.

The size and shape of substrate particles influence many factors that are

important to the successful incubation of embryos and alevins. Some of these

factors -- several of them interrelated--are permeability, porosity, flow of

intra-gravel water, dissolved oxygen concentration, concentration of waste

metabolites such as carbon dioxide and ammonia, the "armoredness" or resistance

to abrasion of the substrate surface, and the degree of embeddedness of large

particles in the substrate surface. Ideal substrate conditions for incubation

of eggs and alevins of chum salmon are hard to specify because different

investigators have used various methods and definitions, making comparisons

difficult. Hunter (1959) found that chum salmon in Hooknose Creek, B.C.,

spawned in gravel 13 to 130 mm in diameter that contained unspecified amounts

of sand and silt. Sano (1959) reported that chum salmon redds in Japan

consisted of > 30% gravel (particle diameter> 31 mm). Burner (1951) wrote

that redds in the Columbia River were composed of 81% gravel (particles

< 152 mm in diameter but larger than sand); 6% mud, silt, and sand; and 13%

8
cobble (~ 152 mm diameter). The substrate in chum salmon redds sampled in the

Susitna River, Alaska, was mainly composed of particles 2 to 76 mm in diameter;

particles < 2 mm comprised 6 to 25% of the total (Vining et al. 1985). Percent

survival of embryos in situ Whitlock-Vibert boxes was zero at percent fines

> 18%.

Sedimentation during the incubation period can be a major source of egg

mortality (Neave 1953; Levanidov 1954; Wickett 1954; McNeil 1966; Rukhlov

1969; Scrivener and Brownlee 1982, in prep.). In one year in the early 1950 1s,

siltation in the River Khor in Siberia killed all of the embryos in 42 chum

salmon redds (Levandov 1954). However, inasmuch as chum salmon often spawn in

areas of upwelling ground water, they may be able to tolerate a higher

percentage of fines if intra-gravel flow is maintained by such upwelling (Sano

and Nagasawa 1958; Helle, pers. comm. in Hale 1981; Vining et al. 1985).

Di 11 and Northcote (1970) found that survi va1 to emergence for chum

salmon in experimental containers was nearly 100~6 in large gravel (51 to

102 mm) but only 31% in small gravel (10 to 38 mm). Koski (1975) found a

highly significant inverse relation between percent sand (fines < 3.327 mm but

> 0.105 mm) and percent survival to emergence of chum salmon in Washington.

Increased percent sand also resulted in earlier emergence, reduced yolk conversion efficiency, and smaller fry. Furthermore, 45~6 silt (fines < 0.105 mm)

reduced survival; this effect was masked by a changing percentage of sand.

Koski also observed a significant inverse relation between DO concentration

and percent sand and silt.

Scrivener and Brownlee (1982, in prep.) reported that after intense

logging in the Carnation Creek watershed, British Columbia: (1) fine sand

particles significantly increased; (2) intragravel DO decreased from a mean of

6.8 mg/l prior to logging to 3.2 mg/l after logging; (3) gravel permeability

declined 50 to 68% to levels below the threshold cited as limiting chum fry

production (3,000 cm/hr); and (4) chum salmon egg-to-fry emergence rates

ranged from 16.3 to 25.0% (x = 20.95%) in the 6 years before logging to 4.6 to

23.5% (x = 12.3%) in the 5 years after logging.

Rukhlov (1969) found that egg mortality of chum salmon on Sakhalin Island

increased as sand content exceeded 14%. He found that average survival rates

were 50% in substrates with> 22% sand, 65% in those with 18% sand, and 85%

in those with 10% sand. Thorsteinson (1965) stated that spawning grounds in

which the percent fines (particles < 0.833 mm) exceeded 12.7 are of poor

quality because permeability is greatly reduced. Johnson (1980) showed experimentally that the permeability of spawning gravels decreased as percent fines

(particles < 0.5 mm) increased. Wickett (1958) related the average survival

of chum and pink salmon eggs and alevins to the permeability of the substrate:

percent survival (eggs in female to fry escapement) was only 1.2% at a

permeability of 1,914 cm/hr but 7.6% at a permeability of 4,035 cm/hr.

Substrate sizes (diameter of particles) suggested for successful spawning and

incubation of chum salmon were 13-152 mm (McNeil and Bailey 1975) and 13-102 mm

(Bell 1973).

9
DO affects the survival, fitness, and timing of emergence of chum salmon

embryos and fry (Wickett 1954, 1958; Alderdice et al. 1958; Koski 1975). The

level of intragravel DO is a function of sediment particle size, rate of

intragravel flow, and DO concentration in the free flowing water (Daykin 1965;

Koski 1975). Critical oxygen levels (the lowest concentration at which

respiration demands are satisfied) for chum salmon eggs at 8 to 10° C ranged

from 0.72 mg/l for early developmental stages to 7.19 mg/l just before hatching

(Wickett 1954; Alderdice et al. 1958). Wickett (1954, 1957) reported that a

DO concentration of 2 mg/l at 5° C was lethal, but Levanidov (1954) and Kogl

(1965) found good survival rates at 2.0 mg/l in areas where springs and upwelling groundwater provided good intragravel flow. Koski (1975) found a

significant correlation between percent survival to emergence of chum salmon

and DO concentration with survival decreasing markedly below 3.0 mg/l. An in

situ incubation experiment using chum salmon eggs in 59 Whitlock-Vibert boxes

Trlartificial redds in the Susitna River, Alaska, resulted in good survival

rates where the intragravel DO level was> 3.0 mg/l and zero survival where DO

was < 3.0 mg/l (Vining et al. 1985). Redds in intertidal areas may be exposed

to low DO levels during tidal cycles; during high tide in Prince William

Sound, Thorsteinson (1965) observed a drop from 7 mg/l to 4.5 mg/l. Reduced

water permeability at one tide level in the O'l scn Creek intertidal spawning

area led to a low DO level (3.6 mg/l) and a low percent survival to emergence

(Bailey 1964). Long dry periods during which stream and intragravel flows are

reduced can also result in low DO. Such conditions in the Traitors River,

Alaska, led to high mortality of embryos at an intragravel DO level of 1.8 mg/l

(Mattson et al. 1964).

Low DO causes reduction in the size of emerging fry and delays hatching

and emergence. Kogl (1965) reported that the dry weight of chum salmon alevins

in the Chena River, Alaska, decreased from 2.3 mg at 4.5 mg/l DO (average) to

0.1 mg at 2.1 mg/l DO (average). Alderdice et al. (1958) reported that early

developmental stages (embryos < 12 days after fertilization) could survive 7

days of exposure to 0.3 mg/l DO but that a hi gh percentage of abnorma 1iti es

resulted; later developmental stages (> 22 days) did not survive. Furthermore,

short-term (7 day) exposures to DO levels below saturation (at 10° C) caused

delays in hatching in early and middle developmental stages but caused

premature hatching when the exposure to low DO occurred just before hatching.

Koski (1975) observed similar delays in emergence of fry when DO was < 3 mg/l.

Delays in emergence could place fry in unfavorable conditions in relation to

food availability, predation, temperature, or streamflow.

Survival of chum salmon embryos seemingly decl ines from the upper to

lower reaches of intertidal zones, Rresumably as a result of higher salinity

(Bailey 1964; Thorsteinson 1965; Thorsteinson et al. 1971). Thorsteinson

(1965) found that egg-to-fry survival was 40 to 50% in the upper section of

the intertidal zone in Prince William Sound and 10 to 15% in the middle

section; the lower section was not used by chum salmon. In laboratory experiments, Rockwell (1956) found that: (1) constant salinities s 6 ppt had no

effect on early developmental stages of chum salmon eggs; (2) significantly

retarded development and 67% mortality occured at 11.6 ppt; and (3) > 12 ppt

led to 100% mortality. Kashiwag; and Sato (1969) found that percent mortality

of chum salmon eggs to hatching was 0 at s 9 ppt, 25 at 18 ppt, 50 at 27 ppt,

10
and 75 at 35 ppt; however, nearly all alevins reared from eggs held at > 9 ppt

died within a few days after hatching. Alevins that were hatched in freshwater

and then exposed to a seawater mixture survived constant salinities of < 9 ppt

but died at higher salinities (Kashiwagi and Sato 1969). Duration of ~xposure,

temperature, dissolved oxygen, and stage of development confound these effects

of salinity on chum salmon eggs and alevins (Rockwell 1956; Thorsteinson 1965;

McNeil 1966). Also, recent observations by Hartman (pers. comm.) in Carnation

Creek, B.C., support McNeil and Bailey's (1975) speculation that Pacific

salmon eggs can tolerate periodic inundations of salinities of 15 to 30 ppt;

Hartman found viable chum eggs in the Carnation Creek intertidal zone 200 m

below the high tide mark where intragravel salinities ranged from 0 to 24 ppt,

depending on stage of the tide.

The presence of upwelling ground water can be an important factor in chum

salmon egg to fry survival (Kogl 1965; Vining et al. 1985). Upwelling ground

water prevents dewatering and freezing when surface flows are low, maintains a

more constant intragravel temperature, and supplies dissolved oxygen. Other

factors, such as pH, CO 2 , alkalinity, ammonia, and H2S, may affect survival to

emergence in altered systems, but the tolerances of chum salmon eggs and

alevins to these factors are largely unknown (see Hale 1981:43-44). The pH of

intragravel water at chum salmon redds was 6,5 in the Chena (Kogl 1965) and

Amur (Levanidov 1954) rivers and 6.3-6.5 in a Hokkaido stream (Kobayashi

1968). McNeil and Bailey (1975) suggested that pH levels of 6.0 to 8.0 are

desirable for Pacific salmon.

Rearing and downstream migration. Habitat requirements of fry from the

period of emergence from the gravel to entrance into the sea center on favorable temperatures, cover from predators, availability of food, and

(eventually), availability of seawater. Smoltification in chum salmon has not

been studied extensively, and it is not clear how environmental factors in

freshwater affect this process and affect the survival of young chum salmon in

estuaries and in the ocean. However, information on other anadromous salmonids

summarized by Wedemeyer et al. (1980) indicated that this stage is particularly

sensitive to environmental stresses. Inferences made here about the factors

that may adversely affect smoltification in chum salmon fry were drawn

primarily from the review of salmonid smoltification by Wedemeyer et al.

Temperature can affect the timing of downstream migration, as well as the

growth, survival, and smoltification of chum salmon fry. The recommended

temperature range for successful downstream migration is 6.7 to 13.30 C (Bell

1973). In Alaska, out-migration of chum salmon fry was observed at temperatures of 3.0 to 5.50 C in the Delta River (Raymond 1981), 0.1 to 14.50 C in

the Susitna River (peak outmigration began at temperatures> 6.0 0 C; Roth

et al. 1984), and 5.0 to 7.0 0 C in the Salcha River during the peak migration

(Trasky 1974). In Hokkaido, Japan, downstream migration began at temperatures

of 2.0 to 3.00 C and peaked in May when temperatures were 6.0 to 10.00 C (Sano

1966). Semko (1954) measured temperatures of 4.6 to 5.50 C during downstream

migration of chum salmon fry in April and May in the Bolshaia River, USSR.

Temperatures > 150 C, which chum salmon fry generally avoid (Brett 1952;

Levanidov 1954), may cause premature downstream migration. Keenleyside and

Hoar (1955) reported that chum fry changed from positive to negative rheotaxis

11
when temperatures rose above 10° C; Mihara (1958) observed this effect at

temperatures above 15° C. The lower and upper 1etha 1 temperatures for chum

salmon fry are near 0° C (Brett 1952; Brett and Alderdice 1958) and 23.8° C

(Brett 1952). McNeil and Bailey (1975) stated that Pacific salmon fry grow at

temperatures of 4.4 to 15.7° C, but that the ideal range is 10.1 to 12.9° C.

Growth is slow at temperatures below 10° C and problems with diseases and

other stress factors develop at 15.7° C or higher; temperatures of 20 to 22° C

are tolerated only for limited periods. Wedemeyer et al. (1980) reported that

elevated water temperatures during incubation accelerate the onset of

smoltification and shortens the smolting period, and thus seaward migration of

salmonid smolts may occur at a time when conditions are unfavorable; they

recommended that temperatures should not exceed 10° C in late winter (to

prevent accelerated smolting) and should not exceed 12° C during emergence and

seaward migration in the spring (to prevent alteration of the smoltification

process and reduce the risk of disease).

The effects of low temperatures on chum salmon fry during rearing and

downstream migration are not well documented. However, temperatures < 8° C

from March through June may render a stream less suitable as potential rearing

habitat. Levanidov (1955) found that feedi ng and growth of the fry were

reduced at temperatures:::; 8° C in the laboratory. Similarly, Kobayashi and

Ishikawa (1964) found that chum salmon fry grew slowly in March-April, the

peri od duri ng whi ch most of the fry migrated to the sea, but that growth was

more rapid after April because feeding increased as water temperatures rose.

Kaeriyama et al. (1978) found that chum fry in the Tokachi River of Japan

increased in weight as much as 310~'; during April and May of 1976; however,

there was no increase in weight during the same months of 1975, when the river

was colder and more turbid. Average weight of stomach contents in 1976 was

four times that of 1975.

Predation is sometimes a major source of mortality in chum salmon fry

(Hunter 1959; Kirkwood 1962). Hunter (1959) estimated that predation losses

accounted for 23 to 85% of the total numbers of fry emerging from the gravel

in Hooknose Creek, British Columbia. Although cover requirements of the fry

are not well understood, the fry are known to hide in interstitial spaces in

the substrate during the day as they migrate downstream in small coastal

streams (Neave 1955). Suchanek et al. (1984) did not find a strong preference

by Susitna River chum salmon fry for cover, but substrate, debris, and undercut

banks were more preferred than aquatic or emergent vegetation. Schooling

(Hoar 1956) and turbidity (Suchanek et al. 1984) also provide protection from

predators. However, chum salmon tend to avoid high turbidity during the

rearing phase (Suchanek et al. 1984). Exposure to high levels of suspended

sediments can lead to "tail rot", as well as reduced gas exchange across gills

as a result of damage, coating, or accumulation of mucus (Smith 1978). Smith

(1978) determined that the 96-hr LC 50 for suspended sediment for juvenile chum

salmon was 15.8 to 54.9 g/l.

In addition to providing cover from predators, rocky substrates are

seemingly necessary to maintain orientation in fry that still have a yolk sac

upon emergence (Emadi 1973; McNeil and Bailey 1975). Emadi (1973) found that

30% of the chum salmon alevins reared on a smooth substrate in the laboratory

12
were malformed; there was no malformation of the alevins reared on gravel

substrate consisting of particles 2 to 3 cm in diameter. McNeil and Bailey

(1975) also stated that hatchery-reared Pacific salmon require a gravel

substrate after emergence.

Terrestrial and aquatic invertebrates are the primary food of chum salmon

that delay seaward migration and rear for a time in fresh and intertidal

waters (Sparrow 1968; Bakkala 1970; Mason 1974). Production of these food

items is high in streams having abundant streambank vegetation and a substrate

with low embeddedness and a low proportion of fines (Mundie 1969; Reiser and

Bjornn 1979).

Both osmoregulatory ability and preference for seawater increase as chum

salmon fry increase in age (Baggerman 1960; Houston 1961; McInerney 1964).

Chum salmon fry that have absorbed their yolk sac can tolerate full strength

seawater (Kashiwagi and Sato 1969). Mason (1974) showed that fry feeding in

intertidal areas are highly tolerant of daily salinity changes. Chum salmon

fry appear to have a physiological requirement for seawater within 3 to 4

months after emergence for normal development (Houston 1961), but have been

raised experimentally to the adult stage in freshwater--with careful handling

(Hoar 1976).

Levanidov (1954) noted that DO levels s 1.5 mg/l at 10° C are lethal to

chum salmon fry, but he found no change in feeding, assimilation, or growth of

fry over the DO range of 5 to 11 mg/l. Lukina (1973) reported 8 to 9 mg/l as

the most favorable concentration at 8 to 10° C. DO levels 5 mg/l cause no

impairment of swimming ability (Dahlberg et al. 1968) or avoidance (Whitmore

et al. 1960) in salmonid fry in general. DO levels 6 mg/l are recommended

for successful rearing of Pacific salmon fry in hatcheries (McNeil and Bailey

1975).

Water velocity requirements for chum salmon fry while in fresh or intertidal waters are not well known. Levanidov (1954) reported that optimum

stream velocities to support the feeding of fry "in the Amur River, Siberia,

were < 20 cm/sec, and Mason (1974) observed fry feeding at a wide range of

velocities < 40 cm/sec in intertidal areas. Similarly, most chum fry in

rearing areas of the Susitna River are found in areas with water velocities

< 35 cm/sec (Suchanek et al. 1984). Out-migration of chum fry is often related

to the rate of river discharge (Brannon and Salo 1982). Chum salmon outmigration in the Susitna River is positively correlated with discharge level

(Roth et al. 1984). McDonald (1960) al so reported that fry out-migration

increased markedly during high flow periods.

HABITAT SUITABILITY INDEX (HSI) MODEL

Model Applicability

Geographic area. This model was designed for use throughout the range of

chum salmon. However, accuracy can be expected to vary greatly, depending on

the watershed.

13
Season. This model was structured to account for seasonal changes in

life stage requirements of chum salmon during the parts of the life cycle when

they inhabit freshwater, which may include all months of the year, depending

on the stream.

Verification level. The model represents our interpretation of how

specific environmental variables combine to determine overall suitability of a

habitat for chum salmon. As a first step in the model evaluation process

(Farmer et al. 1982), we solicited comments from expert reviewers. Solicitation of review comments does not imply endorsement by the reviewers. The

model has not been field tested.

Model Description

Overview. The HSI model presented here represents an attempt to synthesize information on freshwater habitat requirements of chum salmon into a set

of habitat evaluation criteria. The model uses habitat variables based on

parameters known to affect the growth, survival, distribution, abundance, or

behavior of chum salmon. The model produces an index of the ability of a

present or future habitat to meet the requireme~ts of chum salmon during the

three freshwater phases (designated as model components) of upstream migration,

the spawning, embryo, and alevin stages, and rearing and downstream migration.

The relation between habitat variables, model components, and HSI is illustrated in Figure 1.

The following sections document the reasons for choosing a particular set

of habitat variables as a measure of habitat suitability for chum salmon

during a particular phase of freshwater residence. The definition and justification of the suitability levels for each variable are described in a later

section (Suitability Index Graphs).

Upstream migrant component. We included variable one (V1 , Fig. 1) in

this component because temperature can alter timing of upstream migration of

chum salmon (Hunter 1959) and affects the susceptibility of prespawning

sal mon i ds to disease (Wedemeyer 1970); V2 was included because dissolved

oxygen affects swimming abil ity (Davis et al. 1963; Dahlberg et al. 1968) and

avoidance behavior in anadromous salmonids (Whitmore et al. 1960).

No specific variables were included in this component as measures of the

accessibility of the spawning stream. Physical features encountered by chum

salmon while migrating upstream should nevertheless be considered to determine

if the habitat being rated is accessible. Passage criteria for successful

upstream migration of chum salmon have been identified as depths> 0.18 m and

velocities < 2.44 m/sec (Thompson 1972) and, for successful passage from the

mainstem river into natural spawning areas, depths> 0.16 m (Blakely et al.

1985). Use of the Instream Flow Incremental Methodology (Bovee 1982) may be

appropriate to determine if altered discharge regimes provide passage flows.

14
Habitat variables

Suitabi 1ity

indices Component

Temperature ----------- V1 JI-----

Upstream migration

Dissolved oxygen --------- V2

Temperature V3

Dissolved oxygen V4

Spawning, embryo,

Substrate composition Vs alevin HSI

Discharge pattern V6

Salinity V7

Temperature ----------- v, ] Reari ng, down- ----- stream mi grat ion

Dissolved oxygen --------- Vg

Figure 1. Diagram illustrating habitat variables and suitability

indices for the chum salmon HSI model. HSI = the lowest of the

suitability index ratings for variables V1 through Vg •

Spawning, embryos, and alevins component. We included V3 in this component because temperature affects time and duration of spawning (Hunter 1959;

Schroder 1973), survival of embryos (Schroder 1973; Koski 1975), and timing of

emergence and survival of chum salmon fry (Koski 1975). We included V4 because

DO 1eve 1s affect embryo survi va1, i nci dence of deve 1opmenta1 abnorma 1iti es,

and timing of emergence of chum salmon fry (Wickett 1954, 1958; Alderdice

et al. 1958; Koski 1975).

The index Vs was included because spawning will occur only in a limited

range of substrates and because embryo survival, emergence, and production of

fry have been related to substrate composition of spawning redds of chum

salmon (Wickett 1958; McNeil 1966; Rukhlov 1969; Dill and Northcote 1970;

Koski 1975). V6 was included because spawning does not occur where the

velocity is too great and because discharge has been related to survival and

production of eggs and alevins (Wickett 1958; Hunter 1959; McNeil 1966;

15
Cederholm and Koski 1977). V7 was included because salinity affects embryo

survival (Rockwell 1956; Kashiwagi and Sato 1969). Water depth and the

presence or absence of upwelling ground water should also be considered as

factors influencing spawning and incubation.

Rearing and downstream migration component. We included VB in this

component because temperature affects the mortality of chum salmon fry (Brett

1952) and, in general, can alter the timing of seaward migration, smoltification, and the susceptibility of salmonid smolts to disease (Wedemeyer et al.

1980). The index Vg was included because DO concentration could potentially

affect downstream migration by decreasing swimming speed (Dahlberg et al.

1968), eliciting avoidance (Whitmore et al. 1960), or causing direct mortalities of smolts. Model users may also want to consider discharge pattern.

Out-migration of chum fry in the Susitna River is correlated with discharge

level (Roth et a l . 1984). Fish appear to reach a point where they are

physiologically ready to migrate downstream and then an increase in discharge

provides the environmental cue to begin migrating. There may also be some

physical flushing of fish as the heads of rearing sloughs are inundated by

rising water. Food availability should also be considered.

Suitability Index (SI) Graphs for Model Variables

Table 1 lists the sources of information and rationale used in constructing each SI graph. Graphs were constructed by converting available information

on the habitat requirements of chum salmon into an index of suitability ranging

from 0.0 (unsuitable) to 1.0 (optimum or most preferred level). These graphs

should not be construed as graphical presentations of real data, but rather as

hypothetical models of the relation between levels of a particular environmental variable and its corresponding suitability as habitat for chum salmon.

All variables pertain to riverine (R) habitat.

Habitat

R

Variable

Maximum temperature

during upstream

migration.

16

Suitability graphs

1.0

0.8

><

OJ

-0

c

...... 0.6

c-,

+J

0.4 ......

.0

ro

+J 0.2

(/)

0.0

0 5 10 15 20 25

°c
R

V2 Minimum dissolved 1.0

oxygen

( DO) concentration during

x 0.8

upstream migration. Q)

"'0

t::

I-<

e-, 0.6

+->

'r-

"r- 0.4 ..0

rtl

+->

"r-

:::l 0.2 Vl

0.0

4

5 6 7

8

mg/l

R

V3 Extreme intragravel 1.0

temperatures from

I

\ spawning to emer- x 0.8 B I \

gence of fry. Q)

I

\

A "'0

t::

I

\ I-<

A. Maximum

I

\

e-, 0.6

I

\ B. Minimum +->

'r-

,

\

'r- 0.4

f

\ ..0 f \ rtl

5I for

V3= 5I for

A or

Bl +->

,

\ "r-

:::l 0.2

f\

whichever is lower Vl

I\

0.0

I

\

0

4 8 12 16 20

°c

R

V4 Minimum DO concen- 1.0

tration from spawning

to emergence of fry.

x 0.8

Q)

"'0

t::

I-< 0.6

>,

+->

'rr- 0.4 ..0

ttl

+->

'r- 0.2 :::l

Vl

0.0

0

2

468 10

mg/1

17
R Vs Substrate composition 1.0 within riffle and run

areas.

(])

x 0.8

A. Percent of gravel -0

s::

substrate 10-100 mm ....... 0.6

in diameter. >,

+-l

-r-:

r-

'r- 0.4 ..0

co

+-l

0.2 l./)

0.0

a

A

20 40 60 80 100

%

B. Percent fines

« 6 mm)

or

(Percent embeddedness of substrate)

2

SI for Vs = SI for A + S1 for B

2

18

1.0

x 0.8

(])

-0

s::

....... 0.6

>,

+-l

.....

r- 0.4 .....

..0

to

+-l

..... 0.2 l./)

0.0

a 10 20 30 40 50

% fi nes
R Stream discharge

pattern from egg

deposition to downstream migration of

fry.

A. Streamflow stable, < 100-fold difference between

extreme average

daily stream

discharges;

stream channel

stable, with

little shifting.

B. Moderate potential

for flooding: 100 to

500-fold difference

between extreme

average daily stream

discharges.

(Hatch marks

indicate suggested

range of Sl ls for

discharge range,

100-fold equals

0.7,500-fold

equals 0.3).

C. High potential for

substrate scouring: > 500-fold difference between

extreme average

daily stream

discharges during

this period;

stream channel

easily altered

during freshets;

substrate unstable

and easily displaced during

freshets.

D. High potential for

low winter flow or

dewatering, resulting in exposure or

freezing of redds.

19

1.0

x 0.8

Q)

"0C

...... 0.6

>,

+J

-r-

.---

'r- 0.4 oCm

+J

'r- 0.2 :::::l

(/)

0.0

A

B

C or0
R V7 Mean intragravel 1.0 --- salinity. , .... B

x 0.8 .... .... A. For eyed embryos. Q)

"'0 \

B. For alevins. l:: \ ....... 0.6 \

>,

+-' \ 'r- \

'r- 0.4 \ ..0 \ rel

+-' 0.2 \ 'r- \ :::l

V) \

0.0

0 5 10 15 20 25 30 35

ppt

R Va Temperature extremes 1.0

during rearing and I \

downstream migration I \

x 0.8 I \ of fry. Q) B I \ "'0

l:: I A \

A. Maximum ....... 0.6 I \

B. Minimum >, I \ +-'

'r- I \ 0.4 I 'r- \ ..0 I \ rel I +-' \

51 for Va = 51 for A or B, 'r- 0.2 I :::l \

V) I \ whi chever 51 is lower. I

\ 0.0

0 4 8 12 16 20

°c

R Vg Minimum DO concen- 1.0

tration during

rearing and down- 0.8 stream migration x

Q)

of fry. "'0

l::

....... 0.6

>,

+-'

.,...

r- 0.4 'r-

+-'

rel

+-'

'r- 0.2 :::l

V)

0.0

0 2 4 6 8 10

mg/l

20
Table 1. Sources of information and rationale used in constructing

suitability index graphs. Habitat for chum salmon is classified as

"excellent" at 0.8 to 1.0, "good" at 0.5 to 0.7, "fair" at 0.2 to 0.4,

and "poor" at 0.0 to 0.1.

Variable Sources and rationale

Inasmuch as upstream migration of salmon is closely tied to the

temperature regime characteristic of each spawning stream

(Sheridan 1962), we assumed that any deviations from the normal

seasonal temperature cycle during upstream migration would be

suboptimum. Chum salmon migrate upstream primarily at 8 to 14° C

(Hunter 1959; Sano 1966). However, temperatures of 8 to 12° C

were considered excellent, because disease rates for anadromous

salmonids increase markedly at temperatures above 12.7° C (Fryer

and Pilcher 1974; Holt et al. 1975; Groberg et al. 1978). Temperatures 20° C were deemed peor because: (1) temperatures

25.5° C are lethal to anadromous salmonids (Bell 1973);

(2) sublethal temperatures > 20° C are associated with high

disease-induced mortality (Wedemeyer 1970); and (3) upstream

migrations of Pacific salmon are delayed by temperatures> 20° C

(Bell 1973). Temperatures of 15 to 20° C were deemed only fair

because little upstream migration has been observed within this

temperature range (see text).

Dissolved oxygen concentrations that enable undiminished swimming

abilities (> 6.5 mg/l; Davis et al. 1963) and that are recommended

for successful upstream migration of anadromous salmon

(> 6.3 mg/l; Davis 1975) were considered excellent. We considered as poor the levels at which coho salmon swimming speed is

greatly reduced (Davis et al. 1963) or avoidance is high

« 4.5 mg/l; Whitmore et al. 1960), or that are associated with

high mortality of ripe pink salmon in southeast Alaska streams (~ 4 mg/l; Edgington 1981 pers. comm. in Krueger 1981).

Unusually low or high temperatures result in emergence of salmonid

fry at times inappropriate for their survival in the estuary

(Sheridan 1962). Delayed chum salmon fry emergence due to reduced

temperatures has been documented by Koski (1975) and Wangaard and

Burger (1983). Temperatures of 7.2 to 12.8° C were considered

excellent because they were related to high survival (Bailey and

Evans 1971) and normal timing of emergence of pink salmon fry

(Godin 1980). Temperatures that adversely affect survival and

development of chum salmon eggs « 4.4° C; Schroder 1973; Koski

1975; Raymond 1981; Wangaard and Burger 1983) or that inhibit

chum salmon spawning « 2.5° C; Schroder 1973) were considered

poor, as were temperatures above the upper threshold for successful incubation of pink salmon embryos (~ 15° C; Kwain 1982).

21
Variable

Table 1. (continued)

Sources and rationale

McNeil and Bailey (1975) suggested that, like pink salmon embryos

(Bailey and Evans 1971), chum salmon embryos tolerate temperatures

near freezing, provided temperatures exceed 4.40 C for at least

30 days after fertilization.

V4 Low DO levels increase mortality, decrease fitness, and alter

timing of emergence of chum salmon embryos and fry (Wickett 1954,

1958; Alderdice et al. 1958; Koski 1975). Levels corresponding

to high survival, unaltered timing of emergence, and highest

fitness of chum salmon embryos and fry (> 6 mg/l; Alderdice

et al. 1958; Lukina 1973; Koski 1975) were considered good to

excellent. Concentrations corresponding to poor or no survival

« 3 mg/l; Wickett 1954; Mattson et al. 1964; Koski 1975), or

delayed timing of emergence « 3 mg/l; Alderdice et al. 1958;

Koski 1975), were deemed poor.

Vs A. Hunter (1959) reported that chum salmon spawn in gravel 13 to

130 mm in diameter. Sano (1959) reported that chum salmon redds

consisted of > 30% gravel (> 31 mm in diameter). Burner (1951)

reported that chum salmon redds in tributaries of the Columbia

River were composed of 81% gravel « 152 mm). Dill and Northcote

(1970) found survival of chum salmon eggs to be 100% in gravel 50

to 102 mm diameter but only 38% in gravel of 10 to 38 mm. On the

basis of this information, we assumed that a substrate composition

of z 60% gravel 10 to 100 mm in d t ame t e r (and < 10% fines) is

excellent.

B. Sedimentation during incubation is a major source of chum salmon

egg mortality (Neave 1953; Wickett 1954; McNeil 1966; Rukhlov

1969; Scrivener and Brownlee 1982, in prep.). Koski (1975)

observed an inverse relationship between percent fines « 3.3 mm

but> 0.1 mm) and percent survival to emergence of chum salmon.

Higher percentages of fines led to premature emergence of fry,

lower yolk conversion efficiency, smaller size at hatching, and

slower growth. Thorsteinson (1965) reported that redds with

> 13% fines « 0.833 mm diameter) were poor producers of chum

salmon fry because intragravel permeability was reduced.

Scrivener and Brownlee ·(1982) found reduced chum egg-to-fry

survival with increased sedimentation. Thus, levels of fines

associated with high production and survival of chum eggs and fry

« 13%, Thorsteinson 1965; < 14%, Rukhlov 1969) and high survival

and emergence of salmon i d fry in genera 1 « 10%, Hall and Lantz

1969; Phillips et al. 1975) were considered excellent. Levels of

fi nes (> 15%) correspondi ng to lower survi va 1 and emergence of

fry were considered fair to poor. We assumed that, if necessary,

percent fines could be estimated by dividing the percent embeddedness by 2.

22
Variable

Table 1. (continued)

Sources and rationale

Survival of chum salmon embryos and alevins is high in more

stable flow regimes (McNeil 1966, 1969), whereas it is poor in

streams with shifting stream channels and extreme fluctuations in

discharge that result in scouring of redds (Neave 1953; Wickett

1958; McNeil 1966, 1969). Poor survival of chum salmon eggs and

alevins has also been observed during periods of low winter flows

(Hunter 1959; McNeil 1966, 1969). On the basis of the findings

of McNeil (1966, 1969), Cederholm and Koski (1977), and Lister

and Walker (1966), we classified as excellent the streams with a

low probability of redd scouring, i.e., those with stable flow

regimes « 100-fold difference between extreme average daily

stream discharges) and stable stream channels. Considered poor

were streams with a high probability of redd scouring (> 500-fold

difference between extreme average daily stream discharges during

winter) or redd freezing or desiccation, i.e., streams with

unstable stream channels and streambeds and high potential for

flooding, or high probability of very low flows during the incubation period. Streams with characteristics intermediate between

these extremes were assumed to be good to fair.

Rockwell (1956) reported that survival of chum salmon embryos

was highest in constant salinities < 6 ppt (corresponding to

upper and .middle reaches of the study area); mortality increased

to 67% at 6.0 to 11.6 ppt, and 100~& at > 12 ppt. Kashiwagi and

Sato (1969) found that percent mortality of chum salmon eggs to

hatchi ng was 0 at s 9 ppt, 25 at 18 ppt, 50 at 27 ppt, and 75 at

35 ppt. However, nearly all alevins hatched from eggs exposed ta

salinities> 9 ppt died within a few days. Helle (pers. comm.)

found no survival of eggs deposited by chum and pink salmon in a

saline (4 to 8 ppt) stream. On the basis of these somewhat

conflicting results, we assumed < 4 ppt to be excellent, 4 to 9 ppt

to be good to fa i r, and > 9 ppt to be poor. It shoul d be noted

that Hartman (pers. comm.) has observed that chum eggs can survive

tidal inundation by water of up to 24 ppt salinity if there is

periodic (daily) flushing of redds by freshwater. The laboratory

observation by Rockwell (1956) that chum salmon eggs and alevins

can survive in seawater up to 30 ppt for several days at low

temperatures provides corroborating support for this suggestion.

We therefore assumed that short term high salinities caused by

tidal inundation would be a less appropriate measure of habitat

suitability than some measure of average salinity conditions.

23
Variable

Table 1. (concluded)

Sources and rationale

To insure optimum conditions for smoltification, timing of seaward

migration, and survival of chum salmon smolts, temperature should

follow a natural seasonal cycle as closely as possible (Wedemeyer

et al. 1980). Temperatures of 7 to 12° C were considered

excellent because temperatures 12° C were recommended by

Wedemeyer et al. (1980) for seaward migration of salmonid smolts

to prevent altered timing of migration and smoltification, and

because Bell (1973) listed 6.7 to 13.3° C as the temperature

range suitable for downstream migration of chum salmon. Slightly

warmer temperatures (about 8 to 13° C) would be optimum for

growth (Brett 1952; Levanidov 1954; McNeil and Bailey 1975).

Temperatures of 14 to 20° C were considered only fair because the

risk of disease is probably higher (Fryer and Pilcher 1974; Holt

et al. 1975). Temperatures> 20° C were considered poor because

growth of the fry of chum sa 1men (Kepshi re 1976) and other

salmonids (Reiser and Bjornn 1979) ceases in this range and

because mortality occurs at 23.8° C (Brett 1952).

We considered as excellent the DO levels corresponding to high

feeding and growth rates in chum salmon fry (> 5 to 11 mg/l;

Levanidov 1954) and the lack of impairment in swimming (> 5 mg/l;

Dahlberg et al. 1968) and lack of avoidance (> 5 mg/l; Whitmore

et al. 1960). Levels causing high avoidance and reduced swimming

ability in salmonid fry in general, or mortality in chum salmon

fry (1.5 mg/l; Levanidov 1954), were deemed poor.

24
Descriptors for each habitat variable were chosen to emphasize limiting

conditions for each variable. This choice reflects our assumption that

extreme, rather than average, values of a variable most often limit the carrying capacity of a habitat.

Use of the Model

H5I determination. It was assumed that the most l i mtt t riq factor (i .e.,

the lowest 51 score) defines habitat suitability for chum salmon in freshwater;

thus,

H5I = minimum value among suitability indices Vi to V9 •

Field application of the model. General guidelines for measuring aquatic

habitat variables are provided by Hamilton and Bergersen (1984). Detailed

sampling of habitat variables over time would provide the most reliable and

replicable H5I values. The method used to calculate 51 values should be

carefully documented to insure that decision makers understand the quality of

the data used in determining the H5I for a particular chum salmon habitat.

Model Interpretation and Limitations

The model described here is a general description of the freshwater

habitat requirements for chum salmon and should not be expected to discriminate

among different habitats with a high resolution. A positive relation between

H5I Is generated from this model to measurable indices of population abundance

(e.g., standing crop) should not be assumed, unless it has been demonstrated

by testing in habitats similar to those where the model will be applied. The

major reason no relation is assumed is that although the model variables may

be necessary in determining the suitability of habitat for chum salmon, they

may not be sufficient. Other factors not included in this model, including

differences in a watershed or in a run or stock, may determine abundance to a

greater degree than the variables included in "this model. Data describing

measurable population responses of chum salmon to additional factors, however,

are insufficient at this time to use in a chum salmon habitat model. The user

should consult Hale (1981) for detailed discussion of the further research

needed to overcome the deficiencies in the existing knowledge of freshwater

habitat requirements of chum salmon.

The proper interpretation of the H5I is one of comparison. If two areas

have different H5I 's, the one with the higher H5I should have the potential to

support more chum salmon than the one with a lower H5I. Outputs of the model

should be interpreted as indicators (or predictors) of the quality of habitat

for chum salmon: excellent (H5I 0.8 to 1.0); good (0.5 to 0.7); fair (0.2 to

0.4), or poor (0.0 to 0.1). Proper use of this model should follow the admonition by Banks (1969: 131) that lithe consequences of man-made changes (on

anadromous salmonids) ... can be predicted in general terms from the existing

literature, but (due to the formation of local stocks) each situation is

unique ... and requires studies of the special needs of each river system as

well as the flexible application of general principles."

25
ADDITIONAL HABITAT MODELS

No other freshwater habi tat mode 1s for chum sa 1mon were found in the

literature. Habitat suitability index models for the estuarine phase of the

chum salmon life cycle are given by Sheperd and Washington (1982). The user

is referred to Terrell et al. (1982) and U.S. Fish and Wildlife Service (1981)

for general guidance in modifying habitat models to meet specific needs.

INSTREAM FLOW INCREMENTAL METHODOLOGY

Instream Fl ow Incremental Methodology (IFIM) is a process of stepwise

analysis used to assess instream flow problems (Bovee 1982). The Physical

Habitat Simulation System (PHABSIM) model (Milhous et al. 1984), a component

of IFIM, is used to compute the amount of available instream habitat for life

stages of a species as a function of streamflow.

The output generated by the PHABSIM component of I FIM can be used for

several IFIM habitat display and interpretation techniques, including:

1. Habitat Time Series. Determination of impact of a project on a

species' life stage habitat by imposing project operation curves

over baseline flow time series conditions and integrating the

difference between the corresponding time series;

2. Effective Habitat Time Series. Calculation of the habitat requirements of each life stage of a single species at a given time by

using habitat ratios (relative spatial requirements of various life

stages); and

3. Optimization. Determination of flows (daily, weekly, and monthly)

that minimize habitat reductions for a complex of species and life

stages of interest.

Suitability Index Curves as Used in IFIM

PHABSIM utilizes Suitability Index (SI) curves that describe the instream

suitability of the habitat variables most closely related to stream hydraulics

and channel structure (e.g., velocity, depth, substrate, cover, and

temperature) for each major life stage of a given fish species (e.g., spawning,

egg incubation, larval, juvenile, and adult). The Western Energy and Land Use

Team has designated four categories of curves and standardized the terminology

pertaining to the curves (Armour et "al. 1984). Category one curves are based

on literature sources and/or professional opinion. Category two (utilization)

curves, based on frequency analyses of field data, are fit to frequency histograms. Category three (preference) curves are uti 1i zat i on curves with the

environmental bias removed. Category four (conditional preference) curves

describe habitat requirements as a function of interaction among variables.

The designation of a curve as belonging to a particular category does not

imply that there are differences in the qual ity or accuracy of curves among

the four categories.

26
I FIM ana lyses may uti 1i ze any or all categori es of curves, but category

three and four curves should yield the most precise results. Category two

curves yield accurate results if they are transferable to the stream segment

under investigation. If category two curves are not transferable for a

particular application, category one curves may be a better choice.

For an IFIM analysis of riverine habitat, an investigator may utilize the

curves available in this publication. If an investigator believes that spawning habitat utilization in the study stream is different from that represented

by the SI curves, he or she may want to modify the existing SI curves or

co11 ect data to generate new curves. Once the curves to be used have been

defined, the curve coordinates are used to build a computer file (FISHFIL),

which becomes a necessary component of PHABSIM analyses (Milhous et al. 1984).

Modeling of chum salmon habitat by use of PHABSIM has been done for the

spawning (Vincent-Lang et al. 1984) and the rearing (Hale et al. 1984) 1ife

stages.

Availability of SI Curves for Use in IFIM

The IFIM is used for riverine environments, and since chum salmon utilize

streams only for spawning migration, spawning, egg incubation, rearing and fry

outmigration, no SI curves are necessary for the sea life juvenile and adult

stages . All of the SI curves for I FIM ana lyses of chum sa 1mon habi tat presented in Figures 2-4 are category one (except for spawning velocity), and were

developed from data and information found in the literature. Because IFIM

uses English units, all of the SI curves are given in English units. Investigators are encouraged to review the curves carefully, compare them with curves

derived from field data when possible, and modify them if appropriate.

Vincent-Lang et al. (1984) present category 2 and 3 curves for chum salmon

spawning depth, velocity, substrate, and upwelling. Suchanek et al. (1984)

describe category 2 and 3 curves for depth, velocity, and cover during the fry

life stage of chum salmon.

Spawning migration. No curves are presented here for spawning migration

requirements, although variables which may limit migration should be identified

before proceeding with an investigation. Length of migration may vary from

less than one mile to over 1,500 miles and, therefore, critical stream reaches

should be identified which may impede passage to spawning areas. Out-migration

of spawners is not of concern because chum salmon adults die after spawning.

Other variables which may be limiting to migration include water temperature

and dissolved oxygen concentrations. Curves for these variables are in the

HSI model section of this report (VI and V2 , sources and assumptions in

Table 1).

Spawning and egg incubation. There are two approaches for determining

the amount of spawning and egg incubation habitat for a stream reach. One

approach, which treats spawning and egg incubation as separate life stages

(each with its own set of habitat suitability criteria), may be used when

weighted useable area (WUA) does not vary by more than 10% during the

spawning/egg incubation period. If WUA is found to vary by more than 10% (as

a result of streamflow variation), then the determination of effective spawning

habitat (Ni l hous 1982) is the recommended approach (to be discussed later).

27
Coordinates

x y

0.0 0.4

0.3 0.9

1.1 1.0

3.2 0.5

3.6 0.1

5.0 0.0

20.0 0.0

1.0

x

Q) 0.8

-0

s::

......

>,

0.6

+-'

......

r-

...... 0.4 ..Cl

to

+-'

......

:::l 0.2 (/)

x

0.0

0.1

1.0

3.3

10.0

100.0

y

0.0

0.0

1.0

1.0

0.0

0.0

o.0 -l-~-.-~-.-_--.---.---.----"';'~

01234 5

Velocity (ft/s; 0.5 ft above

streambed)

x 0.8 Q)

-0

.s 0.6

;:: 0.4 ......

..Cl

to

0.2 :::l

(/)

o. a-f-o. ......

a 5 10

Water column depth (ft)

x

30.0 0.0

34.0 0.0

43.0 1. a

55.0 1. a

62.0 0.0

100.0 0.0

1.0

x 0.8 Q)

-0

s::

...... 0.6

>,

+-'

......

r-

...... 0.4 ..Cl

to

+-'

......

:::l 0.2 (/)

0.0

30 40 50 60 70

o Water temperature ( F)

Figure 2. SI curves for chum salmon spawning velocity, depth, and temperature.

28
1.0

Coordinates

x

0.0

0.3

1.7

10.0

100.0

y

0.0

0.2

1.0

1.0

1.0

x 0.8

(],I

"'0

c:

t-l 0.6

;:: 0.4 ......

..0

ro

+-'

.; 0.2 V1

0.0

o 5 10

Intragravel apparent velocity

(ft/hr)

1.0

x

0.0

0.1

100.0

y

0.0

1.0

1.0

x 0.8

(],I

"'0

c:

t-I 0.6

...... 0.4 ..0

ro

+-'

:::::l 0.2 V1

-

-

-

0.0

0.0 , I I I

0.5 1.0

Water column depth (ft)

4 8

0.4

0.8

x

(],I

-g 0.6 t-l

..0

.rg 0.2

:::::l

V1 0 . 0 ..............,.......,--r--r-........,.---or-"'T""~

o

0.0

0.3

0.3

1.0

1.0

0.0

0.0

x

0.0

0.4

1.5

2.0

4.0

8.0

100.0

Predominant substrate particle

size diameter (inches)

Figure 3. 51 curves for chum salmon egg incubation velocity, depth,

substrate, and temperature.

29
Coordinates 1.0

x --'L 0.8 32.0 0.0 x 35.0 0.2 QJ

-0

36.0 0.6 s::

...... 0.6

38.0 0.8 >,

40.0 1.0 +oJ

.r- 0.4

56.0 1.0 r-«

.r59.0 0.5 .c

ro

61.0 0.0 +oJ 0.2 .r100.0 0.0 ::::l

Vl

0.0

30 40 50 60 70

Water Temperature (oF)

Figure 3. (concluded).

30
1.0 , ,

Coordinates I ,

x 0.8 - Feeding , OJ , ry y "'0

c: ,

x (feeding) (emigration) ........ 0.6 I Emi grq ti on ,

0.0 1.0 0.0 >, I ,

r-

+-'

0.7 1.0 .r- I ,

1.0 0.0 1.0 .r- 0.4 I , r- .D I , 5.0 1.0 ro

+-' I , 6.0 0.0 .r- 0.2 I , r- :::J

20.0 0.0 0.0 Vl I ,

0.0

0 1 2 3 4 5 6

Mean col umn velocity (ft/s).

1.0

x y x 0.8 OJ

0.0 0.0 "'0

c:

0.5 1.0 ........ 0.6

100.0 1.0 .rr-

.r- 0.4

.D

ro

+-'

.r-

:::J 0.2

Vl

0.0

0 1 2

Water column depth (ft)

1.0

x y x 0.8

0.0 0.0 OJ

"'0

0.5 0.0 c:

........

1.0 1.0 >, 0.6

+-' 10.0 1.0 .r20.0 0.0 .r- 0.4 .D 100.0 0.0 ro

+-'

.r-

:::J 0.2

Vl

0.0

0 10 20

Predominant substrate particle

size diameter (inches)

Figure 4. 51 curves for chum salmon fry velocity, depth, substrate,

and temperature.

31
y y

x (feeding) (emigration) 1.0

31.0 0.0 0.0

39.0 0.1 0.8 41.0 0.9 x

Q)

46.0 1.0 1.0 "'0

c:: 54.0 1.0 1.0 ...... 0.6

57.0 0.8 0.8 >,

+.> 61.0 0.2 0.2 .,.... 0.4

100.0 0.0 0.0 .,....

Ll B ro

+.> A. 0.2 Feeding and growth .,....

::::l B. Emergence and emigration U1

O.0 ...,..--.,....................-r--.....,r-.-...,.~"""""'""

30 40 50 60 70 80

Water Temperature (OF)

Figure 4. (concluded).

32
The SI curve for chum salmon spawning velocity utilization (Fig. 2) is

category two, based on frequency ana lyses of data co11 ected from the Skagi t

River near Marblemount, Washington, during December 1976 (Kurko 1977).

Spawners were observed by boat, and velocity measurements (0.5 ft above the

stream bottom) were taken at the upstream edge of each redd (n = 227). The

resultant velocity curve is similar to that developed by Wilson et al. (1981),

which was based on a combination of literature review, professional opinion,

and field data. The range of velocities selected by spawning chum salmon in

Kurko·s study also encompasses spawning site velocity ranges reported by Smith

(1973). The relative suitability of velocities greater than 1.0 ftls, however,

should be field-verified before using the curve in IFIM studies. Chum salmon

tend to key on areas of upwelling ground water, which influences their selection of surface water velocities (Kogl 1965; Sano 1967; Vincent-Lang et al.

1984).

Chum sa 1mon spawn i ng has been reported to occur in depths rang i ng from

2 inches to 6.3 ft (see HABITAT USE INFORMATION). "Preferred" or recommended

depths have generally been reported to be within the 1.0 to 3.3 ft range

(Smith 1973; Hale 1981). There is general agreement on the suitability of

depths to 3.0 ft (Hale 1981; Wilson et al. 1981), but the relative suitability

of deeper waters is rather uncertain, possibly because of difficulties involved

in sampling. The spawning depth utilization in any given stream may be a

narrow subset of the literature-based curve presented here, and field verification is recommended.

No SI curve was developed for chum salmon spawning substrate suitability.

Therefore, when evaluating spawning WUA as a function of streamflow, use of

the egg incubation substrate SI curve is recommended.

No evidence was found to suggest that cover is a requirement for chum

salmon spawning. Therefore, a cover SI curve is unnecessary for IFIM analyses.

The temperature ranges within which chum salmon spawning occurs may vary

wi th 1oca 1e and season. Among the spawn i ng temperature ranges reported (see

HABITAT USE INFORMATION), temperatures overl apped primari ly wi thi n the 43 to

55° F range, and minimum and maximum temperatures reported were 35 and 61° F.

The water velocities required for successful egg incubation may be

dependent upon several interrelated variables. A surface water velocity of

0.0 ft/s may be suitable if there is adequate groundwater upwelling, if

dissolved oxygen concentrations (V~) remain adequate, and if siltation,

freezing, dessication, and metabolite buildup are not problems. The maximum

velocities tolerable will depend on the "scourability" of the substrate, which

is partly a function of substrate particle sizes. The SI curve for egg incubation velocity (Fig. 3) is based, on information reviewed by Hale (1981), and

represents relative suitability of velocities through the gravel.

The SI curve for water depth during the egg incubation period is based on

the assumptions that (in most situations) eggs must remain submerged for the

duration of the incubation period; that high water temperatures, low dissolved

33
oxygen concentrations, and freezing are not problems at low flows (see Fig. 3

and V4 ) ; and that embryos are at least 3 inches deep within the substrate. If

any of these conditions cannot be met, then the depth curve may have to be

modified to ensure embryo survival.

The 51 curve for egg incubation substrate suitability is based, in part,

on an experiment by Oi 11 and Northcote (1970) who found 100% survi va1 of

embryos to emergence in substrate material 2 to 4 inches in diameter, and 31%

survival in 0.4 to 1.5 inches. The curve is meant to represent relatively

homogeneous substrate with less than 20% sand and silt content.

No 51 curve is necessary for egg incubation cover because eggs are buried

in the substrate. The substrate curve is assumed to take care of embryo cover

requirements.

The 51 curve for egg incubation temperature is based on information

reviewed by Hale (1981). Investigators have reported that temperatures from

40 to 560 F are suitable for egg incubation, and that percent survival of

embryos increases as temperatures increase from 32 to 40 0 F. No investigators

reported finding eggs in water with temperatures greater than 520 F.

Effective spawning habitat. The other approach for determining spawning

and egg incubation habitat measures effective spawning habitat (Milhous 1982)

and is recommended when the spawning/egg incubation period is greater than 2

months and WUA varies by more than 10% during the spawning/egg incubation

period. Effective spawning habitat is habitat that remains suitable throughout

the spawning and egg incubation period. In a given stream reach, the area of

effective spawning habitat is equal to the area of suitable spawning habitat

minus the spawning habitat area that was dewatered, scoured, or silted-in or

otherwise determined to be unsuitable during egg incubation. Factors to

consider when determining habitat reduction because of dewatering include the

depth of the eggs within the streambed, temperature and dissolved oxygen

requirements of incubating eggs, and fry emergence requirements. To determine

habitat reduction from scouring, the critical scouring velocity (Fig. 5) can

be determined by:

(

dbf) 1/6

= 22.35 065

1/2

[K/\ - I)J

1/2

(065)

where V

c

=critical velocity in ft/s

dbf = average channel depth (ft) at bankfull discharge

065 = substrate particle size diameter (ft) not exceeded by 65%

of the particles

K

s = 0.080, a constant pertaining to the general movement of the

surface particles

S = specific gravity of the bed material, and ranges from 2.65

s to 2.80

34
Coordinates

x

o 0

V. - 0.001 0 mln

V . 1 mln

V

c - 0.001 1

V

c

0

20 0

V. is the minimum velocity necessary to mln

prevent siltation of spawning sites; V is

c

the critical velocity, above which scouring

of spawning sites will occur.

x0 0

Dmin - 0.001 0

D

min 1

100 1

D. is either the minimum depth required mln

for egg incubation (~ 0.0) or ice depth

(when ice is present during egg incubation).

1.0

x 0.8

QJ

-0

s::

...... 0.6

>,

+-'

......

...... 0.4 I-

..0

to

+-'

......

::::s 0.2 Vl

0.0 • I

V

min Vc

Velocity (ftls )

1.0

x

QJ 0.8

-0

s::

......

>, 0.6

+-'

......

r--

0.4 ......

..0

to

+-'

......

::::s 0.2 Vl

0.0 I

D . mln

Depth (ft )

Figure 5. 51 curves for spawning/egg incubation velocity and depth,

for effective spawning habitat analyses (Milhous 1982).

35
Factors to consider when determining habitat reduction from siltation

include suspended sediment concentrations, minimum velocities necessary to

prevent siltation (Fig. 5), and dissolved oxygen concentrations among the

embryos. More detailed information about the analysis of effective spawning

habitat is presented in Milhous (1982).

Fry. Habitat for fry may be required for up to 120 days or more after

the onset of hatching, depending on the time of hatching, the duration of

fresh water rearing, and on the distance from the spawning site to the estuary.

After egg hatching, alevins remain in the gravel for 1 to 3 months. While

alevins are in the gravel, we have assumed that SI curves for egg incubation

habitat are also suitable for fry requirements. Fry generally emerge from the

gravel sometime between March and May, and may not begin downstream migration

for a month or two. Out-migration may take another 2 months to complete.

Therefore, habitat for free-swimming fry is required for the time period

between emergence and completion of out-migration to estuaries, the duration

of which is largely dependent on locale. If fry reach the estuary soon after

emergence (as when chum salmon spawn in intertidal areas), then SI curves for

fry are not required.

According to Levanidov (1954), velocities less than 0.66 ft/s are optimal

for feeding by fry. Availability of higher velocities, however, may be

important for assisting out-migration of fry. The SI curve for fry velocity

is based on this information and the assumption that higher velocities will

not injure fry during displacement.

The SI curve for fry depth was based on the assumption that all depths

greater than a minimum are suitable, as long as cover is available, and that

velocities and temperatures are within the range of suitability. Fry of many

salmonid species have generally been found in depths ranging from 1 to 2 ft.

Chum salmon fry in the Susitna River have been observed as deep as 3.5 ft, but

most were found at depths < 1.6 ft (Suchanek et al. 1984). Fry may tend to

select shallower water to avoid predation or to avoid higher water velocities.

The SI curve for fry substrate is based on the fact that chum salmon fry

often use substrate for cover (Neave 1955) and the assumption that fry from

1.2 to 3.0 inches in length would require particles ranging from large gravel

to small boulders (with less than 40% embeddedness and low percent fines) to

provide adequate interstitial spaces. No curve was developed for fry cover.

Requi rements for cover may be part i ally represented by the fry substrate

curve. Suchanek et a l , (1984) did not find a strong preference by Susitna

River chum salmon fry for object cover. Cover requirements of chum fry are

often met by schooling behavior (Hoar 1956) and by turbidity (Suchanek et al.

1984).

The SI curve for fry temperature suitability was taken from Hale (1981).

The curve suggests that minimum temperatures required for feeding and growth

are higher than those required for out-migration.

36
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1966. Salmon of the North Pacific Ocean - III. A review of the

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Natural propagation of chum salmon

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45
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48
50272 -101

REPORT DOCUMENTATION 11..REPORT. NO. [2- PAGE Blo1og1ca1 Report 82(10.108)

... Title and Subtitle

3. Reclplem-s Accession No.

S. Report Date

August 1985 ~abitat Suitability Index Models and Instream Flow Suitability

Curves: Chum Salmon

ft. Author(s) 8. Performin. O..anlzatlon Rept. No.

9. Performln. O..anlzatlon Name and Addre.. Western Energy and Land Use Team

U.S. Fish and Wildlife Service

Drake Creekside Building One

2627 Redwing Road

Fort Collins, CO 80526-2899

10. Project/Task/Work Unit No.

11. Contract(C) or Grant(G) No.

(C)

(G)

12. Sponsorln.O... nizatlon Name and Addr..sWeStern Energy and Land Use leam 13. Type of Report & Period Covered

Division of Biological Services

Research and Development

Fish and Wildlife Service 1~

Department of the Interior, Washington, DC 20240

15, Supplementary Not.. The Biological Report series was initiated in October 1984 to replace the

FWSjOBS series published since 1976. The report number assigned to this model follows

consecutively from the Habitat Suitability Index models published as FWSjOBS-82j10._, now

numbered as Bio1ooica1 Report 82(10. ). 1~ Abstract (Limit: 200 words)

A review and synthesis of existing information were used to develop a Habitat

Suitability Index (HSI) model and instream flow suitability curves for the Chum

salmon (Oncorhynchus keta). The model consolidates habitat use information into a

framework appropriate for field application, and is scaled to produce an index r between 0.0 (unsuitable habitat) and 1.0 (optimum habitat). HSI models are designed

to be used with Habitat Evaluation Procedures previously developed by the U.S. Fish

and Wildlife Service.

17. Document Analysis a. Descriptors

Fishes

Salmon

Habi tabi 1i ty

Mathematical models

II. Identlfiers/Open·Ended Terms

Chum salmon

Oncorhynchus keta

Habitat suitability

Co COSATI Field/Group

lL Availability Statement t Re1 ease un1 imi ted

(1M AHSI-Z39.18)

19. security Class (This Report)

Unc1ass ifi ed

20. Secl4rity ClUJ (This Pa.e) Uncl assi f ied

see Instructiona on Reverae

21. No. of Pa.es

48

U. Price

OPTIONAL FORM 272 (4-7n (Formerly NTIS-35) Department of Commerce

{.( U. S. GOVERNMENT PRINTING OFFICE: 1985-580·444/25368 REGION NO.8
•

••• ..__ .... J~.

' . '· 0 ,,-

Hawaiian Islands (>

-(:( Headq uarters . Division of Biological

Services, Wasnington, DC

x Eastern Energy anO Land Use Team

Leetown , WV

* Nationa l Coastal Ecosystems Team

Slidell, LA

• Western Energy and Land Use Team

Ft. Coll ins . CO

• Locat ions of Regional Off ices

REGION 1

Regional Director

U,S. Fish and Wildlife Service

Lloyd Five Hundred Building, Suite 1692

500 N.E. Multnomah Street

Portland , Oregon 97232

REGION 4

Regional Director

U.S. Fish and Wildlife Service

Richard B. Russell Building

75 Spring Street , S.W.

Atlanta, Georgia 30303

I

J- - - - ---6 1

r-- - L, J

I : --

I I

1- - -'

I I

l,_ I 2 ,, -~-

-_.J

REGION 2

Regional Director

U.S. Fish and Wildlife Service

P.O. Box 1306

Albuquerque, New Mexico 87 103

REGION 5

Regional Director

U.S. Fish and Wildlife Service

One Gateway Cente r

Newton Corner, Massachusetts 02158

REGION 7

Regional Director

U.S. Fish and Wildlife Service

lOll E. Tudor Road

Anchorage, Alaska 99503

- ..~,.

REGION 3

Regional Director

U.S. Fish and WildlifeService

Federal Building, Fort Snelling

Twin Cities, Minnesota 551 JI

REGION 6

Regional Director

U.S. Fish and Wildlife Service

P.O. Box 25486

Denver Federal Center

Denver, Colorado 8022 5
u.s. FISH ..WILDLIFE f>ERVICE

DEPARTMENT OF THE INTERIOR hi?]

u.s.FISH ANDWILDLIFESERVICE "'-r rw T " "

As the Nation's pri ncipal conservation agency, the Department of the Interior has responsibility for most of our ,nationally owned public lands and natural resources . This includes

fostering the wisest use of our land and water resources, protecting our fish and wildlife,

preserving th & environmental and cultural values of our national parks and historical places,

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