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AI,)Sb~ ~l· Of Fo'Sho/ C,.;>Me, 11-J,ifaf 'DI'i,
~or
U.S. FISH AND WILDLIFE SERVICE
1. Chum salmon
2. Pink salmon
3. Broad whitefish
4 . Round whitefish
S. Arctic grayling
6. Dolly Varden chat
7. Threespine
stickleback
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1111 IU:PL 'f ~lFER 'tO;
To:
Unit ~d States Department of rhe Interior
Fl!iri AND WlLDUFE SERVICE
~egtern Alaska Ecological Services
733 Y, 4th Avenue, Suite 101
Anchorage. Alaska 99501
(907) 271-45 7 s
·su.s.
IO'f
Government agencies. private organizations and individuals interested in
Alaskan fish/habitat relationships
Descriptions of freshwater fish habitat relationships for the following
Alaskan species are enclosed:
l.
2.
3.
4,
5.
6.
7.
Chum salmon (Oncorhynchus ketal
Pink salmon (Oncorhynchus goro~scha)
Broad whitefish (~oregonus nasus)
Round whitefish ( roso~TUm cyTlndraccum)
Arctic grayling (Thyma Ius arcticus)
Dolly Varden char tSalvel1.1us malma (Walbaum))
Threespine stickleback {GasterOSieUs aculeatus)
These descriptions were prepared by the Habitat Division of the Alu~ka
Department ~f Fish and Game under contract to the U.S. Fish and Wildlife
Service {FWS). Habitat descriptions for coho salmon and rainbow trout are
crrrently being developed by the FWS.
We hope these descriptions will be a useful reference in your Alaskan natural
resource work. If you know others who could use this infonnatio,,. please
contact:
cc:
Ann Flappoport
Regional HEP Coordinator
U.S. Fish and Wildlife Service
Western Alaska Ecological Services
605 W. 4th, Room G-81
Anchorage, ~laska 995Dl
(907) 271-4575
Sincerely,
~egional H£P Coordinator
see distributfcn list
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Di3tribution l.ist
U.S. Fish and Wildlife Service
lOll E. Tudor Rd.
Anchorage, Alaska 99503
-----
Kcitil Bayha, Assistant Regional Direc tor -Envlronm<>nt
J;m Riffe, Assistant Rc~ional Oircctor -Rcfu~('S
Jon Nelson, Assistant Regional Director -FiRhcrl c!-l
Skip l,add, Wildlife Op e r.:ltions
Tom Rothe, Special Studies
.John Morrison, Biologicnl Services Progrnm
Norval Netsch, Fishery Resources
Dick Wilmot, Fishery Research
Howard Metsker, Environmental Contaminant Evnluntion
Dick Nadeau, Technical Services
James Reynolds, Alaska Cooperative Fisherv Research Unit -Fairbanks
Fishery Resources -Fairbanks
Fishery Resources -Kenai
Fishery Resources -King Salmon
Western AlDska Ecological Services -Anchora~<'
Nor t hern Alaska Ecological Services -Fairbank~
Southeast Alaska Ecological Services -Juneau
~~~ka Department of Fish a~d Gam~
P.O. Box J-2000
Juneau, Alaska 99802
Commissioner
Steven Pennoyer, Director, Commercial Fish Division
Stanley Moberly, Director., FRED Division
Richard Logan, Director. Sport Fish DiviRion
Bruce .Bnker I Director. llnbitat ni vis inn
ALask~~artment of Fis~ and \.~me
333 R~spberry Rood
Anchorage, Alaska 99502
Ron Rc~nnrt, Commcrcinl Fish Division
Al Kingsbury, Commercial Fish Division
Dave Daisy, fRED Division
Russ Redick, Sport Fish Division
Lance Trasky, Habitat Division
Carl Ynnugawa, Habitat IHvision
Alaska Department of Fish and Game
1300 College Road
Fa i rbanks, Alaska 99701
George Van Wyhe, Sport Fish Division
Scott Grundy, lila bit at Division
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A la5h.. !!_m.~n.l....!'_( .f.bh 3nd (:am .. ,
iji)s·. 1:' I..J11klln
Juneau, Alaska 99801
li.Jruld lh:inkcl. FREO Oiv is ion
f:-ank V&Jn llullr. Sport Fish Di,·hdun
Rick Rc(•d. !lahit.H !Hvisiun
J\l.1skll Del!_ilrtment of 1-'ish •md Gnmc p)).-·u·n-; 6S6 ___ -· · -·----~ ·-
Kocli:Jk, Alasl.a <'111111';
J;11'"k L.,•chncr, Commerc·i01l r ish ni\'fsinn
Rout !1t11'T i-;
~.tt icln.tl ~t;lr i n-.• F i slu•r ft.•:< S .. ·rv i rc·
701 C Str~rt, 60x 41
Anchorav.a.', Alask:J 9'J'll!
Du.1nt.· !' .. •t'-·rson
N<l t i.,,,.. 1 ~1.1 r i ne F i sh<'r 1 I'S :ic>rv i •. ,.
1'. {), llox I hh8
Junc<Ju, Aln!-'ka 99RO:?
RichJrJ Sumno.:r
F.nvl r•lO:ll\.'llL.II Prot-.•ct i•'n Al/.t.•nc\
701 C :irr~o.:l, Sox IQ
AnciloraK,. •• Alaska (}9511
Rnn !.<.''-' (!-tS!.~J)
41)!, Rt.•v i ~"" Tea~:~
Enviru!li:IL"nt•tl Prnlt.'•'ticm '·"'""<"''
1200 6th AvtJnuc
Seattle, Washin~ton qRJill
A l l.ov:ws
i\<&tinnal !'a rk Scrvit'l'
51,0 W, Sth t\Vc'OU\.'
AnchorJ~c. Alaskn qq~OI
U_.~_A_r.}DJ.. _Corp~_,,_l~ _1-~n.&~n.c_c_r_s
p .o. ~,,,)( 7002
Ancltnr .tgc, A lasJ....a 9t)'ll 0
tu 1 1 Llt.,ycJ
D~viJ H. Uarro~s
Art G~.•.-1.-tch
U.S. Army Cur;Js of l'n)llncc>rs
~:orth l'.1cffic D!vbion --NPDPI.-ER
P,C', BPX 2R70
PCin I Jntl, Oregon '17208
~ik~ Hlnkcs/Carl ~~ufeldrr
U.S. Uureau of l.•md Mana~t·mcnt
4700 E. 72nd Avenue
Anchor~~~. Alaska Q9S07
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Dave Dunaway
Regional Wildlife Biologist
U.S. Forest Service
P.O. Box 1628
Juneau, Alaska 99801
Ken Thompson
Chugach National Forest
2221 E. Northern Lights, Suite 238
Anchorage, Alaska 99508
Devony Lehner
U.S. Soil ConservaLion Service
2221 E. Northern Lights, Suire 129
Ancho~age, Alaska 99508
Eric Yould
Alaska Power Authority
334 W. 5th Ave.
Anchorage, Alaska 99501
Attn: Richard Fleming
Doug Redburn
Alas~ta Depar~.nent of Environmental Cnnscrvat ion
Poucm 0
Juneau, Alaska 99811
Michael E. Wheeler, Ecologist
Alaska Department of Environmental C~nservation
437 E. St., Suite 200
Anchorage, Alaska 99501
Bill Gissel
Division of Land and Water, Southcentral District
Alaska Department of Natural Resources
Pouch 7-005
Anchorage, Alaska 99510
Dr. Peter Mickelson
1-lildl ife & Fishcrlas Prnr,ram,
University of Alaska
Fairbanks, Ala~ka 99701
Dr. Robert Burgner
Fisheries Research Institute
University of Washington
Seattle, Washington 98195
Alaska Resources Library
701 C. St., Box 36
Anchorage, Alaska 1)9513
Alaska State Library
Pouch G
Juneau, Alaska 99811
T rvi n~ Rl dr,.
-~~~----------------._ __________________ ___ J
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Rasmussen Library
University of Alaska
Fairbanks, Alaska 99 i 0l
Library
University of Alaska
3211 Providence Dr .
Anchorage, Alaska 99504
Ron Dagon
DOWL Engineers
4040 B. St.
Anchorage, Alaska
.Jim Henvning
Dumes & Moore
800 Cordova, Suite
Anchorage, Alaska
Rod Hofflll!an
OTt Water ~ogineers
99503
101
99501
4790 Business Park Blvd.,
Anchorage, Alaska 99503
Gary Lawley
Envirosphere Co.
1227 W. 9th St.,
Anchorage, Alasl ~a
Don Matchett
Suite 222
99501
Bldg. O., Sui tC' J
Stone & Webster Engineering Corp.
Greenwood Plazcl
Box 5406
Denver, Colorado 8021 7
Larry Houlton
\o/oodward-Clyde Consultants
701 Sesame St.
Anchorage, Alaska 99503
Woody Trih,'!y
Alaska Deputment nf Fish & Game -Su ll ydrn Aquntlc Studies
2207 Spena rd Rd.
Am:horugc, Aloska 99503
Bill Wilson
Arctic Environmental Information and Oatn Center
707 A. St.
Anchorage, Alaska 99501
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FRESHWATER HABITAT
RELATIONSHIPS
CHUM SALMON-ONCORHYNCHUS KETA
AlASKA DEPARTMENT OF FISH & GAME
HABITAT PROTECTION SECTION
RESOURCE ASSESSMENT BRANCH
APRIL., 1981
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FR~SHWATER HABITAT RELATIONSHIPS
CHUM SALMON (ONCORHYNCHUS KETA)
By
Stephen S. Hale
Alaska Department of Fish and Game
Habitat Division
ResQurce Assessment Branch
570 West 53rd Street
Anchorage, Alaska 99502
Hay 1981
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ACKNOWLEDGEMENTS
Many people from the Alaska Department of Fish and Game and from the Auke
Bay Fisheries Laboratory of the National Marine Fis~eries Service freely
gave t~eir time and assistance when contacted about t~is project and it is
a pleasure to thank them and fis~ery biologists from ot~er agencies,
especially t~ose who provided unpublis~ed data and observations from their
own work. The librarians of the Alaska Resources Library and t~e U.S. Fish
and Wildlife Service were of great ~elp.
This pr~Jject \lfas funded by t~e U.S. F1s~ and Wildlife Service, Western
Energy and Land Use Team, Habitat Evaluation Procedure Group, Fort Collins.
Colorado. Contract No. 14-16-0009-79-119.
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TABLE OF CONTENTS
I. INTRODUCTION
A. Purpose
B. Distribution
C. Life History Sunmary
D. Ecological and Economic Importance
II. 5PtCIFIC HABITAT RELATIONSHIPS/REQUIREMENTS
A. Upstream Spawning Migration
1. Te~erature
2. Stream Flow, Current Velocity, and Water Oepth
3. Dissolved Oxygen
4. Other Parameters
B. Spawning
1. Temperature
2. Current Velocity
3. Water Depth
4. Substrate Composition
5. Sa 1 i ni ty
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25
C. Intragravel Development of Embryos and Alevins 26 I 1. Temperature 26
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2. Stream Flow, Current Velocity, and Water Depth 29
3. Substrate Composition
4. Intragravel Flow and Penneability
5. Dissolved Oxygen
a. Oxygen consumption rate
b. Effect on survival
c. Effect on fitness
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40
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LIST OF FIGURES
1. Distribution of Chum Salmon in Alaska and Main Study Sites
2. Conceptual Model of Relationship Between Adult Chum Salmon
and Water Temperature During Upstream Migration
3. Conceptual Model of Relationship Between Adult Chum Salmon,
Stream Depth, Current Velocity and Temperature During
Spawning
4. Conceptual Hodel of Relationship Between Chum Salmon Embryos
and Alevins and Intragravel Temperature, Dissolved Oxygen
Concentration, and Salinity
5. Conceptual Hodel of Relationship Between Chum Salmon Fry and
Water Temperature and Salinity During Downstream Migration
LIST OF TABLES
I. Data Table -Upstream Spawning Migration
II. Data Table -SpawninQ
III. Data Table -Intragravel Develop~nt of Embryos and Alevfns
IV. Data Table -Emergence and Downstream Migration of Fry
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58
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68
75
6. Sa 1 ini ty
7. Chemical Parameters
8. Light
0. Emergence and Downstream Migration of Fry
l. Temperature
2. Cover
3. Su bstra t e
4. Food
5. Light
6. Sa 11n1ty
7. Dissolved Oxygen
8 . Stream Flow and Current Velocity
III . CONCEPTUAL SUITABILITY INDEX CURVES
IV. OEFICI~NCIES lN DATA BASE AND RECOMMENDATIONS
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LIST OF FIGURES
Pac.e --
1. Distribution of Chum Salmon in Alaska and Main Study Sites •t
2. Conceptual Model of Relationship Between Adult Chum Salmon 58
and Water Temperature During Upstream Migration
3. Conceptual Model of Relatilnship Between Adult Chum Salmon,
Stream Depth, Current Velocity and Temperature During
Spawning
4. Conceptual Model of Relationship Between Chum Salmon Embryos
and Alevins and lntragravel Temperature, Oi~solved Oxygen
Concentration, and Salinity
!ig
60
5. Conceptual Model of Relationship Between Chum Salmon Fry and 61
Water Temperature and Salinity During Do~mstream Migration
LIST OF TABLES
I. Data Table -Upstream Spawning Migration 62
II. Data Table -Spawning
III. Data Table -Intragravel Development of Embryos and Alev1ns 68
IV. Data Table -Emergence and Downstream Migrat;on of Fry 75
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I. INTRODUCTION
A. Purpose
The purpose of this report is to present available information
about the freshwater habitat to 1 erances, preferences, and
requirements of the chum salmon, Oncorhynchus keta (Walbaum}, and
to evaluate habitat whfch parameters are most important to the
spe~fes or are most often critical to survival or limiting to
production. This information is intended to provide a data base
for habitat evaluation activities. In particular, Section III,
which attempts to relate various levels of certain environmental
parameters to habitat suitability, is intended to be used with
the Habitat Evaluation Procedures and instream flow methods of
the U.S. Fish and Wildlife Service. These procedures and methods
are tools for assessing the effect of land use development
projects on aqu~tic habitat.
Bakkala (1970) has presented a comprehensive summa~ of
biological data on the chum salmn. The present report is
restricted to habitat data and to the fresnwater stage of the
life histo~ and expands upon the relationship between the chum
salmon and habitat parameters. It also considers recent
information and emphasizes data from Alaska, particularly
unpub l ished data and persona 1 communi cat 1 ons from ffshe ~
biologists who have worked with chums fn the Stat~. Habitat
parameters emphasized are those of a physical or chemical nature.
Certain biological factors affecting the well being of the
population such as feeding, predation, competition, parasites,
and dis~ase are not comprehensively treated.
Although information has been examined from throughout the range
of the species, emphasis is placed upon Alaska becaus~ it is
expected that this is where habitat evaluations using this report
will be made. Populations from different geographic areas
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undoubt e dly have habitat need~ which differ to some extent;
however, data to document such differences are available only in
rare instances. Caution must be used when extending information
from one stock to other stocks. Research in Siberia has shown
differences. in habitat preference:s between sunwner chum and fa 11
chum.
Habitat requirements descri bed in this report are for chum
salmon; however, it occasionally was consid~red useful to include
data for other species of Pacific salmon whe.re such infonnation
wa s lacking ft'r chufl'!~. In these cases. the other species are
always mentionP ·1 nah~ or referred to by the phrase "Pacific
salmon" if the fnfonnat ·.on fs of a general nature. Any reference
to salmon without naming a particular species or using thf phrase
11 Pacific salmon .. always refers specifically to chum salmon.
Reviews of factcrs constituti ng the freshwater habitat of Pacific
salmon have been done by Macy (1954), Nicola et al. (1966), and
Reiser and Bjornn (!979).
Chum salmon, also conmonly caned dog salmon in Alaska, are
anadn:.mous as are ~ther North American species of Pacific salmon.
but the time so~nt in freshwater is primrily for reproduction .
Chums migrate to the ocean in their first spring or summer of
life and spend little time, if any, rearing in freshwater as do
king, sockeye, and coho salmon. Most of the chum salmon•s life
is spent at sea .
Four periods in the fr.eshwater life history can be distinguished :
upstream ntigration of adults, spawning, 1ntragrave1 period of
eggs and a 1evins, and downstream m~g r~t i on of fry . The period
from egg deposition to emergence frc.n the gravel is usually the
stage of highest mortality and is probably the stage where
habitat requirements are most critical .
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This report presents a general summary of the life history and
then, in Section II, gives a detailed discussion of habitat needs
for each life stage in the freshwater environment. Section III
attempts to synthesize the relationship between various
environmental parameters and habitat quality.
Aquatic habitat and chum salmor, relationships presented in this
paper are very general. They are not appropriate for
applications to specifi~ watersheds.
B. Distribution
Chum salmon have the widest distribution of any of the Pacific
salmon, 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. Spawni11g has been
documented in streams and rivers 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 fn Japan to
the Lena River in Siberia (Bakkala, 1970). More chum salmon are
apparently produced on the Asian side of the Pacific than on the
North American side (Merrell, 1970). Spawning has been recorded
in at least 1,270 streams in tht United States (Atkinson et al.,
1967). Chums occur only in small numbers on the North American
coast south of central Oregon or north of Kotzebue Sound in
Alaska (Helle, 1979).
Chum salmon generally occur throughout Alaska except for
certain streams in the Copper River drainage and in the eastern
Bro~ks Range (Alaska Department of Fish and Game, 1978; Atkinson
et al., 1967; Merrell, 1970; Morrow, 1980). Chums are abundant
in Southeast Alaska, especially in the nortnern part of the area.
There are both summer runs, primarily in northern Southeast, and
fall runs main1y in southern Southeast. Chums are a~~n~~nt in
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Prince William Sound, where there are early runs which enter the
Sound in late April to early July and spawn in non-lake-fed
streams of the mainland. middle runs which ente r the Sound in mid
to late July and spawn in lake-fed streams of the mainland and in
stre~ms of the outer islands, and late runs whi ch enter the Sound
~"! late July, ente.r freshwater 1n August and early September and
spawn almost exclusively in spring-f~d creeks of the mainland.
Cook Inlet chums enter freshwater between early July and
mid-September. Chums in the southern part of Cook Inlet spawn in
coastal streams, while those of the northern part util~ze the
large river systems. They are unc001110n on the east side. Chums
are abundant in streams of Kodiak Island. They are present in
bays and estuaries of the island from mid June to early September
and in freshwater f ·om mid August to C!arl y Octobe1 • Chums are
also abundant in the south Alaska Peninsula area where they enter
streams from ~id July to mid September. little is known about
the distribution of spawning chums in the Aleutian Islands. The
Nushagak and Togiak areas are the main chum salmon producers in
the Bristol Bay drainage. thums enter streams of Bristol Bay
from mid June to mid August and spawn mostly in the lower portion
of the larger clearwater tributaries of the main river systems.
Chum salmon are the most abundant salmon species in the Kuskokwim
Bay and River area. There are sixteen or more major spawning
streams in the Kuskokwim River system. Chums are also the most
abundant salmon in the Yukon River drainage. The Yukon River
draina£e is the greatest single river system producer of chums in
the state. There are distinct runs of sunwner chums and fall
chums in the Yukon River. The sunwner chums enter the mouth of
the river from late May through mid July and spawn in tributaries
of the lower and middle portion of the r1ver. The Anvik River is
one of the major producers of su~r chums . Fall chums enter the
river from mid July through eai"ly Septegj)er and spawn primarily
in the tributaries of the uppe .· river. In the Norton Sound and
Kotzebue Sound areas, chums are again the most abundant salmon .
Norton Sound chums arrive in lays and estuaries from June to late
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July. Kotzebue Sound has two major rur. components in July and
August. Most of the Kobuk River chums arrive in the Sound and
migrate up river before the Noatak River chums. The Noatak River
is the major chum producer in the area. Although chum salmon
occur in streams along the entire Alaskan coast to the Canadian
border and beyond, there are relatively f~w north of the Kotzebue
Sound drainage and numbers along the Arctic coast are very
limited. This area is the most northern part of the chum
salmon•s distribution in North America (Alaska Department of Fish
and Game, 1978). Figure 1 shows the distribution of chum salmon
in Alaska and indicates some of the major study sites where data
mentioned in this report was collected.
C. life History Summary
Most of this section has been summarized from Ba~kala (1970}.
In the summer and fall. adult chum salmon leave marine waters and
begin the upstream migration to spawning areas. Spawning
migrations vary in length from short distances in some coastal
streams, where spawning may take place in the intertidal zone, up
t~ as much as 3200 km in long rivers such as the Amur River in
Siberia or the Yukon River (Bakkala, 1970; Morrow, 1980).
Many streams have more than one run of chums, including distinct
runs of summer and fall chums.
Fall chums in the Yukon River average about 0.5 kg more than the
summer chums, spawn farther upriver, and are more likely to spawn
in springs or areas of ground water seepage. Similar differences
exist between summer and fall chums in the Amur River of Siberia;
the fall chums also have a greater fecundity (Sana, 1966). The
spawning grounds of fall chums ar~ different from those of summer
chums in the Amur River in that the fanner tend to be shallower,
have a narrower range of temperature variation, have a weaker
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flow, have a lower dissolved oxygen content and a higher c.arbon
dioxide content, and have a lower pH {Snlirnov, 1947).
Adults do not feed on the upstream migration. The rate of
migration in the Amur in Siberia has been found to be 56 to 115
km per day, and in shorter streams, 1.9 to 4.2 km per day
{Bakkala, 1970). Lebida {1964) found that chums in the Yu :.t on
migrated from about 1 to 53 km per day with a mean rate around 20
to 30 krn/ day •
On the spawning grounds , the female excavates the redd in gravel
by turning to one side and rapidly flexing the body·, creating
water currents with the caudal fin. After a depression is
comp 1 eted, the fema 1 e and the dom1 nant of severa 1 ma 1 es in
attendance enter and silll.ll taneously extrude the eggs and milt.
The female excavates another depression slightly upstream which
serves to cover the fertilized eggs just deposited. Tne female
may dig severa 1 nests and spawn. with more than one ma 1 e. After
spawning, she may stay near the redd and protect the site from
other spawners. Males o·ften spawn with mol"f.: than one female.
3oth sexes die within one to six days after spawning.
The eggs are deposited at depths of 8 to 43 em below the surface
of the gravel (Bakkala, 1970; Burner, 19~1). Fecundity of chum
salmon ranges from about 1.000 to 8,000 eggs with 2,000 to 3,000
being most corrmon.. In Asia, sunmer chum salmon are at the low
end: of this range and autumn chums are at the high end (Bakkala,
1970). The average fecundity of 36 females taken in the Noata.k
River was 3,365 (Bird, 1980). Noatak River ch.Jm salmon are
considered to be an autumn race.
Chum salmon spawn from June to January, but the peak for northern
populations occurs from July to early Septecnber and the peuk for
southern populations occurs 1n October of November.
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MAIN sn.cv sna
(D NOATAK RIVER
(2J CHENA ~rvER
aJ DB.rA RIVER
LiJ f!ldbt/KIZHUYAK lliV!RS
IJJ a..5EN ClEEK
liJ SASH~ CREEK
[%J KASh\N lAY~
FJGURE 1. DSTR~ ~ CHUM SALMON IN ALASKA (FROM ALASKA OEPARTMENT
~ ASH AND GAME, 1911) AND MAIN ST\.OY SITES.
The time spent in freshwater (time from entering the stream to
time of expiration) for adult salmon in a stream in southeastern
Alaska was 11 to 18 days (Mattson et al., 1964). The time spent
in freshwater for other stocks such as those spawning hun~reds of
kilometers up the Yukon River could last up to two months. The
actual redd life, or time spent at the SSJawning sites, in
Traitors River was 5 to 9 days (Mattson and Rowland, 1963).
Chum salmon eggs incubate in the gravel 50 to 130 d~ys. After
hatching, the larvae with yolk sacs sti 11 attached are known as
alevins. The alevins remai"' in the gravel until their yolk sacs
are nearly or completely absorbed wh1ch takes about 30 to 50 days
(Bakkala, 1970). The alevins are largely dependent on the yolk
sac for nourishment although Oisler (1953) has reported that
preemergent alevins consume small amounts of b,enthic organisms
such as diatoms and chironomid larvae. The alevins are about 22
rrm long and weigh approximately 0.16 g at hatching; after
absorption of the yolk sac, they are 27 to 32 mm long and weigh
0.20 to 0.23 g {Bakkala, 1970}.
The period spent in the gravel by the eggs and alevins is a time
of heavy rortality. The survival rate from eggs to fry in
natural streams typically averages less thi\n 10%.
The alevins emerge from the gravel in the spring (March, April,
and May). Those that still have remnants ~t the yolk sac absorb
it shortly after emergence. The young chums are nON comnonly
called fry. Host chum salmon fry begin their downstream
migration to the ocean soon after emergence. Some may remain in
freshwater for severa 1 weeks, especially those that are hundreds
of kilometers from the ocean. The outmigration occurs mainly at
night in April and May and is a combination of displacement and
active swimming.
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Most of the fry travel near the surface and in the center of the
stream where the water currents are strongest (Hunter, 1959).
Sana and Kobayashi (1953, cited by Bakkala, 1970) found that chum
fry in a Japanese $tream migrated from three to five kilometers
per day. In large rivers, the fry travel by day as we11 as
night, particularly if the river is turbid.
The bulk of the diet of fry in freshwater consists of benthic
organisms, chiefly aquatic insects such as chironomid larvae,
mavfly nymphs. stonefly nyqths, caddfsfly larvae, and bhckfly
larvae (Bakkala, 1970). Occasionally, planktonic organisms are
taken.
Predation is a major sourcP of mortality to r.hum fry during the
downstream migration period. Commor predators in North American
streams are cutthroat and rainbo\o.' ~"'f)IJt, Dolly Varden, coho
salmon smolts, squawfish, sculpins. and predaceous birds such as
kingfishers, and mergansers.
Chum fry shov an increasing preference for seawater with
increasing age. Those that have completely absorbed their yolk
~acs can tolerate direct entrance into full strength seawater.
When chum fry emerge from the gravel, they are about 30 to 32 mm
long. At the time of entry into marine waters they range from
about 30 to 60 mm in length and their weight ranges from O.ZO to
3.8Z g. Chum fry entering an estuary in southeastern Alaska
averaged 35 mm in length (Bakkala, 1970).
The fry form schools when entering the estuary if they have not
done so before. Entry into salt water is usually complete by
June. The young chum remafn near shore until mid or late summer,
feeding 1n estuarine areas and even moving back into freshwater
areas with the tides to feed (Mason, 1974). The fry switch from
benthic organisms, which were their main food source in
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freshwater, to zooplankton including cladocerans, copepods, and
barnacle ~auplii and cyprids. By the middle of August, almost
all juveniles have left the estuaries. By mid summer, the young
chums are 100 to 150 mm long and by September, about 150 to 225
ITI11 long.
After leaving coastal waters, the immature chum:; become widely
distributed in the North Pacific Ocean and Bering Sea. By their
second sumner at sea, they are distributed from the Asian to
American coasts with a southern 1 imit of about 40° to 44 o f-' and a
northern limit in the Arctic Ocean {Bakkala, 1970). Chum salmon
experience a high mortality, much of it due to predation, during
their early ocean life. The main types of food taken by chums
consist of polychaetes, pteropods, squid, crustacean larvae,
copepods, amphipods, euphausids, and fish (Bakkala, 1970). The
average mean size of immature chums in marine waters in early
summer ranges from a fork length of 34 em and weight of 0.5 kg
for age one fish to 55 em and 1.9 kg for age five fish (Bakkala,
1970). Younger chums have a greater growth rate than older fish.
The immature stage lasts from half a year up to five and a half
years .
Bakkala (1970) considered the adult phase of chum salmon to begin
on January 1 of the year in which the fish matures sexually and
spawns. Therefore, this phase lasts from ab~ut six months for
those adults spawning 1n June to about twelve months for those
spawning in December.
Mature chum salmon range from age one to age seven. Most chums
mature. at age two or three in southern parts of the species range
and at age three or four in northern parts (an age three fish is
one that has spent three winters in the ocean, has three annuli.
and is in its fourth year of life). Age six and seven fish are
rare. Age three fish are usually the most conmon in southeast
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Alaska. Age four fish predominate from Pri nee Wi 11 i am Sound
north~ard, with age three and age five being next in abundance.
Adults range from 45 to 96 em in fork length and from 0.8 to
13.4 kg in weight; the average s1ze for mature fish is about 60
to 75 em and 4,0 to 7.0 kg (Bakkala, 1970; Merrell, 1970; Morrow,
1980). The record fish cited by Bakkala was 108.8 em and 20.3
kg. Age three fish are slightly smaller in northern parts o~
Alaska {55 to 62 em) than in southern parts {67 to 11 em)
{Bakkala, 1970).
Maturing adults begin the migration back to the natal streams in
the last few months of their lives. The majority of them have
spent two to four years at sea. They leave the high seas feeding
grounds during the period front May to Novembtr and enter co as ta 1
aodters. Little tiltll is spent ;n coastal waters before the
upstream migration to the spawning grounds begins.
Economic Importance
The annual collii1E!rcial harvest of chum salmon in Alaska in the
years 1971 to 1976 ranged from 4.3 to 7.7 million fish, weighed
from 32.1 to 64.8 million pounds, and had a monetary value to the
fishermen ranging from 7.5 to 18 6 million dollars. In 1976, the
first wholesale value of the statewide production of chum salmon
was 35.1 million dollars (Alaska Department of Fish and Game,
1979a).
In 1g80, 9.9 million chums were harvested collii1E!rcially in Alaska.
Most of these were taken in southeast Alaska, Kodiak Island, the
Alaska Peninsula, Bristol Bay, the Kuskokwim River, and the Yukon
River. The total statewide production was 65.3 million pounds
(Alaska Department of Fish and Game, 1980b).
-11-
A further harvest of ..:hum salmon produced in Alaskan streams
occurs in the high sea~ fishery of the Bering Sea . ThP Japanese
mothership fishery harvests two to four million fish annually .
Significant numbers of :hese are believed to originate in western
Alaska (Alaska Departmen t of Fish and Game , 1979b).
The commercial salmon catch in Alaska north of Bristol Bay, which
is predominantly chum salmon, may produce a large percentage of
the tota 1 C.J sh income for many vi 11 agers in western and
northwester'n Alaska.
In addition to t he ~ommercial catch, large numbers of chum salmon
are taken for subsistence use . Chums ar~ used for subsistence
throughout the state, but the bulk are taken in westl!rn and
northwestern areas including the Yukon River drainage in the
interior. The annual statewide subsistence nar.,est of chums
ranks in the hundreds of thousands. In former ye-lrS, large
numbers of chums were taken for sled-dog food, but thh use
declined with, among other things, the increasing use of
snowmach i nes starting in the mid 196Ds. Renewed interest in sled
dogs in the last few years has apparently resulted in an
increased chum salmon harvest . In the Kuskokwim area, the annual
subsistence harvest of chums was larger titan the connercial
narvest until 1977 (Alaska Department of Fish and Game, 1979c).
The s.arne was true for the Yukon River unti 1 1970.
Although not a prime target for sport fishen.len in Alaska,
thousands of chum salmon are taken by anglers every year.
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II. SPECIFIC HABITAT RELATIONSHIPS/REQUIREMENTS
A. Upstream Spawning Migration
The habitat requirements of mature adult chum salmon migratin~
upstream to the spawning grounds are relatively broad. The tir~
spent during this phase varies from less than a day for stock;
which spawn in the tidal zone or within a few kilometers of tte
stream mouth, as is comnon in southeastern Alaska and Prince
William Sound, to several weeks for stocks which spawn in
tributaries of the larger rivers such as the Yukon.
1. Temperature
Manzer et a 1. (1965, cited by -akka 1a, 1970) estimate tl1at
chum salmon at sea can probably tolerate temperatures of 1°
to l5°C, but they prefer a range from 2 or 3°C to l1°C.
Bell (1973) listed temperatures from 11.1 to 14.4°C as the
preferred range for adult chums (he was probably referring
to the freshwater phase only).
The temperature range during upstream migration of chums in
some tributaries of the Kuskokwim River is 5.0 -12.8°C
( Al ~ska Department of Ffsh and Game, 1980a). The
temperature range during the peak of the upstream migrati•Jn
1 n the Anvik River 1 s 10.0 -16. 7°C {Trasky, 1974). The
temperature in Traitors River ranges from 4.4 -19.4°C
during upstream migration; for the peak, the range is 8.9 -
14.4°C (Mattson and Hobart (1962).
Hunter (1959) stated that aJults entered Hooknose Cneek in
British Columbia to begin the upstream migration when the
temperature ranged from 8°C to l4°C and that temperatures
within thi5 range did not seem to have any influence on the
upstream migratiJn.
-13-
II. SPECIFIC HABITAT RELATIONSHIPS/REQUIREMENTS
A. Upstream Spawning Migration
The habitat requirements of mature adult ~hum salmon migrating
upstream to the spawning grounds are relatively broad. The time
spent during this phase varies from less than a day for stocks
which spawn in the tidal zone or within a few kilometers of the
stream mouth, as is cannon f n southeastern Alaska and Pri nee
Wi 11 tam Sound, to severa 1 weeks for stocks which spawn in
tributaries of the larger rivers such as the Yukon.
1. Temperature
Manzer et al. (1965, cited by Bakkala, 1970) estimate that
chum salmon at sea can probably tolerate temperatures of 1°
to 15°C, but they prefer a range from 2 or 3°C to l1°C.
Bell (1973) listed temperatures frilll111.1 to 14.4°C as the
preferred range for adult chums (he was probably referring
to the freshwater phase only).
The temperature range during upstream migration of chums in
some tributaries of the Kuskokwim River fs 5.0 -12.8°C
(Alaska Department of Fish and Game. 1980a). The
temperature range during the peak of the upstream migration
in the Anvik River is 10.0 -16.7°C (Trasky, 1974). The
temperature in Traitors River ranges from 4.4 -19.4°C
during upstream migration; for the peak, the range is 8.9 -
14.4°C (Mattson and Hobart (1962).
Hunter (1959) stated that adults entered Hooknose Creek in
British Columbia to begin the upstream migration when the
temperature ranaed from 8°C to 14°C and that temperatures
within this range did not seem to have any influence on the
upstream migration.
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For Japanese streams, Mi~ara et a1.(1951, cited in Bakkala,
1970) reported that chum salmon start to enter the streams
in th~ fall when the water temperature had declined to 15°C;
most enter at temperatures of 10°C to 12°C. During the peak
of the upstream migration, temperatures ranged from 7°C to
11 °C. In northern Japan, stream temperatures during
upstream migration can range from 0.1 to 20.0°C but, during
the peak of migration, are generally around 7 to 11°C (Sano,
1966}. Stream temperatures in the Memu River are 11 to 12°C
(Sano and Nagasawa, 1958).
Levanidov (1954} stated that, while fall chum in the Amur
River of Siberia rarely die before spawning, many sumner
chums died in certain years because of high water
temperatures.
Bell (1973) suggests a water temperature criteria for
successful upstream migration of chum salmon ranging from
8.3 to 15.6°C, with an optimum temperature of 10.1°C.
2. Stream Flow~ Current Velocity, and Water Depth
One of the important parameters affecting upstream migration
is stream flow. Of course, there are many reaches of
streams that have potentially useable spawning grounds which
are not available because of downstream barriers created by
current velocities beyond the swillll'li ng capabi 1 ity of the
adults, or other physical obstructions. However, in those
streams that normally have spawning runs, the relative flow,
whether the stream is in a high or low water stage, is an
1~~J;Jortant factor. Water velocity, and possibly a hea·1y
suspended sediment load. can be limiting to upstream
migration during flooding, and water depth and elevated
water temperatures can be limiting during low water stages.
-14-
There is little infonnation available on the maxiiJ'klm
sustained swimming velocity of which adult chum salmon are
capable. Chum salmon have less ability to surmount rapids
and waterfalls than other species of Pacific salmon (Scott
and Crossman, 1973). Reiser and Bjornn (1g7g) state that
water velocities of 3-4 m/sec are near the upper limit of
the sustained swimming ability of Pacific salmon, although
darting speeds can range up to about 6.7 m/sec (Bell. 1973).
Thompson (1972, cited by Reiser and Bjornn, 1g7g) suggested
that the maximum velocity for successful upstream migration
of adult chum salmon was 2.44 m,lsec and that the minimum
water depth was 18 em.
Adult chums have been observed traveling upstream in shallow
riffles with the upper part of their bodies above water, but
the distance that could be covered in this fashion is
unknown . In southeastern Alaska, where the streams
general :y have low reservoir capacity and the flow is
heavily dependent on rainfall, there have been several
instances during dry periods when migrating chums have been
unable or have had difficulty moving upstream. Similar
situations have occurred fn British Columbia (Wickett,
1958).
The timing of upstream migrations are often correlated
with flow conditions. Lister and Walker (1966) have
observed chum salmon delaying at the mouth of the Big
Qual icum River in British Colull()fa during a period of
extremely low discharge and then beginning a strong run
upriver in a rapidly rising discharge. The amount of water
discharged ft·om power p 1 ants can a 1 so be a major factor
controlling the timing of upstream migration (Mayama 1nd
Takahashi, 1975).
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3.
Kusnetzov {lg28) noted that L ~e run of fa 11 chums in a
Siberian river corresponded with the water level.
In Prince William Sound, Helle (lg6o, cited by Bakkdld,
lg7Q), observeo that although chum salmon were spawning in
certain streams, they were absent from a particular
glacially fed stream until after a heavy flow and silt load
had decreased. This is also the case in the Oelta River
where spawning chum do not enter the stream until after
freeze up when glacial runoff has ended and the only source
of water is upwelling groundwater. These fish spend enough
time tn snallow water in November to accumulate icicles on
their backs (M. Geiger, personal communication).
Dissolved Oxygen
Reduced swimming performance by salmonids can result from
levels of dissolved oxyge~ below air saturation (Reiser and
Bjorn, 1g7g). This probably i3 seldom a limiting factor to
the migration of chum salmon in Alasl(an st'"~ams, but
infonnation is not available. Some small streams in
Southeast Alaska may experience lowered oxygen levels caused
by the decay of spawned-out salmon ( J. Bailey, persona 1
COITI!luni cation).
Winberg (lg56, c;ted by Bakkala, 1g10) found that the
metabolic rate of adult chums swimming upstream an average
of 115 km per day was seven to eight times g;eater than that
of resting adults. In the Amur River of Siberia, the
average daily use of energy was determined by Nikolskii
(1954, cited by Bakkala, 1970), to be equivalent to 25,810
calories (25.8 Kcal) per kg oodyweight for adult males and
28,390 ca~ories per kg bod~~~~nt for females.
-16-
Oxygen consumption for adult chum salmon migrating upstream
was 215 mm 3o2tg body wt ~ hr for males and 236 mm 3o2tg body
wt -hr for females (Winberg, 1g55 cited by bakkala, 1g70).
Mature adults holding position used 71 mm 3o2tg body wt -hr
l~wakura, 1g63, cited by Bakkala, 1970).
4. Other Pa ramet~rs
Adult chum salron can tolerate an abrupt change from
seawater to freshwater (Kashiwagi and Sato, lg70). ~n an
experimental situation, the water content and the osmotic
concentration of blood returned to normal about three days
after such a transfer.
High Sl."per1ded sediment loads could be inhibiting to adults
attempting an upstream migrat1on. Exposure can lead to tail
rot and reduction of gas exchange across gills by physical
damage, coating, or accumulation of mucous (Smith, 1978).
The g6 hour LC 50 values for juvenile chums from Puqet Sound
obtained by Smith are sediment concentrations of 15.8 to
54,g g/1.
B. Spawning
Spawning grounds must consist of suitable substrate as well as
suitable stream conditions. Many stocks of chum salmon,
particularly fall chum, spawn in areas ~here there is a seepage
of groundwater or springwater (Kobayashi. 1968; Kogl, lg65;
Kusnetzov, 1928; Rukhlov, 1969b).
Chum redds have been measured ranging in area from 0.3 to 4.5 m.
The average chum redd area in the upper intertidal zone of Ol~~n
2 Creek in Prince William Sound ranged from 1.0 to 1.8 m
(Tho~steinson, 1965b). Burner (1951) found that chum redds in
tributaries of the Columbia River averaged 2.3 m2 in area.
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Summer chum redds in an Asian stream averaged 1.3 m2 (Vasilev,
1959, cited by Bakkala, 1970). Kusnetzov (1928) found that
autumn chum redds ranged from 125 to 320 em in length 1nd from
106 to 213 em in width in the Kamchatka and Amur areas. (This
would equal a range fn area from 1.3 to 6.8 m2 if the shortest
width was associated with the shortest length.) Chum redds on
Sakhalin Island averaged 1.6 m2 in area with a range from 0.3 to
4.5 m2 (Kukhlov, 1969a). Sull'l'ller chum redds in the Beshenaya
River of Siberia were 1.0 -2.3 m2 in area and fall chum redds
were 0.5 -1.5 m2 in area (Smfrnov, 1947).
Measurements of the total area occupied by a spawning pair (total
area of spawning grounds divided by numer of spawning pairs)
range from 0.3 to 10.1 m2. In Olsen Creek, this area ranged from
about 1.5 -10.0 m2 (Thornsteinson, 1965b).
Burner (1951) suggested that a spawning pair of chum salmon
requires an area of 9.2 i. Wickett (1958) states that the
optimum density of spawning chum varies with the permeability of
the substrate (that is, tha nu~er of embryos and alevfns a given
area of grave 1 can support depends o1: 'the i ntragrave 1 flow rate).
In N;le Creek, B.C., Wickett found that there were 4.8
m2tspawning pair. In McClinton Creek which has gravel of a
higher permeability, there were 1.2 m2tspawning pair. ~"'"~Ito
(1954, cited by Sene, 1966) reported that the spawning area per
female in a river on Kamchatka varied from 0.3 to 10.1 m2 and
Rukhlov (1969a) found 0.8 -10.0 m2 (average 1.7 m2 ) per spawning
pair on Sakhalin • Tho~steinson (1965b) reported that, on the
average, 1.1 females used a square meter of the Olsen Creek
spawning grounds and Bailey (1964), also at Olsen Creek. reported
a cumulative density for the spawning season of 2.0 females m2•
Schroder (1973) stated that the optimum density for maximum egg
deposition is 1.7 to 2.4 m2 tf~le chum.
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1. Temperature
Burner (1951) found chums spawning in tributaries of the
Colunmia River at water temperatures from 5.1 to 6 .7°C .
Chum salmon in British Columbia spawn at temperatures
ranging from 4°C to l6°C (Neave, 1966). In Hooknose Creek,
B.C., they spawn at a temperature of about 12°C (Hunter,
1959). McNeil (1969·) reported a similar val·•e for Sashin
Creek fn southeast Alaska where most chums spawn after the
water temperature has dropped to 13°C or below. Chum salmon
on Kodiak Island spawned at water temperatures ranging from
6.5 to 9.0°C in the K1zhuyak River and at 9.0 to 12.5°C in
the Terror River {Wilson et al., 1981).
Stream temperatures of the Memu River in Japan during
spawning range from 6 to 9°C (Sano and Nagasawa, 1958).
Spawning of summer ch~m in the Amur River of Siberia occurs
during August and September at water temperatures of 9 to
l1°C while spawning of fall chum occurs during the period
mid-September to the end of November at water temperatures
of about 6°C (Sano, 1966). Ivankov and Andreyev (1971)
report that most of the spawning of the South Kuril chum
populations takes place when the water temperature is 5 to
6°C. On Sakhalin Island, spawning occurs ,.t water
temperatures of 1.8 to 8.2°C (Rukhlov, 1969a) and in the
iski River, it occurs at bottom water temperatures of 2 -
9°C (Smirnov, 1947).
Schroder (1973) noted that there was some inhibition 'lf
spawning by chum salmon in Washington at water temperatures
below 2.5°C. About 981 of the females present spawned when
the water temperature was around 7°C while only 82% spawned
at temperatures around 2 .5°C. Nest construction, spawning
behavior, and egg deposition were all affected by the cold
... ater.
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Water temperatures recolmiP.nded by Be 11 (1973) as criteria
for spawning chum salmon range from 7.2 to 12 .8°C. This is
the same range which McNeil and Bailey (1975) state to be
the preferred spawning tP.mperature for Pacific salmon .
2. Current Velocity
Kogl (1965) measured surface current velocities during
the spawning of chum salmon in the Chena River ranging from
0 to 60 em/sec . He believed that surface current was of
less importance to spawning chums than the presence of
springs or groundwater. Sites where chum spawned in the
Terror and Kizhuyak Rivers of Kodiak Island had current
velocities ranging from 0 to 118 .9 em/sec (Wilson et al.,
193~).
At chum salmon redds in Oregon, Thompson (1972, cited by
Reiser and Bjornn, 1979) measured stream velocities of 46 to
97 em/sec and Smith (1973), after s1mpling 214 redds,
measured a mean velocity of 73 em/sec. Collings (1974)
measured velocities of 21 to 101 em/sec at 12 em above chum
redds in Washington streams . In a spawning channel in
Washington, Schroder (1973) measured stream velocities from
22.5 em/sec to, at high tide, 0 em/sec. These differences
did not appear to disrupt normal spawning behavior.
Rukhlov (l969a) reported that chum spawning on Sakhalin most
oft~n takes place at ~urrent velocities from 10 to 90
em/sec. Rukhlov also found that the average number of eggs
deposited per redd decreased as the stream velocity over the
ro ~dd increased. In the My River, chums have spawned where
the stream velocity was 10 to 80 em/sec (Soin, 1959) and 20
to 100 em/sec (Strekalova, 1963). Spawning grounds in the
Bolshaya Ri ver basin have current velocities of 10 to 30
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em/sec (Krokhin and Krugi~s. 1937, cited by Nicola et al.,
1966).
Chum salmon have been reported spawnirtg in the Memu River of
Japan at stream velocities of 10 to 35 em/sec (Sano and
Nagasawa, 1958). Redds were most abundant in areas where
the velocity was 15 to 20 em/sec and least abundant where
the velocity was less than 10 em/sec or greatF'r than 30
em/sec. Sano (1967) states that chums will spa~n in surface
current velocities of less than 15 em/sec if there is a good
intragravel flow of groundwater.
Some recommended criteria for stream velocities at ;pawning
areas are : 46 to 101 em/sec for Oregon chums (Smith 1973)
and 31 to 70 em/sec for chums, chinooks, and cohos in the
Wynoochee River of Washington (Deschamps et al., 1966, cited
by Smith, 1973). A spawning channel at Jones Creek in
British Columbia used by chum salmon was des i gned for a
water velocity of 31 to 76 em/sec (MacKinnon et al ., 1961).
Tautz and Groot (1975) suggest that, i n the selection of a
spawning s i te, chum salmon prefer an area near a rock.
protruding above the substrate where there is an upwelling,
accelerating current . A favorable current regime is also
found at the boundary between pools and riffles. These
irregularities are cond~cive to a good flow of intragravel
water (Reiser and Bjornn, 1979).
The flow of a stream during spawning relative to the flo.,
during the incubation period can be an important factor
affecting survival. Eggs deposited in shallow water durinq
high flow stages are subject to desiccation and/or freezing
during low flow stages.
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3. Water Depth
Water depths where chum salmon spawn fall within a certain
range but pressure itself is probably not a factor limiting
spawning. However, favorable combinations of current
velocities, bottom configur7 ~ions, and substrates (which are
important factors) tend to occur together within a certain
range of water depths.
Kogl (1965) found that chum salmon dug redds in the Chena
River at water depths from 5 to 120 em. Chums spawned at
greater depths in areas of the Chena River where there were
no springs or ground seepage. In certain areas of the main
channel where the upwelling of groundwater was absent, the
chums deposited their eggs at water depths greater than 100
em. Eggs deposited in less than 100 em of water were
subject to winter freezing. Chums have been observed
spawring in the Delta River at water depths up to 61 to 92
em (Francisco, 1976). Water depths measured at sites where
chum salmon spawned in the Terror and Kizhuyak rivers ranged
from 7.6 to 106.7 em (Wilson et al., 1981).
Burner (1951} measured water depths at chum salmon redds in
tributaries of the Columbia River of 5.1 to 76.2 em with a
mean of 25 em. Smith (1973) reported that the average water
depth over chum redds 1n five Oregon streams was 30 em. In
Washington streams, Collings (1974) measured water depths of
15 to 53 em.
In Hokkaido, water depths of 20 to 110 em at chum spawning
sites were measured by Sana (1959, cited by Bakkala, 1970).
For the Memu River in Tokachi, Sano and Nagasawa (1958)
reported that chum redds we.re most abundant where water
depths were 20 to 30 em and least abundant at water depths
greater than 100 em.
-22-
Soin (1954) in the My River of Siberia, measured water
depths over redds of 30 to 100 em. Kusnetzof (1928)
reported that chums favor water depths of 60 to 100 em for
spawning .
Water depths recommended as criteria for chum ~ by Deschamps
et al. (1966, cited b} Smith, 1973) for a Washington stream
range from 23 to 46 em. Smith {1973} suggested that the
water depth for chums in Oregon be no less than 18 em.
Thompson (1972) uses the same figure. An artificial chum
spawning channel at Jones Creek, B.C., was designed for a
water depth of 31 to 61 em (MacKinnon et al., 1961).
4. Substrate Composition
------------
The suitability of stream substrates from the standpoint of
successful reproduction is more related to its qua l ity for
incubation of eggs and alavins than to the ~ase of
excavating redds by spawning females . Chums may deposit
egg s in bottom materials where redds are fairly easily
constructed but which are less than ideal for incubation,
perhaps because of a high s i 1 t content or 1 ow paras i ty.
Certain substrates in which it would be difficult or
impossible to excavate a redd can be fairly good incubation
substrates. An example is loose cobble and boulders over
bedrocli: where the chums deposit their eggs and mi 1 t into
existing crev i ces .
Rukhlov (1969b) reported that smaller chums on Sakhalin
Island tend to have a higher percentage of sand in the redd.
Rukhlov assumed that the weali:er ('5maller} females build
their nests in less resistant gravel .
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Of fferent i nves ti ga tors have used different methods and
definitions to present substrate composition information
which makes comparisons difficult. In general, chum salmon
excavate redds in gravel beds with a particle size of 2 to 3
em diameter, but they will also construct redds in
substrates with particles of a greater size and will even
use bedrock covered with sma:l boulders (Morrow, 1gao; Scott
and Crossman, 1g73). Generally, su~strates wfth a
percentage of fines (particles less than 0.833 nm fn
diameter) greater than 131 are of poor quality because of
reduced permeability (Thorsteinson, 1965). However, chum
salmon often spawn in areas of upwelling ground water and
may therefore be able to tolerate higher percentage of fines
than would seem desirable if some of the fines are kept ~n
su~pensfon by the upwelling water {J. Helle, personal
comment). Sano and Nagasawa (1958) reported that upwelling
water washed away mud and fine sand from chum redds in a
stream in Japan.
The spawning grounds of the Delta River are composed mostly
of particles 1.3 to 12.7 em in diameter with variable
amounts of fines (Francisco, 1976). Sano (1g59, cited by
Bakkala, 1g70) reported that chum salmon in Hakkaido spawn
in a substrate of which 251 is 0.6 to 3.0 em, and 301 is
greater than 3.1 em. Particles less than 0.5 em comprise
less than 201 ot the composition and particles greater than
3.0 em comprise 25 to 53S of chum spawning gravels in the
Memu River {Sano and Nagasawa, 1958). The spawning beds of
My River are composed of gravel mfxed ·;-1 th sand and small
quantities of silt (Soin, 1g54). Substrate samples taken
from chum redds on Sakhalin Island are mostly composed of
particles 0.2 to 1.0 em fn dfa~neter (Rukhlov, 1969a).
Ru~hlov (1969b) reported that a particle size analysis of
143 samples from chum redds on Sakhalin showed the average
sand content to be about 121; gravel content to be around
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35S; shingle content to be around 45l; and fractions greater
than 10 em to be about SS . Cnum redds in tributaries of t he
Columbia River were found by Burner (1951) to have the
fo 11 owing compos it ion: mud I si 1 t, and sand -6. 0~ 1 medium
and small particles greater tttan sand size but less than
15.2 em 81.02:1 and large gravel over 15.2 em -13.02:.
Hunter (1959) reported that the spawning grounds of Hooknose
Creek in British Columbia consisted of particles from 1.3 to
13.0 em., •tiberal" quantities of coarse sand, and '''a
certain amount" of fine sand and s11t.
Mt:Neil and Bailey (1975) suggested a substrate size range of
1.3 to 15.2 em in diaw~ter for the aquaculture of Pacific
s almon . MacKinnon et al ., (1961) used gravel ranging in
:.ize from 0.6 to 3.8 em for a spawning channel for chum
salmon in British Columbia. Bell {lg7J) recocnnended a siZt!
range of 1.3 to 10.2 em for artificial spawning channels tCl
ue used for salmon, including chums.
Duker (1977). in an experimental situation in Washington,
presented female chums with a choice between four sizes of
spawning gravels. The females preferred the two substrates
which consisted mainly of particles 0. 7 to 7.6 em in
diameter , rather than a substrate with mostly smaller
particles or substrate with mostly larger particles.
5. Salinity
Saline water can interfere with fertilization of the eggs of
chum salmon spawning in or near the intertidal zone .
Rockwe·ll (1956) found that sa 1i nit ies of 18 parts per
thousand {ppt) or greater inhibit fertilization of chum
eggs, although a limited amount of fertilization could occur
at salinities up to 30 ppt.
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c. Intragravel Development of Embryos and Alevins
The intragravel period in the development of the chum salmon
consists of two 1 ife history stages with differing habitat
requirements. that of the fertilized eggs or embryos and that of
the alevins. Further, the habitat requirements of each stage
change with the !;tate of development or age. However, most
investigators have treated the intragravel perioJ as a single
stage from the standpoint of measurements of environmental
parameters~ that is how it is treated here. Wherever data are
available, distinctions between the stages will be made.
Mortality is generally higher for the embryo stage than for the
alevin stage {Levanidov, 1954). Infonnation on the density of
chum eggs and alevins per square meter in Alaska has been given
by several investigators {Bailey, 1964; Dangel and Jewell, 1975;
Mattson and Rowland, 1963; Mattson et al., 1964; Roys, 1968;
Thorsteinson, 1965b).
1. Temperature
Water temperature is an important variable to chum salmon
during the gravel incubation period. The time to hatching
and time to emergence are directly related to water
temperature. Cold air temperatures. combined with a low
stream flow or an insufficient ground water flow, can reduce
intragravel water temperature to the point where embryos and
alevins are killed by freezing. Temperature also influences
other parameters such as the solubility of oxygen in water.
The temperature of intragravel water is controlled by the
temperatJre of the stream water and ground water and by the
rate of exchange among these three reservoirs.
Stream water temperatures from slightly below 0° to about
l5°C have been observed during the incubation period of chum
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salmon (Koski, 1975• McNeil, 1966). A site in the Chena
River had intragravel temperatures of 0.5 to 4.5°C (Kogl,
1965). Intragravel water temperatures at several chum
spawning sites in the Noatak R:ver ranged from about 6.1 to
10.0°C in early October to a low of 0.2 to 3.3°C in March
(Merritt and Raymond, in prep.). Francisco (1977) measured
intragravel temperatures ·of 0.6 to 6.7°C in the chum
spawning areas of the Delta R.iver during the incubation
period. Intragravel temperatures fn the Kfzhuyak River
during the month of April vere about 3-5°C (Wilson et al.,
1981). Ba i 1 ey {1964) reported i ntragrave 1 temperatures
{probe 20.3 em deep) in Ols~n Creek of 4°C in October, near
ooc in December and January, and 1°C in mid-March just prior
to fry emergence.
Sana and Nagasawa (1959) reported that the intragravel
temperatures in the Memu River of Japan during incubation
ranged from 7.0 to 11.0°C. Sana (1966} found that autumn
chum in northern Japan chose to spawn in spring areas where
the water temperature in the spawning bed did not become
lower than 4°C. Semko (1954, cited by Sana, 1966) reported
intragravel water temperatures in a river on the Kamchatka
Peninsula of 3.9 to 4.9°C during chum egg incubation and 2.4
to 3,0°C during the alevin phase. Levanidov (1954) found
water temperatures in chum redds in the River Khor of 3.3
and 6.0°C and, in the Bira River, intragravel temperatures
during incubation ranged from 3.5 to s.ooc (Oisler, 1951).
The lcwer end of these temperature ranges may not be ideal
for chum salmon in some areas. Koski (1975) found this to
be the case for chum eggs in a Washington spawning bed and
Schroder (197 3) , working in the same a rea, stated that
temperatures below 4.4°C adversely affected chum eggs.
Schroder et al. (1974, cited by Koski, 1975} reported that
chums in Washington exposed to water temperatures below
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1.5°C during the early stages of errt>ryonic development
experienced 1 over percent su rvi va 1 to emergence. At the
chum salmon hatchery at Clear Air Force Station in i~terior
Alaska, Raymond (1981) reported that the percent mortality
(fertilization to emergence) for one year when incubation
temperatL1res ranged from 2.0 to 4.2°C was 38%. The fol-
lowing year, incubation temperatures were raised about 1°C
to 3.6 to 4 soc and the percent mortality declined to 18~.
It appeared that the increase in temperature was largely
responsible for the decline in mortality. The lower water
temperatures of the hatchery a 1 so caused a de 1 ay in
emerqence of about seven weeks beyond that of wild fry of
the same stock in the Delta River which had incubated at
about 3.9°C. Kirkwood (1962) stated that chum salmon eggs
in Pri nee Wi 11 i am Sound which had not hatched before the
colder months of January and February did not survive. He
found that chum eggs in the upper i ntert ida 1 zone of a
streambed had a higher survi~at rate than eggs above the
influence of the tides and attributed this to the slightly
warmer temperatures of the former areas.
For the water temperature range (! tream water temperature
not i ntra~rave 1 temperature} obsP.rved itt Hooknose Creek,
B.C. (0 to 14°C) Hunter (1959) c;..,uld not demonstrate an
effect ~Jf temperature on the percent survival (eggs in
female to fry emergence) of chum salmon.
Stream temperatures very close to freezing may not pose any
problem as long as there is a sufficient flov of slightly
warmer springwater or groundwater through the incubation
art~a.
Chum salmon, particularly fall chums, actively choose such
areas (Kogl, 1965; Levanidov, 1954; Morrow, 1980; Sana,
1966}. Springs in certain rivers of Kamchatka maintain
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water temperatures in chum salmon spawning gravels of 1°C
even though tie air temperature may be as low as -50°C (Sano
1966). The same i s t r ue in the Chena River (Kogl, 1965 ) and
the Delta River (Francisco, 1977).
McNeil and Bailey (1975) suggest that water temperature
during aquaculture of Pacific salmon be kept above 4.4°C for
at least 10 days after fertilization, and preferably for 20
to 30 days. After this period, the eggs can tolerate
temperatures as low as 0°C as long as the water does not
freeze. Exposure of salmonid embryos to water temperatures
below 4.4°C prior to gastrulation and before closure of the
blastopore can lead to high mortalities (Koski, 1975). Bell
(1973) recommended an incubation temperature for chum salmon
of 4 .4 to 13.3°C.
The time to hatching of chum salmon eggs r anges from about
50 da ys at 7.0 to 15.0°C to about 130 days at 0.0 to 5.0°C
(Bakkala, 1970 ).
Koski (1975) found that annual water temperature variations
in a Washington str~am significantly alters the timing to
chum fry emergence from th ~ substrate. During a three year
period, the t i me to 50 % emergence varied up t o 30 days.
2 . Stream Flow, Current Ve l ocity, and Water Depth
The flow of water in the stream channel is important to
incubating embryos in promoting an ade qu atP. i ntragravel flow
and in protecting the substrate from freezing temperatures.
Hea vy mortal i ty of embryos can occur during periods when
there is a re l ative ly high or a relatively small discharge .
Flooding can cause high mortality by eroding eggs from the
r edds or by depositing fine sediment on the surface of the
redds whi c h can reduce permeability or entrap emerging fry
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(Bailey, 1g64; Dill and Northcote, 1g1o; McNeil, 1g66;
Rukhlov, 1g6gb; Strekolova, lg63; Thornsteinson, 1g65b;
w; ckett. 1958).
Low discharge periods can lead to desiccatiJn of eggs, low
oxygen levels, high temp~ratures, or, during cold weather,
freezing (Neave, 1953; Levanfdov, 1954; McNeil, 1g66; Sana,
1966; Wickett, 1g58).
McNeil and Bailey (1975) recomme~d a stream gradient at
the incubation area of 0.2 to 0.51, a water depth greater
than 15.2 em, a current velocity in the surface water of
30.5 to g1,4 em/sec and a flow of O.J6 m3Jsec per linear
meter of width for the aquaculture of Pacific salmon.
The water depths and current velocities giv~n in the section
for spawning adults apply in general to the intragravel life
history stages as well, although bott'l decline in many
streams during the freezing months, Water depths over chum
redds in some tributaries of the Columbia River ranged from
7.6 -43.2 em and had an average of 21.6 em (Burn~r, 1g51}.
Water depths over summer chum redds in the Beshenya River in
Siberia ranged from 16 -104 em and over fall chum redds
ranged from 3-34 em (Smirnov, 1947).
Lister and Walker (1966) founli an inverse relationship
between chum salmon egg to fry percent survivai and the peak
dai·•y discharge of the Big Qualicum River, British Colunmia,
during the incubation period. No relation between percent
survival and the minimum discharge was apparent from their
data. Wickett (1958) showed a direct relationship between an
index of production (stock + 4 years divided by spawners
times 100} and the discharge of Hooknose Creek, B.C •• in
November. Hunter (1959) could not demonstrate an effect of
stream discharge of Hooknose Creek I B.C. I on chum salmon
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percent survival (eggs in female to fry emergence). On the
Noatak River, low returns of adult chums appear to be
correlated with high water levels {rainfall) during the
brood year (Frank Bird, personal communication). Gallagher
(1979) found a negative correlation between stream flows
during the incubation period and brood year returns in the
Puget Sound area over a fourteen-year period; law flows were
correlated with good returns. Based on these seemingly
contradictory results, it would appear that the effect of a
varying discharge strongly depends upon the particular
stream being examined.
3. Substrate Composition
Infonmatfon on substrate size composition of chum spawning
grounds was presented previous 1 y. The size and shape of
substrate particles and the particle size distribution
influence many parameters which are of importance to the
successful incubation of embryos and alevins. Some of these
parameters. several of them interrelated, are:
permeability, porosity, flow of intragravel water, dissolved
oxygen concentration, concentration of waste metabolites
such as carbon dioxide and a111110nia, the annoredness or
resistance to abrasion of the substrate surface and the
degree of imbeddedness of larger sized particles in the
substrate surface.
Dill and Northcote (1970), using experimental containers,
found a survival to emergence for chum salmon of
approximately lOOt in large gravel (5.1 to 10.2 em) and 31~
in small gravel (1.0 to 3.8 em). They judged that the lower
survival rate in the smaller particle did not result from a
low dissolved oxygen content, but was probably a result of
silt action or entrapment of fry. They found no significant
effect of gravel si~e or egg burial depth (20.3 and 30.5 em)
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on the timing of emergence or on the condition of fry at
emergence.
Koski (1975) showed that there was a highl.v significant
inverse relationship between the percent sand content (fines
less than 3.327 mm but greater than 0.105 mm) of substrate
and the percent survival to ~mergence of chum salmon. An
increasing sand content also resulted in an earlier
emergence, increased the percentage of premature fry
emerging from the substrate, lowered the percent yolk
conversion efficiency, and tended to produce fry of a
shorter length. Koski could not demonstrate a relationship
between the percentage of silt (fines less than 0.105 n~) in
the substrate and percent survival to emergence, primarily
because the effect of silt, which ranged up to about 501,
wa~ masked by a changing percentage of sand. However, silt
contents of 45~ or greater did have an adverse effect on
percent survival.
Rukhlov (1g69b), working on Sakhalin Island, also noted an
increasing egg mortality rate with increasing sand content
above 14~. He found that redds with a sand content of 22~
or greater had a survival rate of so: or lower. Rukhlov
reported that average survival in the Tym• River was 85~
where the sand content was 10~ and 65~ where the sand
content was 181. Kol'gaev (1962, cited by Kogl, 1965) noted
a positive correlation between the duration of incubation
and the particle size of the substrate when chum eggs were
incubated in an experimental container with coarse sand.
Thorsteinson (1965) stated that spawning grounds with
percent fines {particles Tess than 0.833mm) greater than
12.7~ are of poor quality for chums because of reduced water
penneabil ity.
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In hal:chery situations, the absence of a gravel substrate
apparently leads to a less desirable habitat for Pacific
salmon alevins (Bams, 1969; Emadi, 1973; McNeil and Bailey,
1975}. Emadi (1973}, in an experimental situation,
discovered that chum alevins kept on smooth substrate had a
higher percentage of malformed yolks (30%) than those kept
or. gravel of 2 to 3 em diameter (OX) at a water velocity of
100 cm/hr and a temperature of l2°C.
Sedimentation during the incubation period can be. a major
source of egg morta 1 ity ( Levani dov, 1954, Heave, 1953;
McNeil, 1966; Rukholov, 1969b; Wickett, 1954). Siltation
one year in the River Khor of Siberia killed 100~ of the
embryos in 42 chum redds (Levanidov, 1954).
4. Intragravel Flow and Permeability
The rate of flow of intragravel water is an important
variable affecting the survival and fitness of embryos and
alevins. Water flowing through the gravel supplies oxygen,
removes waste metabolites such as carbon dioxide and
aiTII10nia, and is important in regulating temperature . The
rate of intragravel flow is affected mainly by the rate of
stream flow, stream gradient, bottom configuration,
substrate surface irregularities, degree of substrate
surface annoredne-ss, and penneabil i ty of the substrate which
depends strongly on the substrate particle size distribution
{Sheridan, 1962; Vaux, 1962). Johnson (1980) showed
experimentally that the penneabil ity of spawning gravels
decreases with an ;,,creasing percent of fines (particles
less than 0.5 rrm). Different methods of measuremen t of
intragravel flow by dH·:erent investigators has resulted in
order of magnitude differences in results.
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Reiser and Bjornn (1979) reported that the percent survival
of chum to the migrant fry stage increased from about 2~ to
15% as the penneabi 1i ty of the streambed gravel increased
from about 30 to 155 em/hr.
Wickett (1958) surveyed the grave 1 permeability of two
streambeds in British Columbia over several years and found
that chum salmon percent survival (egg in female to fry) in
the streambed with an average pennea.11lity of 1914 cm/hr was
1.2~, while in the streambed with an average penneability of
4035 cm'hr, it was 7.6~.
Kol •gaev (1962, cited by Kogl, 1965) found that the survival
of chum eggs in coarse sand in an experimental apparatus was
directly related to the volume of flow through the sand.
Wickett (1957) stated that the minimum intragravel apparent
velocity for the survival of pre-eyed eggs is 10 cmthr; 50
cm/hr is desirable.
McNeil and Bailey (1975) recommended an intragravel apparent
velocity of at least 200 cm/hr for the aquaculture of
Pacific salmon.
5. Dissolved Oxygen
The supply of dissolved oxygen is of critical importance to
eggs and alevins; a low supply of dissolved oxygen leads to
an increased mortality or a decreased fitness (Alderdice et
31 •• 1958; Koski, 1975; Wickett, 1954, 1957 and 1958).
Apparently, a rate of supply of oxygen less than optimum for
survival and fitness occurs frequently in nature. It is the
rate of supply of dissolved oxygen which is important to the
egg or alevin, not the actual concentration of dissolved
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oxygen in tt.~ water (Daykin, 1965). The rate of su~ply to
the embryos and a 1 ev ins is influenced primarily by the
dissolved oxygen {DO) concentration of the source water and
the rate of water flow through the gravel. Because of a
certain upper limit on possible velocities of intragravel
flow, there is a certain minimum DO concentration below
which it would be difficult to maintain an adequate rate of
oxygen supply. There are also other biological and chemical
oxygen demands within the gravel which compete for the
available supply. The relationship of particle size and DO
concentration was noted by Koski (1975) who found a
si gnificant inverse relationship between minimum observed DO
concentration and maximum observed percent sand and silt in
spawning gravels in Washington.
Water saturated with dissolved oxygen may be regarded as the
optimum condition for eggs and alevins (Alderdice et al.,
1958). The concentration of oxygen in oxygen-saturated
fresh water depends mainly on the temperature.
Oxygen requirements vary considerably for different stages
of development of eggs and alevins. Temperature also has a
strong eJfect on the rate of oxygen consumption.
Often, the oxygen requirements can be met by a fairly low
concentration of dissolved oxygen, about 2 mg/1, as long as
the rate of flow of intragravel water 1s sufficient {Kogl,
1965; Levanidov, 1954). Wickett (1954) measured the
apparent velocity ~f water 12 in~hes down in the gravel with
a standpipe. He detennined that a dissolved oxygen
concentration of 4 p.p.m. at 8°C (50% saturation) would just
maintain full metaboli sm of a pre-eyed chum egg providing
intragravel water flow to the egg is sufficient.
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Insufficient oxygen supply results in an increase in the
mortality rate or decreased fitness in the surviving embryos
or larvae which may be manifested as premature or delayed
hatching or emergence, production of monstrosities,
decreased length, or less efficient yolk conversion
{Alderdice et al., 1958; Koski, 1g75). As with many
envi ronmenta 1 parameters, the duration of the period of
stressful levels of oxygen and the interaction of other
stresses is important.
a. Oxygen consumption rate
Wickett {lg54) measured the oxygen consumption of chum
salmon eggs and found a rate of 0.0001 to 0.000~
mg/egg-hour for pre-eyed eggs 0 to 12 days old,
(temperature 0.1 -8.2°C), 0.0002 to 0.008 mg/egg-hour
for faintly eyed eggs 67 to 85 days old (temperature
0.1 -4,goc), and 0.0002 mg/egg-hour for eyed eggs 103
days old (temperature 5,g-6.1°C).
Alderdice et al. (1g58}, using chum salmon eggs at four
developmental stages at l0°C, reported oxygen
consumption rates from o.ooog3 mg/egg-hour for middle
stages to 0.00521 mg/egg-hour for later stages. They
also showed that, while the oxygen consumption per
embryo increases with age because of increasing embryo
mass. the actual rate of oxygen consumption per gram of
embryo initially decreases and then more or less levels
off with increasing age. An early developmental stage
had a rate of about 0.006 mm 3/g-min, mid-stages were
about 0.040 mm 3tg-min, and late stages were about 2.100
rrm 3tg-mfn.
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Lukina (1973 measured oxygen consumption rates of
0.00015 to 0.00951 mg 02/egg-hour at 3.5 to 4.5°C for
chum embryos and 0.0132 mg 02/larvae-hour at 5 to 6°C
for chum larvae.
b. Effect on survival
Critical oxygen levels (the lowest concentration
at which respiration i s just satisfied) for chum
salmon eggs have been found in laboratory
experinents to range f rom 0. 72 mg/1 for early
develop.-nental stages to 7 .19 mg/1 for later
developmental states at 8 to l0°C (Alderdice et
al ., 1958; Wickett, 1954). Whether or not these
concent""at ions are sufficient to satisfy the
oxygen demand of the eggs depends on the flow rate
of the intragravel water . Alderdice et ~1. (1958 )
reported that the incipient medi in lethal level
for dissolved oxygen ranged from about 0 .4 mg/1
for early developmental stages to 1.0 to 1.4 mg/1
previous to hatching.
Levanidov (1954) stated that t~e fall chum of the
River Khor in Siberia have an ability to adapt and
to develop under a continuous oxygen deficiency.
Oxygen cJncentration in one spawning gravel
dropped to 4 mg/1 (30% saturation) in March and
the oxygen content ,.f an'lther area which had a
strong outflow of groun d water dropped to 2 .5 to
3.0 mg/l (.20 to 25'X saturation) at the end of
February. The rate of egg surviva 1 and fry
development was high at both of these sites. A
third site. where the surviva l rate of eggs was
also good. had an oxygen concentration in
mid-December of 2.0 mg /1 . Experiments conducted
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by levani~ov (lg54) showed that fall ch~m eggs 10
to 15 days after fertilization can survive two
hours exposure to water of zero oxygen content.
Eggs which are a fev days from hatching can
survive (two hours?) exposure to water with an
oxygen concentration of 0.45 to 0.50 mg/1. larvae
24 hours after hatching can survive an oxygen
concentration of 0.28 mg/1.
Disler (1g51) stated that the DO concentration of
some chum redds in the Bira River of Siberia doe!.
not exceed 3-4 mg/1. In Holckaido streams,
Kobayashi (1g68) found oxygen saturation values cf
31.3 to 73.9% in the intragravel waters.
Intragravel DO concentration in the Chena River
during incubation of chum eggs ranged from 0.6 to
6.5 mg/1 (Kogl, 1965). There were low survival
rates at the lower concentrations 1nd high
survival rates at the higher C'lncentrat~ons.
Koski (1975) found that the rate of chum salmon
survival to emergence at a site in Washington wa!
significantly correlated with DO concentration an.f
that survival decreased rapidly at 00
concentrations below 3.0 mg/1. DO levels in the
i ntragrave 1 water of Indian Creek ( Kasaan Bay)
have an annua 1 range from 0. 0 to 12. 7 mg/1
{McNeil, 1962). long dry periods in southeastern
Alaska c~n lead to low intragravel DO levels as a
result of reduced stream and intragravel flow.
Such ct .. ditions in Traitors River one year led to
an intragravel 00 level of 1. 77 mg/1 which
resulted in a high mortality rate of embryos
{Mattson et al.. 1964). In another situation, a
low water permeability at one tide level in the
intertidal spawning ar~a of Olsen Creek led to a
low intragravel DO value (3.6 mg/1) and a low
percent survival to emergenc~ (Bailey. 1964).
Wickett (1954 and 1957) stated that chum salmon
eggs, in the absence of other stresses, have a
letha 1 1 imi t of about 2 mg/1 at 5° C. On the other
hand, both Levanidov (1954} for the Amur K1ver of
Siberia and Kogl (1965) for the Chena River of
Alaska reported good survival rates in gravel with
DO concentrations of 2 mg/1 as long as there was a
strong flow of groundwater or springwater. Kogl
(1965) was not able to demonstrate a relationship
between egg survival and DO concentration at
concentrations greater than 2 mg/1.
Cheyne {1941, cited by Nicola et al., 1966)
reported tllat chum salmon eggs developed at about
the same rate in waters with DO levels from 3.6 to
7.6 mg/1. Wickett (1957} stated that a 00 level
of 5 mg/1 is low for chum eggs and that a level of
13 mg/1 is desirable . Based on experimental
studies, Lukina (1973) indicated that a DO
concentration of 6 to 8 mg/1 was the most
favorable for the tievelopment period of chum
embryos and larvae at a temperature of 4·8°C.
McNeil and Bailey (1975) suggest, for aquaculture
of Pacific salmon, that alevins be provided water
with 00 concentration of at least 6 mg/1 o2 • Chum
salmon embryos and alevins in redds in the
intertidal zone of streambeds may be exposed to a
lowering of DO lev~ls during high tide. At the
2.1 m tide level in Olsen Creek, the DO level in a
redd dropped from about 7 mg/1 at low tide to 4.5
mg/1 at high tide (Thorsteinson 1965a).
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c. Effect on fitness
Kogl (1965) reported that low DO concentration in
the intragravel water of the Chena River affected
the size of chum alevins. The dry weight of
alevins decreased from about 2.3 mg at a site
where the DO concentration had averaged 4.1 mg/1
to about 0.1 mg at a site where the DO
concentration had averaged 2.1 mg/1. Cheyne
(1941, cited by Nicola et al., 1971) stated that
chum eggs held at a DO level of 7.6 m/1 resulted
in iarger fry than those held at 3.6 mg/1.
Conversely, Koski (1975) was able to show little,
if any effect of DO concentrations less than 6.0
mg/1 on mean fry lengths and weights at emergence.
Alderdice et al. (1958) found that early
developmental stages of chum salmon eggs (12 days
after fertilization) could survive seven days
e~posure to a DO concentration of 0.3 mg/1 but.
upon hatching. had a high percentage of
monstrosities (deformed alevins). The abnormality
took the form of a shortening of the vertebral
column posteriorly. Eggs at a later developmental
stage (22 day!; after fertilization) did not
to 1 erate seven days exposure to the same DO
concentration (0.3 Mg/1) and exhibited 1001
mortal fty.
Alderdice et al. (1958) further showed that seven
days exposure to oxygen levels below saturation
produced delays in the mean hatching rate of chum
salmon eggs at 10°C. Early and middle
developmental stages were most affected. Eggs of
the most advanced developmental stage showed the
opposite effect, premature hatching. Koski (1975)
-40-
found that the low dissolved oxygen (3.0 mg/1 or
less) during the incubation period appeared to
delay the timing of chum fry eme-.-gence from the
gravel of spawning beds in Washington. A delay in
emergence could place the fry in an unfavorable
situation with regard to feeding, predators, or
temperature. Beall {197"), using the spawning
channel at Big Beef Creek, Washington, incubated a
group of chum eggs in a trough with gravel and
another group in an open trough, with the result
that the fry produced in the open troug~ were, on
the average, shorter by 1 to 2 mm and lighter by
0.04 g. Beall found that the predation rate by
coho salmon (Oncorhyncus kisutch) was
significantly higher on the smallP.r group of fry.
6. Sa 1 i nity
Seawater incursion on chum salmon embryos and alevins
in gravel in or near the intertidal zone can have an
adverse effect. Presumably, 100J freshwater is ideal
for chum salmon emryos and alevins, but Rockwell
{1956) suggested that, based on laboratory experiments,
hatching ~f eggs may be slightly improved by a small
percentage of seawater mixing with freshwater. He
found that chum eggs and larvae survived for several
days in seawater of up to 30 parts per thousand (ppt}
salinity, that they could survive the higher salinities
at Tower temperatures, and that larvae withstood higher
salinities than eggs. Important variables affecting
survival at various salinity concentrations were time
of exposure and temperature. For eggs in early
developmental stages, exposure ~o 6 ppt salinity had no
effect; however, exposure to 11.6 ppt significantly
ret~rded develo~nt and led to a 67~ mortality. Fully
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developed eggs had a SOt survival rate at salinities of
6.0 to 8.5 ppt through hatching.
Data obtained by Kashiwagi and Sato (1969) indicated
that percent survival of eggs to hatching ~as inversely
related to percent seawater concentration. Eyed eggs
had a lOOi hatching rate in water of about 9 ppt
salinity; this decreased to 75~ for 18 ppt, 50S for 27
ppt, and 25S for 35 ppt. The time of incubation was
increased from about 10 days in water of 0 to 18 ppt to
about 14 days in water 27 to 35 ppt. However, almost
all of the alevins which did hatch from eggs exposed to
water of 9 to 35 ppt died within a few days. Alevins
which were hatched in freshwater and then exposed to
seawater mixtures could survive a salinity of 9 ppt.
but died at higher salinities.
Shepard (1948) demonstrated that chum alevins, given a
choice between two flows. showed a preference for
stronger current velocities. Given a choice between a
seawater flow and a freshwater flow, they chose the
1 atter. Salinity was more important than current
velocity in influencin~ the alevins.
In the intertidal area of streambeds of Olsen Creek
(3.7 m tide) there is constant residual intragravel
salinity of 3 ppt from the 2.1 m level on down
(Thorsteinson, 1965a). Salinities up to 25 ppt have
been observed at all levels. The salinity of the
i~tragravel water fluctuates with the tide. At the 2.1
r. level, there was a 7 ppt increase at high tide. The
egg to fry survival of chum salmon was 40 -50S fn the
upper section of the intertidal zone and 10 -1St in
the middle section. Chums do not use the lower
section. The mortality rate is influenced by oxygen
-42-
and temperature differences as well as by salinity
differences.
McNeil and Bailey (1g75) suggest that Pacific salmon
eggs can tolerate salinities of 0 to 9 or 11 ppt and
that chum eggs can survive periodic inundations by
water of 15 to 30 ppt as long as there is freshwater
flushing in between the seawater inundations.
7. Chemical Parameters
There is a limited amount of data available on the
deshable levels of soluble substances present in
fntragravel water, or on their effects on chum salmon
embryos and alevins.
The pH of Chena River intragravel wcter at chum redds
was 6.5 (kogl, 1g65); this is similar to the Amur River
in Si berfa (Levani dov, 1g54) and a Hokkai do stream,
which was pH 6.3 to 6.5 (Kobayashi, 1968). McNeil and
Bailey (1975} suggest that pH 6.0 to 8.0 is a desirable
range for Pacific salmon.
McNeil (lg62) measured intragravel C0 2 levels in some
streams of sou thea stem A 1 as ka of 2 to 24 mg/1.
LeV1 n 1 dov (1954 ) reported that the C0 2 con tent of
groundwater in the Amur River was about 25 to 30 mg/1.
He found that concentrations greater than 20 to 25 mg/1
inhibited the development rate of chum salmon eggs.
Alderdice and Wickett (1958) observed that co 2
concentrations greater than about 10 mg/1 decrease the
percent survival to hatching. The uptake of oxygen is
inhibited at C0 2 concentrations greater than 125 mg/1.
Koski {1g75) measured co 2 concentrations of up to 48
mg/1 in intragravel water in a Washington incubation
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area. but the effect of this concentration on survival
was not studied. Cisler (1951) found co 2 levels in the
intragravel water of the Bira River of 16 to 20 mg/1
and Kobayashi (1968) found levels of 17.8 to 25.2 mg/1
in a Hokkaido stream. McNeil and Safley (1975)
reconnended that the C0 2 concentration in Pacific
salmon hatcheries be kept below 5 mg/1.
The alkalinity c,f intragravel water in chum salmon
redds in the Chena River was 86 to 103 mg/1; iron
content was 0.1 to 0.3 mg/1 (Kogl, 1965). Good
survival rates were noted at these sites.
Ammonia is a waste product of metabolism which could he
highly toxic to embryos and alevins if not removed by
intragravel flow. McNeil and Bailey (1975) recommend
that Pacific sa 1 man hatcheries keep anmon i a 1 eve 1 s
below 0.002 mg/1.
Hydrogen sulfide {H 2S) is produced by sulfur-reducing
bacteria in waters of low oxygen content and 1s highly
toxic to embryos and alevins. Koski (1975) found H2S
levels in intragravel water of a spawning channel in
Washington of 0.4 to 0.5 mg/1 but did not measure the
effect on chum embryos or alevins. McNeil and Bailey
(1975) suggest that H2s 1 eve 1 s in Pacific sa 1 mn
hatcheries be kept below 0.5 mg/1 in acidic waters.
Springs often have higher levels of C0 2 and H~S and
lower levels of DO thar. stream water. In some areas,
these substances may be at such levels that they are
detrimental to the incubation of chum eggs and alevins.
-44-
Cisler ,1953) stated that direct su~)1ght i~ fatal to
chum salmon embryos. Indirect sunlight can retard the
rate of development, affecting earlier stages more than
later stages of development (~oin, 1954).
0. Emergence and Downstream Migration of Fry
This section will consider the period between the time of fry
emergence from the gravel and the time the fry enter estuarine
waters. Most chum fry begin the seaward migration shortly after
emerging from the gravel in the spring and essentially all have
left freshwater by mid-summer. Habitat requirements during this
period center on favorable temperatures, cover from predators,
availability of food, and eventually, availability of seawater.
Down.stream migration occurs as a result of the re·sponse of the
fry to physical and chemical parameters including light and
photoperiod, current, temperature, salinity, and objects in the
environment (Hoar, 1956 and 1957). The outmigration from streams
in British Columbia results from a combination of active swimming
and displacement at night when visual orientation is diminished.
Streams in northern Alaska or Siberia have very little darkness
during the time of outmigration. Response to the various
environmental stimuli (phototaxis, reheotaxis) can change. An
increase in temperature can change a positive rhl!otaxis to a
negative rheotaxis. Mihara (1958, cited by Bakkala, 1970) found
that chum fry in Hokkaido streams displayed a negative rheotaxis
and moved downstream quickly when the temperature reached 15°C.
This was interpreted as an adaptive response to avoid the high
summer stream temperatures.
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i. Temperature
Bams {1969) demonstrated a very close correspondence between
water temperature and the numbers of chum fry emerging from
the gravel in a lat~ratory in British Columbia. There were
about eight fry emerging per hour at 5.3°C and about 48 per
hour at 7.2°C.
Emergence and outmigration of chum fry from the Delta River
has been observed at temperatures of 3.0 to 5.5°C, (Raymond,
1981). The temperature range observed during the peak of
chum fry outmigration from the Salcha River was 5 -7°C
(Trasky, 1974}.
Downstream migration of chum fry in Hokkaido begins in early
April at water temperatures of 2.0 to 3.0°C but does not
reach a peak until early to mid-May when water temperatures
are 6.0 to 10.0°C (Sana, 1966). There are other factors as
well, such as the rise in water level from melting snow in
May. Semko (1954, cited by Sana, 1966) measured water
temperatures of 4.6 to 5.5°C during the downstream migration
of chum fry in April, May and June in the Bolshaia River.
levanidov (1g54) reported tha~ chum salmon fry in the Amur
River cf Siberia ~an survive water temperatures up to about
15°C but that they prefer temperatures of 8° to l0°C.
Brett (1952) noted that young chum salmon (three to six
months after hatching) in t!xperimental tanks generally
avoided temperatures above l5°C and preferred temperatures
of 12 to 14°C. He found a lower lethal temperature for
young chums of ooc and an upper lethal temperature of
23.8°C. The young chums could acclimat~ to 23°C, but not to
24°C. In further experiments, Brett and Alderdtce (1958)
showed that the ultimate lower lethal temperature is 0.1°C.
-46-
Bell (1973} g1ves the migration temperature range of
juvenile chum salmon as 6.7 to 13.3°C.
McNeil and Bailey (1975) state that Pacific salmon fry will
grow in the temperature range 4.4 to 15. 7.,C. but that the
ideal range is 10.1 to 12.9°C. Below 10.l°C, growth i~ slow
and at 15.7°C and above, there are problems with diseases
and other stresses. Fry can tolerate temperatures of 20° to
22°C for limited periods but prolonged periods at 18.5°C and
above cause severe stress and prolonged periods at 21.3°C
and above can cause mortality.
In experiments examining the culture of juvenile chums from
Oregon in heated seawater, Kepshire {1976) found that growth
was similar at 12.9, 15.7, and 18.5°C, but gross food
conversion efficiency decreased with an increase in
temperature. There was good survival at these temperatures,
but not at 21.3°C.
levanidov (1956, cited by Bakkala, 1970) found in the
laboratory that the growth rate of Amur River chum fry was
three percent of body weight per day at 8°C and five to six
percent at 14 to 20°C. Temperature can also modify
behavioral patterns. Keenleyside and Hoar (1955) found that
most chum fry in an experimental situation exhibited a
positive rheotaxis (swimming into the current) at 6 to 9°C
I and a negative rheota x 1 s ( swimming wi th the current ) at 10
to 13°C. A higher percentage exhibited negative rheotaxis
at 17 to 19°C and an even higher percentage did so at 18 to
21°C.
2. Cover
Chum fry in short streams migrate mainly at night and seek
cover in the substrate during the daytime if the journey is
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not completed in one night (Neave. 1g55}. Observations by
Hoar {1g56) indicate that chum fry. after schooling, use the
protection of the schools and no longer seek protection in
the substrate. Saltwater increases the schooling tendency
of the fry (Shelbour,, 1966).
Predation can be a major source of mortality (Hunter, 1g5g;
Kirkwood. 1962). Hunter (lg5g) estimated that predation
ranged from 23 to 85 percent in Hooknose Creek, B.C.
Predators in Hooknose Creek include: coho sa 1 man smo lts.
Aleutian sculpin, prickly sculpin, Dolly Varden char,
cutthroat trout and steelhead trout. Kingfishers,
mergansers, and certain mammals have also been menti1ned as
predators (Scott and Crossman, 1973).
3. Substrate
In addition to providing cover from predators, gravel
substrate is appare'ltly necessary to maintain orientation
for those fry that still have a yolk sac upon emerging. 1n
an experimental sittJation, Emadi (1973) found that chum
alevins kept on a smooth substrate had a higher percentage
of malformed yolks (30S) than another group kept on a gravel
substrate of particles 2 to 3 em in diameter (no malformed
yolks), both in a current velocity of 100 em/hr. McNeil and
Bailey (1g75) have also pointed out that when Pacific salmon
are being artificially raised in hatcheries, the alevins and
fry nust be provided with a gravel substrate after
emergence.
4. Food
In short streams where the outmigratfon is completed in a
day or two, chum fry may feed very lit:tle, if at all. In
longer streams and river systems they do feed and gain
-48-
weight during the course of their seaward migrat1on. Fall
chum fry of the Amur River may spend up to a month feeding
in the spawning grounds (Levanidov, 1954). Chum fry from
the Noatak River apparently feed in the lower 10 km or so
until at least early August (Frank Bird, pers. conrn.). This
is also the case at the mouth of the Yukon River (W. Arvey,
pers. conm.). Benthh: organisms, mainly aquatic insects,
are t 11e dominant food species in freshwater. Prey species
include chi ronomid larvae, nymphs of stoneflys and mayflies,
caddisfly larvae. blackfly larvae, blackflies. mosquitoes,
mites, and thrips, ar.d other terrestrial insects.
Levanidov (195·1) ca 1 culated that there was enough food in
the River Khor to support 40 chum fry per square meter. He
found that the food requirements of fall chum fry amounted
to 8 to 10% of their body weight per day at 6 to soc and 17
to 20% of th~ir body weight at 17 to 18.0°C. Le~an1dov also
states that certain chum fry in freshwater with high food
requirements would be unl ik.ely to survive more than two or
three weeks without food. Levanidov and Levanidova {1951)
calculated that chum fry in Lake Teploe consumed about 27"'
of their body weight daily.
Volov1k (lg68) estimated that chum fry in the rivers of
Sakhalin require 18.0 to 123.0 mg food per day, which was
5.2 to 9.0% of body weight.
s. Light
Chum fry exhibit responses to light (phototaxis) which aid
them from streams 1 n reaching the ocean during their
outmigration. Loss of visual orientation at night leads to
downstream displace,..~nt. An increasing daylength in the
spring affects the thyroid system which increase·s salinity
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6.
tolerance and preference and the development of a caoability
for osmoregulation (Baggerman, 1960~ Shelbourn, 1S56). Hoar
(1957) demonstrated that schools of fry sh0w a preference
foA light, but exhibited no marked response to changing
li~~t intensity. Strong light is avoided (Neave, 1955).
Salinity
The preference of chum fry for higher salinities and their
ability to osmoregulate in seawater increases with
increasing age (Baggerman, 1960; Houston, 1961). At the
onset of downstream migration, chum fry in an experimental
s i tua ti on preferred sa ltwa t~r to freshwater { Baggerman,
1960).
Chum sa 1 mon fry that have absorbed their yo 1k sac can
tolerate full strength seawater {Kashiwagi and Sana, 1969;
McNeil and Bailey, 1975; Weisbart, 1g68). Salinity
tolerance increases with age after hatching an~ is related
to an increasing capability for osmoregulation (Kashiwagi
and Sano, 1969}. Kashiwagi and Sano (1969) found that chum
salmn 60 days after hatching and older can survive an
abrupt tr~:1s fer to 100~ seawater ( 35 ppt). Converse 1 y,
Levanidov (1952) found that fry captured during their
seaward migration in the Khor River could survive transfer
to water with salinities of 7.0 ppt and 17.1 ppt, but 8 out
of 10 died in a salinity of 35.0 ppt. The period of
acclimitization was characterized by an increase in the rate
of respiration. Houston {1959) noted that an abrupt
transfer of chum fry to seawater of 22 to 24 ppt. reduced
their cruising speed to about 68~ of the normal rate. After
about 36 hours, the cruising speed was l"learly back to
normal, which ranged from about 19 em/sec for 3.9 em long
fry to about 36 em/sec for fry 4.9 em long. Houston
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suggested that this depression in cruising speed might be
related to osmoregulatory adaptation.
Shepard (1948} conducted experimental studies on the
interaction of current velocity and temperature in relation
to salinity preferences of chum fry. The fry were initially
presented with a choice between two flows. When one flow
was freshwater and the other was seawater, they chose the
seawater flow, regardless of relative strengths of the flows
(flow difference of about 1.3 em/sec). When both flows were
freshwater and one was l0°C and the other 12.5°C, they chose
the l0°C flow (they had been acclimated to 10°C). If the
12.5°C flow was seawater, they still initially preferred the
l0°C freshwater flow, but gradually changed to the 12.5°C
sa 1 ine flow . These experiments demonstrated that no one
~arameter has a definitive effect and that the interaction
of the thr~e parameters plays an important role in governin~
the seaward migration of fry.
Chum salmon fry ~poear to have a physiological requirement
for seawater wi tl•; ·
normal developmer i
access to the oc""
raised experiment •
this required e• ·
Chum fry in natu•
three or four months after emergence for
fhis is not related to their need .for
for normal feeding. They have been
~J the adult stage in freshwater but
. y careful handling (Hoar, 1976).
• t erns whose access to the ocean was
restricted would pr .04u iy not survive.
7. Dissolved Oxygen
Levanidov (1954) stated that the threshold for the asphyx ia
of chum fry at l0°C occurs at a dissolved oxygen
concentration of 1. 5 mg/1. He found no change in the
feeding intensity, assimilation, or growth rate of fry over
the range 5 to 11 mg/1 . Lukina (1973) reported that 8-9
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mg/1 was the most favorable concentration fCir chum
fingerlings (temperature 8-10°C).
McNeil and Bailey (1975) suggest that the dissolved oxygen
level for Pacific salmon fry in hatcheries be kept no lower
than 6 mg/1.
The oxygen requirements of fry vary with the temperature.
Chum fry used 188 mm3o2;g body weight-hour at 8.6 to 9.0°C,
228 mm3 at 10.0°, and 445 mm 3 at 20°C and fingerlings used
144 mm 3tg body weight-hour at 9.0 to 9.4°C (Awa~ura, 1963
and levanidov, 1955; both cited by Ba~~ala, 1970). Volovik
(1968) found that chum fry about 4 em long used about 0.26
mg o2;g live weight-hour (8.0°C) and fry about 5 em long
consumed around 0.61 mg o2;g 1ive weight-hour (7 .3°C).
Oxygen consumption varied during different hours of the day.
8. Stream Flow and Current Velocity
In experimenta 1 chambers in which chum salmon fry were
presented with a choice between two channels with different
11 ~ ami na r•• fl ows , MacK i nnon and Hoar (1953 ) found that the
fry preferred 350, 500, 600 and 700 ml/min flows to a flow
of 200 ml/min; the greatest response was toward the 500
ml/min flow. In another experiment with 11 turbulent 11 water
f1 ow, the fry seemed to prefer flows of about 5,000 to
12,000 ml/min over either lesser or greater flows. They
always exhibited a positive rheotaxis.
Levanidov (1954) stated that optimum stream velocities to
support the feeding of fry in the Amur River are less than
20 em/sec.
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III. CONCEPTUAL MODEL OF HABITAT SUITABILITY
Suitability index curves in a conceptual fonm are presented for water
temperature~ stream depth, current velocity, dissolved oxygen
concentration and salinity in Figures 2 through 5. The suitability
index for each environmental parameter ranges from zero to one. An
index of one indicates an optimum or preferred level of that
particular parameter and an index. Gf zero indicates a completely
unsuitable level. These curves should not be construed as a graphical
presentation of actual data. Rather, they are intended to be
hypothetical models of the relationship between chum salmon and
certain environmental parameters. As with any untested hypothesis,
they must be tested and verified before being applied to any
particular situation. The curves are based on published and
unpublished data and on conversations with fishery biologists who have
worked with chum sallll)n. Both experimental laboratory data and field
measurements and observations were used. The published data base is
summarized in Tables I through IV.
A problem encountered in constructing the curves is that much of the
data in the literature concerning environmental parameters of chum
sallll)n habitat does not relate various levels of the parameters to
some measure of habitat suitability. Often, ranges of the parameter
are given based on measurements taken throughout the area occupied by
chums but there is no indication that one point on the range is any
better or worse than any other point in terms of habitat suitability.
A good example fs stream velocity measurements in spawning areas. A
measurement of spawner density fs needed from throughout the range of
stream velocities where spawners are found. This kind of fnfonmation
is rare in the literature. More useful data has been collected in the
case of measur~nts of dissolved oxygen concentration of intragravel
water, which are frequently linked with percent survival to emergence.
The curves are drawn usi~~ data f~ throughout the natural range of
the chum salmon. Although there are differences in habitat
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preferences and tolerances for different stocks in different
geographical areas or even in different streams of the same
geographical area, there is not enough data to support drawing
separate curves at this time. However, one must be aware that any
point on the curve~ especially toward either extreme, may be more
suitable or less suitable for a particular stock. It may even be that
the extremes are completely unsuitable. How far the stock d~Jiates
from the curve must be detennif'!l!d by field measurements and
experimentation with that particular stock.
A second precaution regarding use of the curves concerns the
interaction of various parameters. A given level of one parameter can
have a different effect on the fish as the level of another parameter
varies. For example, a dissolved oxygen concentration of 5 mg/1 may
be suitable at a water temperature of 5°C 1 but unsuitable at a
temperature of 20°C. ldeallyt given enough data, a separate dissolved
oxygen curve should be drawn for each of several different
temperatures. The overall suitability of any particular habitat is a
summation of interacting effects of many parameters.
A third precaution to consider is that the effect on the fish of less
than optimum levels of any parameter depends strongly on the duration
of exposure. Exposure to seawater may have little effect on chum
embryos in the upper sections of the intertidal zone of a stream where
they are exposed for short periods at high tide, whereas longer
exposure to seawater in the lower sections of the intertidal zone may
have a severe effect on the embryos.
Also, because d 1 fferent 1i fe stages have di iferent requ f rements,
information on embryos and alevins (Figure 4) should not be dr3wn as
one curve. Because of insufficient data, it was drawn this way and,
if nothing else, points out the need for more field measurements
during these life stages. In fact~ there should be different curves
for different ages of embryos and for alevins of different ages after
hatching. It has been shown that at least the oxygen requirements of
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chum salmon embryos and alevins vary considerably for different ages
(Alderdice 1!t al ., 1958).
Despite the fact that there is not enough known about the habitat
needs of the chum salmon to construct accurate suitab il ity index
curves, the ones that are presented in this section do serve a useful
purpose . Overall, they provide an indication of what would ma ke a
des i rab 1 e chum habitat and what ~u 1 d make a 1 ess des i rab 1 e habitat .
Also, although they a .·e general, they are probably specific enough to
show that there are differences in habitat needs among chum salmon and
the other species of Pacific salmon. Further, the process of
constructing these curves is beneficial in def i ning those areas where
more data is needed . Lastly, these curves can aid in the design of
experiments and samp l ing programs . As more data becomes available,
these hypothetical curves can be further refined. At the present,
they a r e only conceptual models which synthesize available data in the
fonn of graphs.
The overall suitabil i ty of a particular reach of a stream as habitat
for chum salmon is a sumrr~tion of all the suitability index c ~rves for
part i cular parameters and would i nclude other parameters fe r wtotich
there was not enough information to construct curves. For example, the
habitat suitability of a particular spawning substrate would be an
aggregation of suitability index (SI) curves for all parameters during
spal'.''ling and during incubation of embryos ~nd alevins. The most
likely aggregation function which would define a habitat suitability
index (HSI) for chum salmon would be that of limi ting factors. This
aggregation function takes the form:
HSI z minimum (SI 1 + SI 2 + ••• + Sin).
In other words, the habitat suitability index could not be any higher
than the lowest suitability index value of the spawning substrate for
any one parameter. If the dissolved oxygen regime of the intragravel
water has a suitability index value of 0 .5, then the HSI, or the
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ability of the spawning subst·ate to support the successful production
of chum salmon relative to other areas, could be no higher than 0.5
even if all other parameters (substrate si .t:e, water depth,
temperature. and so on) had optimum values (SI = 1.0).
Different environmental parameters have different degrees of impor-
tance in contributin~ to the quality of chum salmon habitat.
Parameters which were not drawn fn the form of curves for the upstream
migration phase of the life history because of insufficient
information were: current velocity, water depth, physical
obstructions, and turbidity. These four are all important factors and
perhaps, in Alaska, are more important than water temperature (which
is shown graphically), and should be included in any habitat
suitability index which might be constructed.
During -~he spawning phase of the life history, one of the most
important factors that determines the habitat quality of a particular
reach of stream is the composition of the streamed. It was not
possible to satisfactorily show the relationship between the particle
size distribution of the streamed and habitat quality in a two-
dimensional plot. Water depth, current velocity, and temperature are
also important factors during spawning.
The development of embryos and alevins in the substrate is perhaps the
most critical stage in the freshwater life history of the chum salmon.
One of the most fmportant factors during this stage is the
permeability of the substrate or the amount of water flow through the
substrate. This parameter has a strong influence on many other
parameters such as intragravel temperature, oxygen supply. and
concentration of waste metabolites. Unfortunately, there are few
measurements of penneability or intragravel flow in chum spawning
gravels and it was not possible to construct a curve. The particle
size distribution of the substrate is the most important factor
determining the permeability. Curves were not constructed for surface
current ve 1 oc i ty or stream depth. These two are of much 1 es 'i
-56-
importance than intragravel flow. Salinity, which was placed in the
form of a curv~, can be a major 1 imiting factor in intertidal
streambeds. Chemical parameters such as carbon dioxide concentration
and pH, which were not graphed, can be limiting factors in some areas
where upwelling ground water have values outside the limi ts preferred
by chums.
Curves were constructed for temperature and salinity, which were
graphed for the emergence and outmigration of fry stages. The amount
and duration of daylight is important but is ~ given factor for this
time of y~ar. Stream flow is also important, but the appropriate
discharge rate is unique to each stream .
In surmary, more infonnation is needed on severa 1 environmenta 1
factors before it will be ;>ossible to assign a habitat suitability
index to any particular area. More detailed infonnation on the
deficiencies of the data base is provided in Section IV .
-57-
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CHUM SALMON
ADULTS
1.0
i
• • .. c Oa6 -,.. -= • • -"i • 0
0 a 10 15 20
Water temperatur•
See text tor qualifications
for use of this curve.
(NOT •aCDMMI!NDI!D I' OR AP~LICA TION
TO INCIPIC WATIRSHID8 WITHOUT
1'11&.0 va•I•ICA TION)
21
•c
30
U.atreant
Migration
Figure 2. Cet~Ceptual model of relatlonattlp ttetween adult Cttum
aalmon and water temperature.
-58-
CHUM SAl...,.
ADUL?8
• • I
,.
5 • c
~
5 •
IPAWNING
t.O
...
'
See text tor qualifications
for u•• of these curves.
(NOT .ICOMMINDED '01t
A'~LICATtON TO lltiCIP'IC
WATIAaH.D8 WIThOUT ,.ILD
¥1JtiPICATION J
1-+--------~------~------~-• .. ... 1M . ........... ..
'
• • •• , .. 110
'
0 I tO 11 10 21 30
"s a L C •• 1 Ql 0 •• ••• ., nlatlo11elatp lletwee11 adult Ctlum •all" on and
•••• ... -. ......... aMI water telllpe•·ature.
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... ···MOtr •cytATION
• 31108 ..
.ll.lw.a
.. a&.
• • I
t
1.0
•
•
•
•
~•••-e•bryoa aiNI ••tna
.. , ...
See text for qualifications
for use or these curves.
t•OT ••cOMM.ND•D 'OR
A~LICATIOII TO INCI,.C
WAT.IIIH.D WnHOUT PIELD
YDIPIICATION J
-----~----::r"-~ /
/
/
,/
/
"' •
/
/
/
•
I
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•
,
Yuloa, dapo"cUne on etage
ef HWelo .. ottt. to•porature.
..., rate of now of lfttr•er•••• . ... ,
10 11 ......... ,_. . .,.
•
...,,.., af "o day ' ......... ...
lallllty tolerUHio
lllereaaoo wHII a .. : alao
4 ........ -..... ,.tur. ,. ,. .. .. .. ..
• 5 ......
t a,., P _tl., -t ,.. ••••• ._ "•••• c:.-••..._ ._, .... ,~
_SF -••r 5 •• '7 Jlr•t~ ........ d oay .. • •-Mtntklra.
.. 7 ••
-60-
CHUM SALMON
FRY
)I(
w
Q
!
>
~ -= CD c
!:: ::>
"
1.0
0.5
0
0
1 .0
0 .5
Emergence
11nd
outmigration
I "" ,. \
\
........ F•edlno
' and
growth
5 10 15 20
Water temperature,
See text for Qualifications
for use of these curves.
{NOT ~ECOMMENDED FOA
APPI..ICATION TO SPECIFIC
WATERSHEDS ',YITHC'! UT II'IELD
VERIFICATION)
25
•c
30
Sa.eltabtllty at high
aaUn,ty levels
dependa on age
and temoerature
0~---+--~----~--~--~--~--~~
0 10 15 20 25 30 35
Salinity, oot
Figure 5. Conceotual model of relationshi p between Chum salmon fry
and water temoefature and saJinl1y.
-61-
f
t
--------------------
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Table I.
Parameter
Tempe~ature c
Observed Values
4.4 -19.4
8.9 -14.4
5.0 -12.8
10.0 -16.7
8 -14
8.3-21.1
10.1
o. 25.6
0.1-20.0
7 -11
11 -12
CHUM SALMON
Adults -Upstream Mfgrat1on
Remarks
total range
range during peak migration
range during upstream migrati~
during peak ~f upstream migration
range for the species
optirrRJm value
lower and upper lethal limit
total range
range during peak of migration
Locatfon
Traitors Rher
Reference
Mattson and
Hobart ( 1962)
Tribs. of Kuskokwim AOF&G (198Da)
River
Anvik Rher Trasky ( 1974)
Hooknose Ck., B.C. Hunter (1959)
Bell (1973)
Northern Japan Sana (1966)
Memu River, Japan Sana and
Nagasawa (1958)
...
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0\ w •
Table II.
Parameter Obser~ed Values ..;....;;;..;...;;;.;..;=.;.;;..;.... __ ...;;.;;;;..;;..;
Temperature oc 6.5 -12.5
~13
ca. 12
4 -16
2.5
<15
4.4 -6.7
7.2 -12.9
1.8 -8.2
5 - 6
9.5 -12.8
9 -10
ca. 6
2 -9
6 - 9
CHUM SALMON
Adults -Spawning
Remarks
most spawning
inhibited spawning behavior
range during part o·f spawn1 ng
season
suggested criteria for species
'!lOSt spawning
mean temperatures, s ummer chum
sunmer t~hum
autumn chum
bottom water
Locat i on Reference
Terror and Wilson et al.
Kizhuyak R. (1'981)
Sashin Creek McNeil (1964)
Hooknase Creek, Hunter ( 1959)
British Columbia Neave ( 1966)
Big Beef Creek, Schroder (1973)
Washington
Big Beef Creek, Koski (1975)
Washington
tribs of lower Burner (1951 )
Columb i a
Bell {1973)
Sakhalin, USSR Rukhlov ( 1969a)
South Kuril Is 1. lvankov and
USSR Andreyev (1971)
Amur River, USSR So in (1954)
Amur River, USSR Sa no {1966)
Iski ~1ver, USSR Smirnov (1947)
Hemu River , Japa n Sano and Nagasawa
--
-------------------
Table II Cont'd. CHUM SALMON
Adults -Spawning
Panneter Observed Value Remarks Location ~eference
Water depth. 61.0 -91.5 partial range Delta River Franci sea { 1976}
em
5 -120 Chena River Ko91 {1965)
7.6 -10€. 7 Terror and Kizhuyak R. Wilson et al.
( 1981)
5.1 -72.2 range Tribs of lower Burner ( 1951)
25.4 mean Columbia River
2:18 Oregon streams Thompson ( 1972.
cited by Reiser
and Bjornn(1979)
I
0\ 18 minimum depth Oregon streams Smith ( 1973) .$:o
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15 -53 preferrP.d depth Western Washington Col11ngs ( 1974)
23 -46 recommended depth Wynochee River. Wash. Oec;champs et <!l.
(1966, cited by
Smith, 1973)
60 -100 preferred depth Amur River, Kusnetzov (1928)
Kamchatka, USSR
30 -100 sunmer chum My River, USSR So in ( 1954)
20 -110 Northern Japan Sana (1959,cited
by Bakkala,l970)
<20 -100+ range Memu River, Japan Sana and
20 -30 redds most abundant Nagasawa (1958)
<100 redds least abundant
Table II -Cont'd. CHUM SALMON
Adults -Spawning
Parameter Observed Values Remarks Location Reference
Stream 0 -60 Chena River kogl (lg65)
Velocity,
em/sec 0.0 -118.9 Terror and K1zhuyak Wilson, et al.
Rivers {1981)
46 -g7 Oregon streams Thompson ( 1g72,
cited by Rieser
& Bjornn (lg7g)
46 -101 range, V 12.2 em above bed Oregon streams Smfth ( 1g73)
7'J mean
I 21 -101 v 12.2 em above bed Western Washington Collings (lg74)
0\ c.n
I 31 -70 recommended range Wynoochee R., Wash. Oeschamps et al.
( 1966 , c fted by
Smith, 1973)
10 -80 My Rf v~r, USSR Sui n ( 1gs4)
20 -100 My River, USSR Strekalova
(1963)
10 -go most spawning Sakhalin, USSR Rukhlov (1g6ga)
10 -30 characterfs t1 c Bolshava R. Basin krokh1n and
USSR krogius ( 1937,
cited by N1cola
et a 1 . , 1 ':>66 )
<10 -35 range Hem R., Japan Sano and
15 -20 redds most dbundant Nagasawa ( 1958)
<10 & 30-35 fewer redds
-
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Table II -cont'd. CHUM SALMON
Parameter
Substrate
Particle
Size
Adults -Spawning
Observed Values
gravel 2-3 em in dfam ••
also use coarser stones and
even bedrock covered with
small boulders
Remarks
gravel mostly 1.27-127.0 em spawning grounds
with variable amount of fines
stones 1.3-13.0 em, coarse
sand, fine sand and silt
Location Reference
Alaska Morrow ( 1980)
Delta River Francisco (1976)
Hooknose Creek, Hunter (1959)
B.C.
gravel I > 2.69 em (98i) grave 1 size!; II and JJ 1 B f g 8'-. Creek Ouker (1977)
gravel II 1.35-7.61 em
(g7s)
gravel III 0.67 -2.69 em
(951)
grave~ IV 0.02 -0.57 em
(g6s)
1. 3 -10.2 em
selected by 751 of spawning
females; gravel I selected by
201. gravel IV selected by 51
substrate size criteria
for species
Bell ( 1973)
Table II -Cont'd CHUM SALMON
Adults -Spawning
Parameter Observed Values Remarks
Substrate gravel mixed with sand,
Particle small amount of silt
She
"sand" (0 .8 -52 .8%,
av. 12.0 -12.7 %)
spawning grounds
"gravel" (10.0 -50.1%,
avg. 33 .0 -45.91)
"shingle" (6.0 -84.1~,
avg. 41.4-44.4%)
>10 em (0.0-73.9S,
I avg. 0,0 -lO.OS)
0\ .....
I "sand" {14 -221) decreased survival
emergence
0.5 em (251), 0.6-3.0 em
(451), 3.1 em (301)
particles 0.5 em
(always 20S)
spawning grounds
particles 3.0 em
(25 -53S)
--
to
location
Amur River.
USSR
Sakhalin
Sakhalin
Northern Japan
MelllJ River,
Japan
Reference
Sotn ( 1954)
Rukhlov (1969b)
Rukhlov (1969b)
Sano (1959, ctted
. f Bakkala, 1970}
Sano and Nagasawa
( 1958)
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Table III CHUM SALMON
~ryos an1 Alevins -Intragravel Development
Parameter Observed Values
Temperature 9 0.5 -4.5 oc
2.0 -4.2, 3.6 -4.5
2.0 -4.5
0.2 -10.0
0 - 4
0.4 -6.7
<1.5
4.4
4.4 -13.3
4.4
3.5 -5.0
Remarks
higher mortality at
lower range
emergence delayed beyond
that of w1ld fry incubated at
3.9°C
probe 20.3 em deep
during early development
stages. leads to higher
mortality
lowest limit for good
survival
suggested criteria for
species
lowest temp. prior to closure
of blastopore (Pacific
salmon), then can go as low as
0°C and still have good
survival
range, fertilization to
emergence
Location
Chena River
Clear hatchery
Noatak River
Olsen Creek
Jelta Rfver
Big Beef Creek.
Washington
Big Beef Creek,
Washington
Bira River, USSR
Reference
kogl (1965)
Raymu11d ( 1981)
Merritt and Raymond
{in prep.)
Ba 11 ey ( 1964)
Francisco (1977)
Schroder et al ••
(1974, cited by
Koski. 1975)
Schroder (1973)
Bell ( 1973}
McNeil and Bailey
(1975)
Disler (1951)
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Table III Cont'd. CHUM SALMON
Parameter
Tell1).
Cont•d
Substrate
Partic.le
Size
Embryos and Alevfns -Intragravel Development
Observed Values
3.9 -4.9
Z.4 -3.0
7.0-11.0
4.0
fines <0.0833 em (>12.7%)
silt and sand (61},
<!Scm (81%).> 15 em (131)
Remarks
egg stage
alevtn stage
lcwest temp. in redd
dut tng wfnter
poor quality substrate
redds
Location
Bolshaia River,
USSR
Memu Rfver, Japan
Northern Japan
Prince William
Sound
Reference
Semko (1954, cited
by Sano, 1966)
Sano and Nagasawa
( 1958)
Sano (1966)
Thorsteinson (1965)
tributaries of Burner (1951)
lower Columbia River
5.1 -lO.Z em
1.0 -3.8 em
greater survival to emergence Robertson Creek,
in larger gravel British Columbia
Dill and Northcote
(1970)
sand, 0.0105-0.33Z7 em
(ca. 5 -502:)
mostly O.Z-1.0 em,
some partfc1es>Z.O em
lower survival to emergence
and smaller fry at higher
percentages
redds -also, redds had le~s
of the 0.025-0.1 em
fractions, but more sand
and mud than spawning gravel
11 Sand" (1 .8 -30.01, avg. redds
10.0 -13.51)
11 grave1" (13.4 -60.01. avg.
33.5 -40.31)
Btg Beef Creek,
Washington
Sakhalin
Sakhalin
Koski (1975)
Rukhlov (1969a)
Rukhlov (1969b)
-------------------
Table III -Cont'd. CHUM SALMON
Embryos and Alevins -Intragravel Development
Parameter 06served Va1ues ~emarks [ocat1on ~ef!erence
Substrate wshingle" (13.7 -75.9l,
Cont 1 d avg. 39.2 -53.0l)
> 10 em (0.0 -50 .0l, avg .
0 .7 -9.5l)
Dissolved >2 good survival of eggs and Chena River Kogl (1965)
Oxygen, alevins (strong flow of
mg/1 groundwater)
0 .6 -3.0 low survival
2.8 -6.5 high survival
2.1-4.1 smaller alevins produced at
lower end of range
I ..... 0.0-12.7 annual range Twelvemile Creek McNeil (1962 0
I 5.4 -8.9 September and November Indian Creek
l. 77 -6.80 July -September, lowest Traitors River Mattson et al.
caused by long dry period (1964)
3.6 -8.3 upper intertidal area July -01 sen Creek Bailey (1964)
Septen.~er, low percent
survival at low end of range
<3.0 survival to emergence de-Big Beef Creek, Koski (1975)
creases, ~:!mergence is WasMngton
delayed
3.6-7.6 same rate of egg development, Washington Cheyne (1941, c ited
but eggs held at the highest by Nicola et al.,
value produced the largest fry 1966)
0 . 72 -3 . 70 critical values for embryos Britis.h Columbia Wickett (1954)
0-85 days old
2 lethal limit for hatching
eggs and alevi~s at 5°C
Table Ill -Cont'd. CHUM SALr«lN
Embryos and Alevins -Intragravel Development
Parameter Observed Values Remarks Location Reference
Dissolved 0.72-7.19 critical values for eggs British Columbia Alderdice et al.
Oxygen, with temp . un i ts (0 f) ( 1958)
mg/1 from 4 . 0 -452 • 4
2 lethal limit fnr eggs Briti sh Columbia Wickett ( 1957)
5 low
8 desirable
>2 good survival of eggs and Amur River, USSR Levanidov (1954)
alevins as long as there is
a strong ground water outflow
• ~ 6 -8 most favorable level for N. Okhotsk, USSR lukfna (1973) ......
I entire development period
of embryos and larvae at
4-8°r
3 -4 lowest value 1n redds 81ra River, USSR Dis 1 e r (1951)
--
__________ .. _______ _
Table III -Cont*d. CHUM SALP«lN
Embryos and Alevins -Intragravel Development
Parameter Observed Values Remarks Location Reference
Water 7.6-43.2 depth of redds, range tributaries of Burner (1951)
depth, em 21.6 average depth of redds lower Columbia
River
16 -104 sunmer chum Beshenaya River Smfrnov {1947}
3 -34 fall chum
Stream 0 - 7
Chena River Kogl (1965)
Velocity 1
em/sec
I ..... Intragrave 1 10 minimum for survival British Columbia Wickett {1957) r\)
I apparent
velocity 1 50 desirable level
cm/hr
200 minimum recommended for McNeil and Bailey
Pacific salmon hatcheries {1975)
ca. 25 -150 percent survival to emergence Reiser and Bjornn
increases as velocity increases {1979)
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I II -Cont • d.
Parameter
Sa lfni ty
(ppt)
Observed values
0 -25
0 -11
15 -30
<6
6 -8.5
> 11.6
9 -35
0 -35
<9
CHUM SALMON
Embryos and Alevins -Intragravel Development
Remarks
eggs a"d alevins exposed to
lower levels had higher
survival (te.p. and oxygen
1 evel also factors)
eggs can tolerate continuous
exposure
eggs can tolerate
intenmittent exposure
no effect on early egg
stages
50% of fully developed
can survive through
hatching
retarded egg development
rate of hatching for eyed
period eggs declined from
lOOX at 9 ppt to 25 '' at 35
ppt; alevins hatched at highe r
salinities did not survive
delay in hatching and
decrea s ing survival as
salinity increased
good survival of alevins
location
Prince Wi 11 iam
Sound
Washington
(laboratory)
Washington
(laboratory)
Japan
(laboratory)
Reference
Thorsteinson (1965)
McNeil and Bailey
(1975)
Rockwell (1956)
Kashiwagi and
Sato (1969)
,_
..,
- -- --- -------------
Table III Cont'd CHUM SALMON
Embryos and Alevins -Intragravel Development
Parameter Observed Values Remarks Location Reference
ph 6.5 intragravel Chena River kogl ( 1965)
"slightly acidic 11 i nt ragrave 1 Amur River, USSR levanidov (1 g54)
6.3 -6.5 intragravel Hokka1do. Japan Kobayashi (1968)
co 2 • mg/1 > 10 percent survival to British Columbia Alderdice and
hatching decreases ( 1 abora tory) Wickett (1958)
48 high value in some sections Big Beef Creek. Koski (1975}
of intragravel watPr Washiflgton
I 25 -30 range for ground water Amur River. USSR lev ani dov (1954)
......
~
I > 20 inhibitory to development
rate of eggs
16 -20 tntragravel '-later Bira River. USSR Di~1er {1951)
17.8 -25.2 intragravel water Hokkaido. Japan Kobayashi (1968)
I ...,
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Table IV. CHUM SALMON
Fry -Emergence and Downstream Migration
Parameter Observed Values Remarks Location Reference ~~~~-------~~~~~~~------~~~~------------------------~~~~~-----------·------·-·
Temperature. 3.0 -5.5 oc
5 - 7
5.3-7.2
12 -14
> 15
-0.1, 23 .8
12.9-18.5
21.3
6.7-13.3
8 -10
<15
4.5 -5.5
emergence and outmfgration
range during peak of fry
outmtgratfon
greater rate of emergence
at higher end of range
preferred range,
three months after hatching
generally avoid
ultimate lower and ~pper
lethal temperature
good survival; growth rate
similar over this range but
gross food conversion
efficiency decrease~ as
temperature increases
~urvfval is not good
preferred range for species
preferred temp (over the
range 5.2 -19.0°C)
survive
outm1gration
Delta River
Salcha River
Port John,
British Columbia
British Columbia
( 1 a bora tory)
British Columbia
( 1 a bora tory)
Oregon
(laboratory)
Amur River, USSR
Bolshafa River,
USSR
Raymond (1981)
Trasky {1974)
Barns ( 1969)
Brett (1952)
Brett & Alderdice
(1958)
Kepshire (1976)
Be 11 ( 1973)
Levanidov (1954~
Semko (1954, cited
by Sa no, 1966}
-------------------
Table IV-Cont•d. CHUM SALP«lt~
Fry -Emergence and Downstream Migration
Parameter Observed Values Remarks locations Reference
Temp. 2.0 -3.0 fry become active when temp. Hokka1do, Japan Sana (1966)
Cont•d reaches this level
6.0-10.0 peak of downstream migration
Sa ltn1ty t 35 tolerated by fry that have McNeil and
ppt absorbed yolk sac Bailey (1975)
7 .o, 17.1 tolerated by fry at least USSR levanidov (195Z)
up to 30 hours (laboratory)
I ...... 35.0 8 out of 10 fry died
01
I 35 tolerated by fry 90 days Japan Kash1wagi and
from hatching and older (laboratory) Sato (1960)
IV. DEFICIENCIES IN DATA BASE AND RECOff1ENOATIONS
Chum salmon are the least studied of the five species of Pacific
salmon found in North America (Merrell, 1970). There are only a few
studies conducted in the laboratory on physiolog ical tolera"ces of
chum salmon to various physical and chemical parameters . Because of
the interaction of the effects of several parameters in natural
environments, it fs important to isolate each parameter in the labora-
tory and expose the fish to a wide range of values for that parameter.
This will provide an understanding of the true response of the fish
and o f the lethal limits. It is also important to examine the
interaction of the effects of two or more parameters in a controlled
situation. Much of the data in this report comes from field
observations . It would be incorrect to refer to these as absolute
reqLd rements of the species because the extremes of the particular
par"arneters may not have occurred at the time the field measurements
were made.
Much of the actual data in this report comes from Japan or the Soviet
Union. Another large block of data originated in Washington and
British Columbia. Of all the published references taken as a source
of data for generation of the curves in Section III, only four or fi~e
reported on work in Alaskan streams. This is the greatest deficiency
in this document which is intended to focus on habitat conditions for
chum salmon in Alaska. As a consequence, it is not possible to
document differences fn habitat preferences among different stocks or
different geogranh1cal areas within the State, although many fishery
biologists suspect that such differences exist. In fact. it was not
possible to conclusively demonstrate differences in habitat preference
between A 1 askan chums and those from Washington or Hokka; do or
Kamchatka.
Infonnation on the timing and abundance of chum salmon during the
upstream migration and, to a lesser extent, during the outmigration of
f~y is availa~le in varying degrees of completeness from throughout
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volume) is graphed ve~us particle size. Several different spavning
gravels plotted in this fashion result in an envelope which describes
the range. One further step is needed to make this method well suited
to habitat evaluation and that is to outline those areas of the
envelope that are optimum habitat and those areas that are less than
optimum. This could be done quantitatively by using some measure of
habitat quality such as the density of spawners or the number of eggs
deposited per square meter or the percent survival to emergence per
square meter. Presumably, the areas outside the envelope describe
those substrates that are unsuitable for chum salmon.
The intragravel development of embr~os and alevins is probably the
period of the life history which has the greatest influence on
production. Variation in one of several different parameters during
this critical stage can dramatically influence the survival rate and
can lead to strong or weak brood years. Yet, this stage of life
history, occurring in winter, is the least studied in Alaska.
In fonna ti on is needed on vi rtua 11 y a 11 the important env i ronmenta 1
parameters during intragravel development. Intragravel temperature,
dissolved oxygen concentration. flow, and substrate composition data
are most needed. More information is needed on the differences in
habitat tolerances and preferences that exist among different stages
of development of embryos and alevins.
There is little information available on use of cover by fry during
outmigration. Although chum fry do not overwinter in freshwater as do
kings, cohos, and sock.eyes, they do exper·ience a high ·ate of
mortality during their seaward migration.
In sunmary, much information is neet1ed on habitat requirements,
to 1 e ranees, .. nd prefe~nces of chum sa 1 man during their entire
freshwater life history 1n Alaska. A research program directed toward
the intragravel period would greatly assist our understanding of the
needs of chum salmon and also our understa.1ding of the annual
variation in their abundanc£.
-79-
the State. This infonnation needs to be related to physical~
chemical, and biological parameters of the environment in order to be
useful for habitat evaluatio~.
No data were found on the swimming ability of adult chums during the
upstream migration other than distance coverad per day. Information
on the maximum velocity possible over time and on their ability to
overcome obstacles would be useful in determining whether certain
streams are accessible to spawning chums.
Measurements of the current velocities and water depths in Alaskan
streams d~ring the upstream migration would ~e useful in establishing
the range for those parameters. Daily stream temperature and dis·
charge data from the time of breakup unti 1 after the upstream runs are
completed, correlated with the number of spawners moving upstream, are
needed to establish the optimum levels of those conditions when the
peak run occurs.
Information is needed from Alaska on all parameters associated with
spawning: water temperature, current velocity, water depth, and
substrate composition.
Probably the greatest deficiency in the data base for the species
throughout its range is information on the substrate used for spawning
and for development of embryos and alevins. Much can be determined
about the habitat quality of a particular reach of stream by the
particle size distribution of the substrate. Much of the literature
reviewed for this report described sub!':rate in semi·quantitative
terms such as "large" or "coarse" gravel. Methods used for describinq
substrates should be standardized in order to make comparisons with
other areas more feasible. Some investigators use dry sieves, others
use wet sieves. Some report on the percent passing each sieve size by
volume, others use the percent passing by weight. A good method to
use to describe spawning gravels would be that used by MacKinnon et
al. (1g61) where the cumulative percent sediment passing by weight (or
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This report and the conceptual suitability index curves which were
presented are based in part on a review of published and unpublished
data which was collected by the original authors for purposes other
than constructing such curves. The curves presented here need to be
field tested. A research program is needed to check the accuracy of
these curves • to further refine and narrow their ranges. and to
determine differences in habitat relationships a~~g different stocks.
~80-
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LITERATURE CITED
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Alaska Department of Fish and Game. 1979b. Annual Management Report.
Yukon area. Co~~rcial Fisheries Division--pp.
Alaska Department of Fish and Game. 1979c. Annual Management Report.
Kuskokwim area. Commercial Fisheries Division--pp.
Alaska Department of Fish and Game. 1980a. Annual Management Report.
'<uskokwim area. Commercial Fisheries Divisior1--pp.
Alaska Department of Fish and Game. 1980b. 1960 Alaska commercial salmon
fisheries review. Unpublished preliminary data. Juneau.
Alderice, D. F., W. P. Wickett, and J. R. Brett. 1958. Some effects of
temporary exposure to low dissolved oxygen levels on Pacific salmon
eggs. J. Fish. Res. Bd. Can. 15(2):229-249.
Alderice, 0. F. and w. P. Wickett. 1958. A note on the response of
developing chum salmon eggs to free carbon dioxide fn solution. J.
Fi~h. Res. Bd. Can. 15(5):797-799.
Atkinson, C. E., J. H. Rose, and T. 0. Duncan. :967. Salmon of the North
Pacific Ocean-Part IV. Spawning populations of North Pacific
salmn. 4. Pacific salmon fn the u.s. Int. N. Pac. Fish. Comm. Bull.
No. 23:43-223.
'
Baggerman, Bertha. 1960. Salinity preference, thyroid activity and the
seaward migrat i on of four species of Pacific salmon (Oncorhynchus).
J. ~ish. Res . Bd. Cdn. 17(3):295-322 .
Bailey, Jack E. 1964. Intertidal spawning of pfnk and chum salmon at
Olsen Bay, Prince William Sound, Alaska. U.S. Fish and Wildl. Serv.,
Bureau of Comm. Fish. Manuscript Report 1964-6, Auke Bay. 23 pp.
Bakkala, Richard G. 1970. Synopsis of biological data on the chum salmon,
Oncorhynchus keta (Walbaum) 1792. FAO Species Synopsis No. 41. U.S .
Fish and Wildl . Serv., Bureau of Comm. Fish. Circular 315. Wash.,
D.C. 89 pp.
Bams , R. A. 1969. Adaotations of sockeye salmon associated with
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Symposium on salmon and trout in streams . Univ. of B.C., Vancouver .
388 pp.
Bea 11, Edward P. 1972. The use of predator-prey tests to assess the
quality of chum salmon Oncorhynchus keta fry. M.S. thesis, Univ. of
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Army Corps of Engineers, North Pacific Division. Portland, Oregon.
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investigations in Kotezbue Sound in 1979. Alaska Dept . of Fish and
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Brett J . R. 1952. Temperature tolerance in yot.mg Pacific salmon, genus
Q1corhynchus. J. Fish. Res. Bd . Can. 9(6):265-323 .
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Brett, J. R. and 0. F. Alderice. 1958. The resistance of cultured young
chum and sockeye salmon to temperatures below zero degrees C. J.
Fish. Res. Bd. Can. 15(5):805-813.
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Collings, M. R. 1974. Generalization of spawning and re~ring discnarges
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Geological Survey. Open-file report. 39 pp.
Dangel, James R. and John F. Jewell. 1975. Southeastern Alaska pink and
chum salmon pre-emergent fry data file, 1963-75. Alaska Dept. of Fish
and Game. Technical Oata Report No. 21. Juneau. 226 pp.
Daykin, P. N. 1965. Application of mass transfer theory to the problem of
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factors on survival, condition, and ~iming of emergence of chum salroon
fry (Oncorhyncnus keta). J. Fish. Res. J~. Can. 27{1):196-201.
Cisler, N. N. 1951. Ecological and morphological characteristics of the
development of the Amur autumn ch··m salmon -Oncorhynchus ::""'~
{Walb.). (Transl. from Russian). Pages. 33-41.!.!!. Pacific Salmon:
Se 1 ected art i c 1 es from Soviet perf od i ca 1 s. Is rae 1 Program for
Scientific Translations. 1961. '34 pp.
Disler, N. N. 1953. Development of autumn chum salmon in the Amur River.
(Transl. from Russian). lsrael Program for Scientific Translations,
Cat. No. 763. Office of Tech. Serv .• U.S. Dept. of Commerce, Wash.,
D.C. 14 pp.
Duker, Gary J. 1977. Nest site selection by chum salmon (Oncorhynchus
keta) in a s pawning channel. M.S . the s is, Univ . of Washington,
Seattle. 113 pp .
Emadi, H. 1973. Yolk-sa c malformation in Pac i fic salmon in relation to
substrate, temperature, and water velocity. J . Fish. Res . Bd . Can .
30(8):1 249-12 50.
Fran c isco , Kim . 1976 . Fi rst i nterim report of the Commer ial
Fish-Technical Evaluation Study. Joint State/Federa l Fish and
Wildlife Advi s ory Team. Special Report No . 4. Anchorage, Alaska .
85 pp .
Franc i sco , Kim. 1977. Second interim report of the Commercial
Fish-Technical Evaluation Study. Joint State/Federal Fish and
Wildlife Advisory Team . Special Report No . 9. Anchorage, Alaska .
46 pp .
Gallagher, Arthu r J ., Jr. 1979. An analysis of fac t or s affecting brood
year returns in the wild stocks of Puget Sound chum (Oncorhynchus
keta) and pink salmon (Oncorhynchus gorbuscha). M.S. thesis, Univ . of
Wash., Seattle. 152 pp.
Ha rt , J.l. 1973 . Pac i fic fishes of Canada. Fi sheries Research Board of
Canada . Bull . 180 . 740 pp.
Helle, John H. 1979. Influence of marine environment on age and size at
maturity, growth, and abundance of chum salmon, Oncorhynchus keta
(Walbaum), from CJisen Creek, Prince William Sound, Alaska. Ph.D.
dissertation, Oregon State University. 118 pp.
Hoar, WilliamS. 1956. The behavior of migrating pink and chum salmon
fry. J . Fish. Res . Bd. Can . 13(3):309-325 .
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Hoar, W. S. 1957. Reactions of juvenile salmon to light. J. Fish. Res.
Bd. t::an. 14(6):815-830.
Hoar, Willilm S. 1976. Smolt transformation: evolution, behavior, and
physiology. J. Fish. Res. Bd. Can. 33:1234-1252.
Houston, Arthur A.
(Oncorhynchus
Can. J. Zool.
1q59, Locomotor performance of chum salmon fry
keta) during osmoregulatory adaptation to sea water.
37:591-605.
Houston, Arthur H. 1961. Influence of size upon the adaptation of
steelhead trout (Salmo gairdniri) and chum salmon (Oncorhynchus keta)
to sea water. J. Fish. Res. Bd. Can. 18(3):401-415.
Hunter, J. G. 1959. Survival and production of pink and chum salmon in a
coastal stream. J. Fish. Res. Bd. Can. 16{6):835-886.
Johnson, R. A. 1980. Oxygen transport in salmon spawning gravels. Can.
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Kashiwagi, Masaaki, and Ryuhei Sato. 1969. Studies on the osmoregulation
of the chum salmon, Oncorhynchus keta (Walbaum). I. The tolerance of
eyed period eggs, alevins and fry of the chum salmon to sea water.
Tohoku Journal of Agricultural Research. 20(1):41-47.
Kashiwagi, Masaaki, and Ryuhei Sato. 1970. Studies on the osmoregulation
of the chum salmon, Oncorhynchus keta (Walbaum). I. The tolerance of
adult fish reared in a salt water pond to fresh water. Tohoku Journal
of Agricultural Research. 21(1):21-31.
Keenleyside, Miles, H. A. and William S. Hoar. 1955. Effects of
temperature on the responses of young sa 1 mon to water currents.
Behavior 7(2/3):77-87.
K~pshire, Bernard M., Jr. 1976 . Bioenergetics and survival of chum
(Oncorhynchus keta) and pink {Oncorhynchus gorbuscha) salmon in heated
sea water. Ph.D. 01ssertation, Oregon State University, Corvallis.
112 pp.
Kirkwood, J . B. 1962. Inshore-marine and freshwater life history phases
of the pink sa 1 mon and the chum sa 1 mon f n Pri nee Willi am Sound,
Alaska . Ph.D. Dissertation, Univ. of Loiusville. 300 pp .
Kobaysahi, Tetsuo . 1968. Some observations on the natural spawning ground
of chum and pink salmon in Hokkaido. {In Japanese, English summary).
Sci. Rept. Hokkaido Salmon Hatchery. 22 :7-13.
Kogl, Dennis Raymond. 1965. Springs and ground-water as factors affecting
survival of chum salmon spawn in a sub-arctic stream. M.S. thesis,
Univ. of Alaska, Fairbanks . 59 pp.
Koski, K. Victor. 1g7S. The survival and fitness of t wo stocks of chum
salmon {Oncorhynchus keta) from egg deposition to emergen ce in a
controlled-stream environment at Big Beef Creek. Ph.D. dissertation,
Univ. of Washingtcn, Seattle. 212 pp.
Kusnetzov, I. I. 1928. Some observations on spawning of the Amur and
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Scientif1c Fishery Research Station, Vladivostok . 2(3):1 -124 . Fish.
Res. Bd . Can. Transl . Ser., No. 22.
Lebida, r obert C. 1972 . Yukon Rive r anadromous fish investigations, July
1971 to June 1972 . Alaska Dept . of Fish and Game, Division of
Commercial Fisheri es . AYK Fishery Bull. No . 14 . Anchorage, Alaska.
Levanidov, V, Y. 1952. The osmoregulatory ability of young fall chum
salmon migra ti ng downstream . (T ransl. from Russian}. Pages .245-247
in Pacific Salmon: Selected articles from Soviet periodicals. Israel
Program for Scientific Translations . 1961 . 284 ~p.
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levanidov, V. Y. 1954. Ways of increasing tf.e reproduction of Amur chum
salroon. (Transl. from Russian). Akademiya. Nauk SSSR,
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Levanidov, V. Y. and I. M. Levanidova. 1951. The food of young Amur chum
salmon in fresh water. (Transl. fr.Jm Russian). Izv. Tikh, Nauch. -
Issled. Inst. Ryb. Khoz. Okeanog. 35:41-46. Fish. Res. Bd. Can.
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Lister, D. B. and C. E. Walker. 1966. The effect of flow control on
freshwater survival of chum, coho and chinook salmon in the Big
Qualicum River. The Canadian Fish Culturist. 37:3-25.
Lukina, 0. V. 1973. Respiratory rate of the North Okhotsk chum salroon
[Oncorhynchus keta (Walb.)]. Journal Ichthyology 13(3):425-430.
MacKinnon, D., l. Edgeworth and R. E. Mclaren. 1961. An assessment of
Jones Creek spawning channel, 1954-1961. The Canadian Fish Culturist.
30:3-14.
Ma;Kinnon, Dixon and WilliamS. Hoar. 1953. Responses of coho and chum
salmon fry to current. J. Fish. Res. Bd. Can. 10(B):523-538.
Macy, Paul T. 1954. A preliminary review of the factors influencing
freshwater survival and distribution of Pacific salmon (genus
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.
Mason, J. C. 1974. Behavioral ecology of chum salmon fry (Oncorhynchus
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Mattson, Chester R. and Richard A. Hobart. 1962. Chum salmon studies in
southeastern Alaska, 1961. U.S. Fish and Wildl. Serv., Bureau Comm.
Fish. Manuscript Report 62-5, Auke Bay. 32 pp.
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Mattson, Chester R. and Richard G. Rowland. 1963. Chum salmon studies at
Traitors Cove Field Station, June 1960 to March 1963. U.S. Fish and
Wildl. Serv., Bur. of Comm. Fish. Manuscript Report 1963-11. Auke
Bay. 32 pp.
Mattson, C~ester R., Richard G. Rowland and Richard A. Hobart. 1964. Chum
salmon studies: is southeastern Alaska, 1963. U.S. Fish and WildT .
Serv., Bureau of Comm. Fish. Manuscript Report 1964 -8. Auke Bay.
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Mayama, Hiroshi, and Toshimasa Takahashi. (1975?). Ecological
observations on the adult salmon. I. Diurnal variation of upstream
migration of the adult chum salmon in the Chitose River. (in
Japanese, English abstract). Pages 21-
McNeil, William J. 1962 . Variations in the dissolved oxygen content of
intergravel water in four spawning streams of southeastern Alaska.
U.S . Fish and Wildl. Serv., Special Scientific Report -Fisheries No.
402. Washington, D.C. 15 pp.
McNeil, William J. 1964. Environmental factors affecting survival of
young salmon in spawning beds and their possible relation to logging.
U.S. Fish and Wildl. Serv., Bur. Conm. Fish. Manuscript Report 64-1,
Auke Bay. 25 pp.
McNeil, William J. 1966. Effect of the spawning bed environment on
reproduction of pink and chum salmon. Fishery Bulletin 65(2):495-523.
McNeil, William J. 1~69. Survival of pink and chum salmon eggs and
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and Trout in Streams. Univ . of British Columbia, Vancouver. 388 pp .
McNeil, William J. and Jack E. Bailey. 1975. Salmon rancher's manual.
Ntl. Mar. Fish. Serv., Northwest Fisheries Center, Auke Bay Fisheries
laboratory. 95 pp.
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Herrell, Theodore R., Jr. 1970. Alaska's fishe~ resources. The chum
salmon. U.S. Fish and Wildl. Serv. Bur. Comm. Fish. Fishe~ leaflet
632. Washington, 0.~. 7 pp.
Merritt, P. and J. A. Raymond (in prep.). Early life history of chum
salmon in the Noat~k River and Kotzebue Sound. Manuscript. Alaska
Dept. of Fish and Game, FRED Division, June~u
Morrow, James E. 1980. The freshwater fishes of Alaska. Alaska Northwest
Publishing Co .• Anchorage. 248 pp.
Heave, Ferris. 1g53, Principles affecting the size of pink and chum
salmon populations in British Columbfa. J. Fish. Res. Bd. Can.
9(9) :450-491.
Neave, Ferris. 1955. Note~ on the seaward migration of pink and chum
salmon fry. J. Fish. Res. Bd. Can. 12(3}:369-374.
Heave, F~rris. 1966. Salmon of the North Pacific Ocean -Part III. A
review of the life history of North Pacific sdlmon. 6. Chum salmon
in British Columbia. International N~rth Pacific Fisheries Commission
Bull. No. 18. pp 81-85. Vancouver, B.C.
Nicola, Stephen J., Robert G. Mausolf and Donald E. Bevan. 1966. An
annotated bibliography on the ecology of salmon spawning gravels.
U.S. Fish and Wildlife Service. Special Scientific Report. Fisheries
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Raymond, J. A. 1981. Incubation of fall chum salmon Oncorhynchus keta
(Walbaum) at Clear Air Force Station, A1a~ka. Alaska Dept. of Fish
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I
Reiser, 0. W. and T. C. Bjomn . 1979. Influence of forest and range·land
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Rockwell, Julius, Jr. 1956 . Some effects of sea water a.nd temperature on
the embryos of t he Pac fie salmon, Oncorhynchus gorbuscha (Walbaum)
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Rays, Robert S. 1968. Forecast of 1968 pink and chum salmon runs in
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Rukhlov, F. N. 1969a. The natural reproduction of the autumn chum salmon
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Rukhlov, F. N. 1969b. Materials characterizing the texture of bottom
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Sano, S. 1966 . Salmon of the North Pacific Ocean -Part III . A review of
the life hi.story of North Pacific salmon. 3. Chum salmon in thP Far
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Sana , S. 1967 . Salmon of the North Pacific Ocean-Part IV. Spawning
popu~ations of North Pacific Salmon. 3. Chum salmon in the Far East.
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Sana, Seizo and Ariaki Nagasawa. 1958. Natural propagation of chum salmon
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Schroder, Steven L. 1973. Effects of density of the spawning success of
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:ii1elbourn, J. E. 1966. Influence of temperature, salinity, and
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Shepard. Michael Perry. 1948. Responses of young chum salmon,
Oncorhynchus keta (Walbaum), to changes in sea water content of the
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Sheridan, William L. 1962. Waterflow through a salmon spawning riffle in
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Smirnov, A. G. 1947. Condition of stocks of the Amur salmon and causes of
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Smith, Allan K. 1973. Development and application of spawning velocity
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Smith, David W. 1978. To le,·ance of juvenile chum salmon (Oncorhynchus
keta) to suspended sediments. M.S. thesis, Univ. of Wash., Seattle.
124 pp.
Soin, S. G. 1954. Pattern of development of summer chum, masu, and pink
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Strekalova. I. I. 1963. Observations on spawning of pink salmon
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Tautz, A. F. and C. Groot . 1975 . Spawning behavior of chum salmon
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Thorsteinson, Fredrik V. 1965a. Effects of the Alaska earthquake on pink
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Thorsteinson, Fredrik V. 1965b . Some aspects of pink and chum salmon
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Anchorage, Alaska.
Vaux, Wal t er G. 1962 . Interchange of stream and intergravel water in a
salmon spawning riffle. U.S. Fish and Wildl. Serv., Special
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Volovik, S. P. 1968. Oxygen consumption and food rations of young salmon
in the rivers of Sakhalin. (Transl. from Russian). Izv. Tikh. Nauch.
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Weisbart, Melvin. 1968. Osmotic and ionic regulation in embryos, alevins,
and fry of the five species of Pacific salmon. Can. Journal Zool.
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Wickett, W. Percy. 1954. The oxygen supply to salmon eggs in spawning
beds. J. Fish. Res. Bd. Can. 11(6):933-953.
Wickett, W. P. 1957. The development of measurement and water quality
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95-99 in A. W. Johnson (ed. ). Proceedings Eighth Alaskan Science
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Wickett, W. P. 1958. Review of certain environmental factors affecting
the production of pink and chum salmon. J. Fish. Res. Bd. Can.
15(5): 1103-1126.
Wilson, William J., F.. Woody Trihey, Jean E. Baldrig~. Charles D. Evans,
James G. Thiele and David E. Trud9eon. 1981. An assessment of
environmental effects of construction and operation of the proposed
Terror Lake hydroelectric facility, Kodiak. Alaska. Instream flow
studies. Final report prepared by Arctic Environmental lnfonnation
and Data Center, Univ. of Alaska, Anchorage. 419 pp.
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FRESHWATER HABITAT
RELATIONSHIPS
PINK SALMON-ONCORHYNCHUS GORBUSCHA
I AlASKA DEPARTMENT OF FISH" GAME
HABITAT PROTECTION SECTION I RESOURCE ASSESSMENT BRANCH
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APRil, 1981
FRESHWATER HABITAT RELATIONSHIPS
PINK SALMON (ONCORHYNCHUS GORBUSCHA}
By
Steven W. Krueger
Alaska Department of Fish and Game
Habitat Division
~esource ~ssessment Branch
570 West 53rd Street
Anchorage. Alaska 99502
Hay 1981
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Acknowledgements
Fisheries biologists from the Alaska Department of Fish and Game, the U.S.
Forest Service, U.S. Fish and Wildlife Service, National Marine Fisheries
Servi ce-Auke Bay Laboratory, Bureau of Land Management, and the Arctic
Environmental Information and Data Center provided insight into this
project and deserve recognition. The assistance of librarians at the
Alaska Resource Library, the u.s. Fish and Wildlife Service
Library-Anchorage and the Commercial Fisheries Library-Alaska Department of
Fish and Game-Juneau is heartily appreciated.
This project was funded by the U.S. Fish and Wildlife Service, Western
Energy and Land Use Team, Habitat Evaluation Procedure Group, Fort Collins,
Colorado. Contract No. 14-16-0009-79-119.
TABLE OF CONTENTS
I. INTRODUCTION
A. Purpose
B. Distribution
C. Life Hi story Su111111ry
D. Economic Importance
II. SPECIFIC HABITAT REQUIREMENTS
A. Upstream Migration
1. Water temperature
2. Stream discharge, current velocity
and t~~ater depth
8. Spawning
1. Water temperature
2. Current velocity
3. Water depth
4. Substrate composition
5. Spa~er Densities
C. Intragravel Development of Eggs and Alevins
1. Water temperature
2. Stream discharge. current velocity
and water depth
3. Substrate composition
4. Dissolved oxygen
5. Sa 1 i ni ty
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D. Downstream Fry Migration
1. Cover
2 . Discharge
3. Water temperature
III. CONCEPTUAL SUITABILITY INDEX CURVES
IV. DEFICIENCY IN DATA BASE
V. RECOMMENDATIONS AND FURTHER STUDIES
VI. LITERATURE CITED
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I. INTRODUCTION
A. Puroose
The purpose of this project is to describe how selected physical
and chemical features of lotic habitats within Alaska influence
the survival and behavior of the freshwater life stages of pink
sa 1 mon, Oncorhynchus gorbuscha ( Wa 1 baum).
Objective,s of this project are:
1) To gather data from published and unpublished sources within
Alaska, Canada, U.S.S.R., the Pacific Northwest, and Japan (with
emphasis on Alaska), and from fishery biologists from Alaska and
nearby areas concerning aquatic habitat -pink salmon
relationships;
2) To develop an Alaskan data base composed of narrative and
habitat suitability index (HSI) models following U.S. Fish and
Wildlife Service techniques to better understand lotic
habitat-pink salmon relationships; and,
3) To identify data gaps and reconmend appropriate research to
fill these gaps.
This report discusses the habitat relationships (with emphasis on
the physical and chemical habitat components) of the fresh water
life history stages of the pink salmon which include:
upstream spawning migration;
spawl'li ng;
egg, alevin development; and,
downstream fry migration.
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B. Distribution
Pink salmon are the most abundant of the five species of Pacific
salmon. They spawn in North American and Asian streams bordering
the Pacific and Arctic Oceans and are more abundant in Asia than
in North America (Scott and Crossman, 1973). In North America
this fish ranges from northern California to northern Alaska and
eastward to the Mackenzie River, Northwest Territories. Within
Asia, the pink salmon is found from K~rea and Hokkaido, Japan
northward and westward to the Lena River, Siberia. Pink salmon
have been introduced to and are maintaining populations in Lake
Superior, Newfoundland, and northern Europe.
Pink salmon are widely distributed along coastal Alaska; major
production areas include streams within the southern Alaska
Peninsula area eastward to southeastern Alaska including the
Chignik area, Kodiak Island, Cook Inlet, and Prince William Sound
(Alaska Department of Fish and Game, 1978). More localized pink
salmon production occurs in the lower Kuskokwim and Yukon River
drainages and certain Norton and Kotzebue Sound streams. Pink
salmon are found in the Yukon River drainage up to the Anvik
River. Pink salmon abundance in streams above the Arctic Circle
is relatively 1 ow, but they have been documented in Hmited
numbers as far east as the mouth of the Mackenzie River.
C. L He Hi story Sunmary
Pink salmon have the shortest life cycle of any Pacific salmon.
They spend about 15 months in the sea and grow to about 1 to 3 kg
(2.2 to 6.6 lbs), then migrate to coastal areas and usually
ascend their natal streams to spawn and die. Explanations of
timing and orientation of ocean migration by pink salmon are not
thoroughly understood. Olfactory cues probably aid pink salmon
in detecting their natal streams after the fhh reach coastal
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waters. However, pink salmon tend to stray more than the other
species of Pacific salmon (Hasler, 1971; Morrow, 1980).
Pink s.almon may enter Alaskan streams from June through October,
but arrival and subsequent s.pawning times in a given stream are
similar from year to year (Sheridan, 1962 b). Early, middle and
late runs of pink-salmon occur in southeast Alaska and Prince
William Sound {Alaska Department of Fish and Game, 1978). The
two year 1 ffe eye 1 e prevents consecutive., • eve.n 1 and 1 odd' year
stocks from interbreeding and produces genetically distinct
1 even 1 and 'odd' year runs. The abundance of these runs often
varies considerably.
Distance of from the estuary to spawning areas may influence the
timing of upstream migration of pink salmon in large river
systems. Pink salmon spawning in the upstream reaches of the
Skeena River, northern British Colt.dnbia, traditionally arrive
about two weeks earlier than fish that spawn in the lower reaches
of the Skeena River (Neave, 1966).
The extent of upstream migration of pink salmon within streams is
variable. They ge.nerally spawn within 40 miles of the ocean, but
may ascend streams for considerable distances. Pink salmon have
ascended the Yukon River as far as 160 km (100 mi), the Fraser
and Skeena rivers up to 500 km (311 mi) and the Amur River, China
up to 700 km {435 mi) (Scott and Crossman, 1973).
The relative abundance of fish can influence pink salmon
distrih·1tion and the extent of upstream migration. During years
of high abundance of pink salmon in various streams ('on' years),
reaches of a stream or adjacent streams which nonnally support
few, if any of these fish, may support substantial numbers of
pink salmon (Neave, 1966).
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FIG.1 . DISTRIBUTION OF PINK SALMON IN ALASKA
(ALASKA DEPT. OF FI ,SH & GAME, 1978)
-4-
The spawning behavior of pink salmon is similar to other
salmonids. The female is solely responsible for construction of
the redd and the male defends the immediate redd area from other
male pink sal~~n. The female excavates the redd by turning on
her side and repeatedlt flipping her caudal fin against the
substrate, displacing silt, sand and gravels which are carried
downstream. Some of the displaced substrate material fonns a
characteristic lip on the downstream end of the redd. Dimensions
of the finished redd vary with the size cf tnt:! fema 1 e, current
velocity, and substrate composition and imbeddedness. Redds can
be as long as g1.5 em {36 in) and up to 45.7 em (18 in) deep.
The spawning pair then descend into the center of the redd, with
mouths agape and fins erect and release eggs and sperm. The
fertilized eggs, 0.6 em (0.24 in) in diameter, settle into
interstices within the redd. Egg numbers may range from 800 to
more than 2,000. The fecundity of pink salmon is variable and
influenced by the size of the fish, the geographic area and the
year (Scott and Crossman, 1g73). After the pair completes
spawning in the first redd, the female may begin construction of
a ntw redd immediately upstream of the previous one. This
activity displaces gravels into the first redd, cove .. ing the
fertilized eggs. McNeil (1g62) determined that pink salmon eggs
within selected redds of several southeast Alaska streams were
overlain by 0.08 to 0.41 m (0.25 to 1.33 ft) of gravel. This
gravel layer protects the fertilized eggs from sunlight and
predation, reduces mechanical disturbance by anchor ice and other
objects, allows water to transport tixvgen to and metabolir wastes
from the deve 1 oping eggs. The fema 1 e guards the redd from
disturbance by other females. A female may spawn with different
males and both sexes die soon after spawning.
Streams used by pink salmon may range f~om extremely small, short
coastal streams to large river systems like the Yukon River.
Tidally and non-tidally influenced stream reaches are used for
spawning. Intertidal spawning is especially characteristic of
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pink salmon populations within Prince William Sound and southeast
Alas~a. (He11e, Williamson and Bailey, 1964).
Hatching of fertilized pink salmon eggs occurs in the spring.
Hatching is followed by a period of alevin growth as the attached
yolk sac gradually shrinks. Eggs and alevins remain in the redd
gravels from 6 to 8 months.
Substarttial mortality of pink salmon eggs and alevins (sometimes
exceeding 75S, occasionally 90S} has been documented in selected
reaches of various streams in Alaska and British Columbia
(Wickett, 1952~ Hunter, 1959• McNeil, 1962; Helle, Williamson and
Bailey, 1964}. McNeil {1962} detennined greater mortality
generally occurred during the egg than the a levin stage of
development. Egg and alevin mortality appears to be the major
1imitation to production (McNeil, 1962; Bailey, 1969}.
The fry emerge from the gravels in the spring (March to June}
when development 1s complete. The newly emerged fry range from 3
to 4.5 em (1.2 to 1.8 in} in length (Scott and Crossman, 1973}.
They immediately begin migrating downstream to the sea (Neave,
1966}. Emergence and subsequent downstream movement of fry
generally occurs shortly after dusk and ceases before dawn (Hoar,
1956; Godin, 1980}. Pink salmon migrating down short coastal
streams may reach the estuary within the first night of travel.
Pink salmon fry migrating long distances downstream may cease
mrlgration during the day and conceal themselves in the substrate.
They res~ downstream movement at dusk. L im1ting migration to
dusk or dark probably minimizes predation (Heave, 1966}. Pink
salmon fry may migrate downstream during the day at flood flows,
periods of high turbidity, or during the latter portion of fry
emergence {Neave, 1966; MacDonald, 1960; Godin, 1980}.
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0. Ecological and Economic Importance
Pink salmon are extremely important to the conmercial fishery,
and to a lesser extent, to the recreational and subsistence
fisheries. Although their relative value (price/pound) to
commercial fishermen is lower than for most other Pacific salmon,
their abundance makes them economically important. FI"'R'' 1962
through 1976, pink salmon contributed 301 of the income brought
to commercial fishermen from all Pacific salmon caught in Alaskan
waters. Their mean annual value to commercial fishermen during
this period was nearly 16 million dollars. They are the 'bread
and butter' commercial fish species within the southern Alaska
Peninsula, Chignik area, Kodiak Island, Cook Inlet, Prince
William Sound, and southeast Alaska, especially during years of
abundance (Morrow, 1980~ Alaska Department of Fish and Game,
1979}. Pink salmon are harvested commercially by beach and purse
seines, and drift and set gill nets. Pink salmon are also
harvested by trolHng, primarily in southeast Alaska (Alaska
Department of Fish anci Game, 1979). Pink salmon are usually
canned, the roe is processed for exportation, and the remaining
material used for fertilizer and pet food.
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II. SPECIFIC HABITAT RELATIONSHIPS/REQUIREMENTS
A. Upstream Spawning Mi~ration of Adults
l. Water temper~ture
Sheridan (1962b} "bserved pink salmon returning to and
spawning in various streams at different times of the year.
He determined that pink salmon spawned in colder streaDIS
earlier in the season than in wanner streaiiiS. Sheridan
attributed variations in spawning times to adaptations of
pink salmon to the water temperatures.
Upstream migration of pink salmon has been documented in
streams with water temperatures ranging from 7.2 to 15.5°C
(45 to 60°F) (Bell, 1g73). Various researchers have not
found significant positive correlations between upstream
migration of pink salmon and water temperature (Neave,
1966). Upstream spawning migration may also be delayed by
water temperatures exceeding 21°C (70°F} (Bell, 1g73).
2. Stream discharge, current velocity and water depth
Stream discharge, current velocity, and water depth can
influence the upstream spawning migration of pink salmon 1~
a variety of ways.
High stream discharges with associated high current
velocities can surpass the swimming capabilities of pink
salmon and prevent upstream movement. If current velocities
near constrictions or falls exceed the swimming capabilities
of adult fish, the upstream migration will cease. Pink
salmon can negotiate maximum current velocities of about 2.1
mtsec (6.6 ft/sec). Waterfalls that may block fish passage
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at some flow conditions may not limit fish at other flows
(Thompson, 1972).
Conversely, low flows with characteristic shallow water
depths can also prohibit fish movement upstream, depending
on the severity of dewatering (Neave, 1966; Reher and
Bjornn, 1979). Minimum water depths needed for upstream
passage of a.dul t pink salmon a.re about 0.1B m (0.6 ft)
(Thompson, 1972). These values will vary with the size a.nd
condition of adult pink salmon a.nd the length of stream
reach with shallow water. Pink salmon have been observed
passing over shallow riffles less that 0.09 m (0.3 ft) deep
along the Kizhuya.k and Terror Rivers (Wilson, Trihey,
Baldridge, Evans, Thiele and Trudgen, 19B1).
Davidson a.nd Vaughan {1943) examined the relationship
between adult pink salmon numbers a.nd discharge rates in two
streams in southeast Alaska. and one in British Columbia.
They found a. positive correlation between di scha.rge a.nd
numers of ascending adult pink salmon in Sashin Creek,
British Columbia.. ln this creek the fish reside in the ba.y
r.ear the stream when maturing, then enter the stream,
usually during freshets. No positive correlation was found
between numers of ascending pink salmon and discharge rates
in the other two streams. Differences in these findings may
be attributed to the relatively shallow water of Sa.shin
Creek and the relatively deep pools of the other two streams
(Banks, 196B).
Hunter (1959) showed that flow conditions in Hooknose Creek,
British Columbia, accounted for about 34% of the variation
in upstream fish movement. However, only data from 1952
wer _ presented.
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Pritchard (1948) also found a significant positive
correlation between numbers of adult pink salmon migrating
in this stream and rainfall and discharge in McClinton
Creek, British Columbia.
B. Spawning
1. Water temperature
Sher!dan (1962b) observed pink salmon spawning in southeast
Alaska streams at water temperatures from 7.2 to 18.40°r. (45
to 65°F). Bailey (1971) reported that pink salmon spawning
activity commenced in Grace Creek, near Ketchikan, Alaska in
mid-August at maxin~~m water temperatures of about 10°C
(50°F). McNeil (1964) stated that pink salmon spawning in
Sashin Creek generally occurs after water temperatures
decline to 13°C (55.4°F) or below.
Pink salmon are reported to spawn 1n Hooknose _Creek, British
Coluneia, in water temperatures cooler than 12°C (58°F)
(Hunter, 1959). Webb (1978 a, b and 1980) reported pink
salmon spawning in Buckland, Shaktoolik and lnglutalik
Rivers in water temperatures of 16°C (61°F), 9°C {48°F) and
10°C (50°F), respectively. Bailey and McNeil (1975) stated
"mature salmon generally ripen and spawn as water
temperature declir.es from its s111111er maximum; the preferred
range for spawning is 7.2°C to 12.8°C (45 to 55°F)!• Bell
(1973) also stated that the preferred spawning temperatures
of pink salmon range from 7.2 to 12.8°C (45 to 55°F).
Relatively high stream water temperatures and low dissolved
oxygen levels associated with drought conditions havP.
apparently killed many ripe pink salmon in several southeast
Alaska streams. Water temperatures exceeding l7°C and
dissolved oxygen levels of 4 ppm or less were responsible
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for the mortality of severa1 thousand adult pink salmon in (
Staney Creek, Alaska during mid-August, 1979 (personal
communication, J~hn Edgington , 1981).
2. Current velocity
Asian pink salmon spawn in a variety of current velocities,
ranging from about 0.3 to 0.8 m/sec (1.0 to 2.6 ft/sec).
They ttave also been observed spawning in a.reas of upwelling
water with no current during years of abundance ( Setnko,
1939; Krchkin and Krugius, 1937; Ovinin, 1952).
Current velocities utilized by pi n k salmon in selected State
of Washington streams are reported to be 0.21 to 0 .99 m/sec
(0.7 to 3 .30 ft/sec) (Co11ings, 1974). These velocity
mea.surements were taken 0.12 m (0.4 ft) above the substrate.
Hourston and MacKinnon (1957) described c:urrent velocity and
water depth preferences of spawning pink salmon in the Jones
Creek spa.wning channel on Va.ncouver Island, British
Columbia . The 615 m (2,000 ft) long spawning channel was
divided into 15 sections of equal length and three "complete
depth-velocity c r oss sections were, made ;, each section of
tne channel " at a discharge of about 7 m3/sec (246 cfs)
prior to the arrival of the fish.
Current velocities within the channel apparently exceeded
preferrt:a velocities of s.pawn1ng pink salmon. The first
fish to enter the experimental channels usually chose the
slowest velocities available in the channel . Only 40& pi·nk
salmon entered the channel in 1955; density conditions were
extremely low with only one spawning female per 10 .5 m2 of
channel.
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Pink salmon spawning in Terror River, Alaska selected
current velocities ranging from 0.19 to 0.66 m/sec (0.6 ~o
2.0 ft/sec} and within this range, they preferred velocit~es
from 0.35 to 0.47 m/sec (1.1 to 1.5 ft/sec) {Wilson, Trihey,
Baldridge, Evans, Thiele and Trudgen, 1981). All values are
expressed as mean column velocities.
Pink salmon in the upper Skagit River, Washington spawned in
areas with current velocities ranging from 0.10 to 1.32
m/sec (0.3 to 4.3 ft/sec) (Graybill, BuMJner, Gislason,
Huff~n, Wyman, Gibbons, Kurlso, Stober, Iagnan, Sta~n and
Eggers, 1979). Eighty percent of the fish were observed in
current velocities ranging from 0.37 to 1.0 m/sec (1.2 to
3.2 ft/sec). All measurements were taken 0.16 m (0.5 ft)
above the substrate.
3. Water depth
Selection of water depths by spawning pink salmon is
associated with current velocity, substrate type and spawner
densities. Kuznetsov (1928) stated that pink salmon in the
Amur River and streams of West Kamchatka spawn in depths
ranging from 0.2 m (0.7 ft) in relatively fast current
velocities to 0.3 m (1 ft) in slower currents. Ovinin
(1952) observ~d pink salmon in streams of southern Sakhalin.
He found that in uncrowded conditions the fish spawned in
water depths of 0.5 m (1.7 ft) and in crowded conditions
they utilized water depths from about 0.1 to 1.2 m (0.3 to
3.9 ft).
Water depth preferences of pink salmon in selected
Washington State streams are about 0.15 to 0.53 m {0.5 to
1.75 ft) (Collings, 1974). Hourston et al. determined water
depth preferences of spawning pink salmon in the Jones Creek
spawning channel, using the transect method.
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They observed that the first fish to enter the spawning
areas chose mean water depths of 0.42 ~ {1.38 ft), and the
subsequent 100 spawning fish occupied slightly shallower
mean water depths of 0.39 m (1~28 ft) (Table 1). The mean
water depth of the entire spawning channel was 0.37 m (1.20
ft}. No fish utilized the relatively shallow depths (0.30
to 0.34 m, 1.0 to 1.1 ft) which also had rather high current
velocities (0.73 to 0.98 m/sec, 2.4 to 3.2 ft/sec). Female
pink salmon density within the channel was extremely low
with one female per 10.5 m2•
Spawning pink salmon in the Terror River, Alaska used water
depths ranging from 0.09 to 1. 2 m (0 .3 to 3.8 ft) and
demonstrated a preference for depths from 0.37 to 0.63 m
(1.2 to 2.0 ft) (Wilson et al., 1981).
Spawning pink salmon in the Skagit River, Washington were
observed at water depths ranging from 0.09 to 1.32 m (0.3 to
4.2 ft} and 80% of those fish were in water ranging in depth
from 0.28 to 0.78 m (0.9 to 2.5 ft) (Graybill et al., 1979).
4. Substrate composition
Pink salmon spawn over a variety of substrate materials.
The size, shape, density and imbeddedness of this material,
current velocity, water depth and fish densities can
influence substrate selection. Adult pink salmon usually
spawn in small gravels, sometimes reaching 10 em (4 in} in
diameter. Pink salmon have also been observed spawning over
much 1 arger substrate in selected Prince Wi 11 iam Sound
streams during •on• years (R. Nickerson, personal
communication, 1980).
Hourston and Mac Kinnon (1957) examined substrate samples
taken from •good• pink salmon spawning areas along the Adams
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and Okanagan Rivers and Jones Creek. They found that gravel
from 0.6 to 3.8 em diameter provided the best substrate for
in artificial spawning channels. Lucas (1959) used
substrate sizes from 2 to 10 em diameter in spawning
channels.
Spawner Densities
Redd superimposition influences egg survival when high
densities of pink salmon spawn in streams with limited
spawning habitat. Pink salmon redds may be partially or
totally torn apart and reexcavated by newly arriving
females. McNeil (1962) formulated a hypothetical model
predicting the effects of pink salmon spawner density on egg
survival in a selected reach of the Harris River. At a
spawner density of 4.5 fish/9.3 m2 (100 ft2) redd
superimposition accounts for about 251 of tot11l egg
mortality. At a density of 10.6 fish/9.3 m2 (100 ft 2 )~ egg
mortality was predicted to be about 501. Helle (1966) also
recognized that redd superimposition increased pink salmon
egg mortality in Olsen Creek~ Prince William Sound.
C. 1ntragravel Development of Eggs and Alevins
1. Water temperature
Rates of egg development and hatching times are strongly
controlled by temperature regimes and by the number of
degree-days accumulated since egg depo~ition. A degree-day
is the number of degrees above a base temperature (usually
the freezing point) in the average temperature of one day.
It has been estimated that at least 500 degree days (°C) are
required for the hatching of pink salmon eggs (Scott and
Crossman. 1973). This is equivalent to 50 days at 10°C or
71 days at 7°C. Abnormally warm or cold temperatures can
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accelerate or depress development rates and cause premature
or delayed salmon fry emergence.
There are upper and lower temperature limits for successful
incubation of salmonid eggs. Pink salmon eggs are more
sensitive to cold water {0°C, 32°F) at initial developmental
stages than later stages. Combs and Burrows (1957) and
Combs {1965) reported that hatchery pink salmon eggs could
tolerate water temperatures as low as 0.5°C (33°F} provided
temperatures exceeded 5.5°C (42°F) during the initial month
of development. Bailey and Evans (1971) concluded from
laboratory tests with pink salmon eggs from Grace Creek near
Ketchikan, Alaska that water temperatures above 4.5°C (40°F}
are necessary through the gastrula stage to ensure
development.
Bai 1 ey and Evans (1971) found that egg mort a 11 ty and the
occurrence of spinal deformities in alevins were inversely
related to increases in water temperatures between 2.0°C
(36°F} and 8.5°C (47°F). Water temperatures of 4.5°C (40°F)
and 3.0°C (37°F} caused substantial egg mortalities and
spinal deformities among alevins. Complete mortality of
eggs occurred at 2.0°C (35°F). No egg mortalities or alevin
spinal deformities were found at 8.5°C (47°F). After pink
salmon eggs reach the gastrula stage they can tolerate water
temperatures close to freezing.
The upper and lower estimated threshold temperatures for
pink salmon eggs are 13.3°C and 4.4°C (Bell, 1973).
Mortality is expected to increase if these thresholds are
exceeded. Eggs will survive and develop normally at lower
temperatures than 1nd1cated, provided initial development of
the embryo has progressed to a stage that is tolerant of
co I der water.
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Timing of the downstream migration of pink sal.,n fry is
probably correlated with habitat conditions in the estuary
in late spring or early sumner (Sheridan, 1962). Fry
err,J?rging earlier or later than nonnal could encounter
:uboptlenal physical conditi 'lns and lower food availability
in the estuary.
2. Stream discharge, current velocity and water depth
Discharge alterations can adversely affect developing pink
sal.,n eggs and alevins. Spates can mechanically remove
substrate material and developing fish {McNeil, 1966).
McNeil (1966) observed damaged pink salmon redds and
displaced pink sal.,n eggs and alevins in several
southeastern Alaska streams after intense rains between
October and December. He estimated that these floods
destroyed about 50 to 90S of the developing pink salmon eggs
and alevins within these streams. Pink salmon spawning
areas within the mainstem Terror River have been scoured
during storm flow conditions (Wilson, 1981). This
phenomenon is probably more comnon in .,derately high
gradient streams used by spawning pink salmon.
Summer and winter low flows can adversely affect developing
pink salmon eggs and alevins, depending on the severity of
f 1 ow reduction, temperatures , and other factors. McNeil
(1966) believed that significant .,rtality of chum and pink
salmon eggs and alevins occurred in one of two study streams
in southeastern Alaska during extremely cold weather and low
flows. He noted that discharge in Indian Creek was
considerably less than in Twelvemile Creek. Indian Creek is
characterized by an extreme range of discharges {500 fold};
in contrast, Twelvemile Creek exhibited only an 80 fold
difference.
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Pink salmon eggs deposited along the stream margins may t~
subjected to high mortality with winter low flows. Thi·;
could cause redds to become exposed, desiccated, or froze1.
Summer drought conditions, and the attendant low flows an~
high temperatures, can cause reduced intra-gravel di ssobect
o~ygen levels. McNeil (1966) attributed the high .artalfty
of pink and chum salmon in several southeastern Alaskl .
streams in 1957 to 1 ow flow, unseasonal ty warm s~~~~~er
temperatures and tow dissolved oxygen levels.
3. Substrate composition
Substrate composition, particularly the percentage of s-.11
particles (called 'fines') can influence the intragra"rel
environment and condition of developing eggs and atevi1s.
Fines have been defined as materials less than 6.4 mm (1.26
in), 3.0 mn (0.12 in) and 0.833 mn (0.03 in} mini11111
diameter by McCuddin {1977), Phillips (1975) and McNeil and
Ahnell (1964}, respectively. Increased amounts of fines
within the substrate interstices where eggs and alevins are
developing, can reduce water penr.eabil ity and intragravel
flow rates (apparent Vt!locity) (Wickett, 1962}. This
influences the rate at which oxygen is transported to a~d
metabolic products from the developing eggs and alevins.
McNeil and Ahnell (1964} classified the pink salmon
production capability of six streams in southeastern Ala!:ka
by the permeability of their spawning areas. They conctu~ed
that the substrate of 'productive • streams generally
contained less than 5i by volume of fines (0.833 mn) and had
associated -,ermeabfl fty rates of 24,000 cm/h (787 ft/h).
Less productive pink salmon streams were characterized bJ
151 or more fines w1th associated permeability rates of less
than 1 ,300 cm/h ( 43 ft/h}. Wickett (1958) and Reiser an(l
Bjornn (1979) reported 121 survival of pink and chum salmcn
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to emergence at penneability rates of 96,000 cm/h {3,150
ft/h) and 2i survival at rates of 1,800 cm/h (59 ft/h}.
Pink salmon egg and alevin survhal in relation to tidal
influence (with associated sediment, salinity, dfssol ved
oxygen and temperature conditions) has been examined by
several researchers. Scud and Hanavan (1954) detennfned
that pink salmon egg and alevin su."'Vival in tidally
influenced reaches of Lover's Cove Creek were as high or
higher than in upstream non-tidal influenced reaches.
kirkwood (1962) detennined that egg and alevin survival
decreased from higher to lower tidal levels in several
Prince William Sound streams. Helle, Williamson and Bailey
(1974) examined Olsen Creek in Prince William Sound. They
report'!d decreased survival at lower intertidal reaches
associated with increased proportions of fine sediments.
Helle (1970) corroborated previous findings in Olsen Creek.
He attributed greater egg and alevin mortality rates from
higher to lower intertidal areas to increased fine sediments
and salinity. Dissolved oxygen and temperature were
addft1onal factors influenc1ng mortality.
Fines can influence emergence of alevins from the redd.
Survival and emerge~ce of chinook salmon and steo,head trout
fry were adversely affected when fines (6.4 mm) exceeded 20i
of the volume of the substrate in laboratory troughs
(Bjornn, 1969; McCuddin, 1977). Koski (1966) documented
that coho salmon fry were unable to emerge from natura 1
redds with substantial accumulations of fines.
4. Dissolved Oxygen
Dissolved oxygen is supplied to developing eggs and alevins
within the redd by intragravel flow. Dissolved oxygen
levels within the redd are influenced by the dissolved
-18-
oxygen levels within the stream, rate of intragravel flow,
biological oxygen demand of material within the redd such as
detritus and dea.d eg~s, water temperature, metabolic rates
of the deve 1 oping eggs and a 1 evi ns, density of eggs and
alevins, and other factors.
Oxygen consumption generally increases as embryo development
progress {Bailey, Rice, Pella and Taylor, 1980). Bailey et
al. (1980) planted ~Various densities of fertilized, eyed
pink salmon eggs (O; 1,600; 6,400; 1 2,800 and 2:5,600
eggs/0.015 m3 ) in experime.ntal egg incubators . Dissolve·d
oxygen content of inflow water was maintained at relatively
constant levels during the test: 9.16 mg/1 in December and
8. 08 mg/1 in Apri 1. Oxyge ,n consumption rates measured at
pre-hatching, post-hatching and pre-emergence progressively
increased with time and developmental stage. For example,
within the incubator containing 6,400 eggs, oxygen
cons.umption rates increased from 0 .003 to 0.010 mg/h per egg
pl':"ior to and following hatching and reached a maximum of
0 .027 mg/h per alevin inmediately prior to fry emergence.
Resu 1 tant di s ·so 1 ved oxygen 1 eve 1 s within the various
incubators reflected egg densities and oxygen consumption
rates .
Dissolved oxygen levels generally decreased with time and
were progres.sively lower within incubators containing more
eggs or alevins (Bailey et al., 1980). For example,
dissolved oxygen levels immediately prior to emergence of
fry within incubators containing 1,600, 6,400 and 12,800
alevins ·were 7.0, 5.5 and 4.3 mg/1 respectively. An
exception occurred in the incubator containing 25,600 eggs
where alevins emerged prematurely. Premature emergenc.e was
probably due to the combined effect of low dissolved oxygen
and high a11111onia levels . The dissolved oxygen concentration
at emergence was only 6.2 mg/1.
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Dissolved oxygen levels exceeding 6.0 ~/1 are recommended
for the successful deve 1 opment of pink sa 1 mon eggs and
alevins (Bailey et al., 1980). Dissolved oxygen levels
below 6.0 mg/1 apparently caused premature emergence,
decreased size and low survival of fry, especially at higher
densities. Fifty percent of the fry in incubators
r.:ontaining 12,800 and 25,600 eggs emerged 7 and 82 days
earlier, respectively, than fry in the incubator with 1,600
eggs. Emergent fry at the two highest densities also were
shorter than fry from the other incubators. Survival was
only 501 in the highest density incubator.
Coble (1961) and Phillips and tampbel 1 (1961} determined
that intragravel dissolved oxygen concentrations at or above
8 mg/1 are necessary for high survival of steelhead trout
and coho salmon eggs. McNeil (1962j attributed low survival
of pink salmon eggs in 195 7 in ievera 1 southeast A 1 ask a
streams to low flows and associated le-w dissolved oxygen
levels and, possibly, other factors. Reiser and Bjornn
(1979} recommended dissolved oxygen levels near saturation
with momentary reductions no lower than 5 mg/1
(approximately 40-451 saturation} for eggs and alevins.
5. Salinity
Pi n k sa 1 nmn conmon 1 y spawn in the in terti da 1 reg i ons of
streams. Noerenberg (1963} estimated that over 501 of the
pink salmon spawning activity in Prince William Sound occurs
in tidally influenced areas. Fluctuations in sa11 nity,
dissolved oxygen, water temperature and fine sediment may
influence pink salmon egg and a levin survival. Various
researchers have recognized that pink salmon fry production
within intertidal stream reaches is generally highest in the
upper reaches, moderate in the intermediate reaches and low
or non-existent 1n the lower reaches. Little research has
-20-
focused on the effect of just salinity on pink salmon egg
and alevin survival (Hanovan and Skud, 1954; kirkwood, 1962;
Hell, Williamson and Baily, 1964i Helle, 1970).
Bailey (1966} conducted laboratory experiments to simulate
salinity conditions of developing eggs and alevins at the
2.4 m {8 ft), 1.8 m (6 ft) and 1.2 m (4 ft) tide levels in
Olsen Creek, Alaska. Test salinity conditions ranged from
12-30°/oo. He concluded that developing pink salmon eggs
fertilized in fresh water showed no adverse affects to
salinity conditions at the 2.4 m (8 ft) tide level. Pink
salmon fry production was severely limited at the 1.8 m (6
ft) tide level and was extremely limnted or nonexistent at
the 1.2 m (4 ft) tide level.
D. Downstream Fry Migration
1. Cover
Pink salmon fry may reach the estuary along short coastal
streams within the first night of travel, but fry migrating
down r~latively long strea~ need adequate cover during the
day. Cover could be from gravel interstices, overhanging
riparian vegetation or other instream cover.
2. Discharge
A wide range of discharges are sui tab 1 e for successful
downstream fry migration. Discharges must be high enough to
maintain adequate water depths in the channel and to enhance
the swimmnng of the fry.
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3. Water temperature
Water temperatures encountered during downstream migration
and in the estuaries can influence dissolved oxygen levels,
swimming perfo~nces and growth rates of the fry, and the
ability to capture and digest food (Reiser and Bjornn,
1979). Brett (1952) detenn1ned that the upper lethal
temperatures for pink salmon f~ (40 to 50 mm total length)
were about 21, 22.5, 23 and 24°C at acc11met1on temperatures
of 5, 10, 15 and 20°C, respectively. Preferred water
temperatures were about 11, 13 and l7°C at acc11mat1 on
temperatures of 10, 15 and 20°C, respectively. Low
temperature tolerance limits of pink salmon f~ were not
detem:1 ned.
--------~--
III. CONCEPTUAL SUITABILITY INDEX CURVES
The hab i tat suitability curves presented in this section are derived
pri mar i ly from threshold values reported i n the literature. Avoidance
behavior, abnormal development and increased mortality are expected to
occur if the threshold points are exceeded. A major limitation of· the
suitabi lity index curves is that they do not represent the increments
of habitat quality between the two threshold points . Because of the
paucity of data, particularly between the threshold points, the curves
are simplistic and have limited applicat~on.
The conceptual han ·~at suitability curves depict the inflllence of
certain physical and chemical lotic habitat components on the survival
and behavior of different pink salmon life stages . Most lotic habitat
components collectively influence all life stages. For example, water
permeability within the redd influences the i ntra-gravel environment
and the well-being of developing pink salmon eggs and alevins.
Dissolved oxygen and water temperatures also affect intra-gravel water
quality. Other lotic habitat components, sucn as current velocity,
water depth and substrates collectively influence spawning site
selection by f~.~le pink salmon .
The habitat sui tabi 1 ity curves presented in this document do not
account for interactive effects of chemical and physical parameters.
For example., a dissolved oxygen concentration of 5 mg/1 may be
suitable at a water temperature of 5°C, but unsuitable at a
temperature of 20°C. Ideally, a se.parate dissolved oxygen curve
should be drawn for each of several di fferent temperatures or
dissolved oxygen levels represented as percent saturation .
Additionally, the effect on the fish of less than optimum conditions
depends on the duration of exposure and on the particular life stage
of the fish.
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The conceptual habitat suitability curves are not constructed for
application to specific watersheds. Field data collection techniques
(when described) often varied. For example, three techniques for
measuring current velocity preferences of spawn~ng pink salmon were
found in the literature: the mean column velocity, the current
velocity 0.12 m above the redd and the current velocity 0.16 m above
the redd. These values may vary significantly among techniques. I,
addition, field data were presented as means, ranges, frequencies of
occurrence or the range of values utilized by a given percentage of
the sample population. Certain suitability index curves were based
upon laboratory tests, such as the water temperature--pre-gastrula
curve.
The value of each habitat component vhich was regarded as 'ideal' was
given a 1.0 rating on the Y-ax1s of the suitability index. The
threshold points were conn£cted by a solid line and dashed lines were
used to delimit the known upper and lower threshold points.
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Table 1 -AQUATIC HABITAT CRITERIA USED FO J>. CONCEPTUAL SUITABILITY INDEX CURVES
REGARDING UPSTREAM PASSAGE OF PINK SAlr«lN
Parameter
Water
temperature
Wate'r depth
Observed Values
0 21.1 c
O.lOtn
0.09 m
Re~~arks
Preferred water
temperatures for pink
salmon
May cause mtgratton
delay for pi nk salmn
Minimum passage re-
quirement for pink
salmon. Thts value
will change w1th actual
size and condition of
fish.
Pink salmon ,passage
noted along riffles
of several streaJ!s on
Kodiak Island less
than 0.09 m (0.3 ft)
Location
Oregon
Alaska
Be~l (1973)
Bell (1973)
fhOIIIJ.ISOO ( 1972)
Personal com-
nm1cation.
Jean Baldridge
(1981)
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---------------Table 2 -AQUATIC HABITAT CRITERIA USED FOR CONCEPTUAL SUITABILITY INDEX CURVES
REGARDING UPSTREAM PASSAGE OF PINk SALMON
Parameter
Water
teq>erature
Current
veloctty
Observed Values
7.2 -18°C
0.19-0.66 m/sec
0.35-0,47 m/sec
0.10 m/sec to
1.32 m/sec
0.37 m/sec to
1.0 m/sec
Remarks
Southeast Alaskan
stream temperatures
at time of pink
salmon spawning
Minimum value for
sucessful pink
salmon egg develop-
ment
Range of current
velocities chosen
by spawning pink
salmon. Mean column
velocity
Current velocities
preferred by spawn-
ing pink salmon.
Mean column velocity
Range of current
velocities chosen by
spawning pink salmon.
0.16 m above substrate
Current velocities
chosen by 80S of
detected spawning
pink salmon.
Location
Southeast
Alaska
Grace Crk.
eggs subjec-
ted to con-
trolled envi-
ronmental
conditions
Reference
Sheridan
( 1962b)
Ba1lel' I Evans
(1971)
Terrot• River, Wilson, Trihey,
Alaska Baldrfdve,Evans
Thiele &Trudgen
(lgal)
T~rror River Wilson, Trihey,
Alaska Baldridge,Evans
Skagit River
Washington
Skagtt River
Washington
Thiele &Tnadgen
(1981)
Graybill,Burgner
Gislason,Huffman
W,YNn, Gibbons,
kurko, Stober,
Fagnan, Stayman,
and E9gers
(lg79)
Graybill,Burgner
Gislason,Huffman
W,YNn, Gfbbons,
kurko, Stober,
F agna n , S tay11111 n
and E9gers
(lg7g)
---
Table 2 -AQUATIC HnBITAT CRITERIA USED FOR CONCEPTUAL SUITABILITY INDEX CURVES
cont•d REGARDING UPSTREAM PASSAGE OF PINK SALMON
Parameter Observed Values Remarks location Reference
0.21-0.99 m/sec Values measured 0.4 ft Washington Collings
above substrate State (1974)
0.3-0.8 m/sec Assorted streams; Sakhalin Dvfnin (1952)
method not stated Peninsula
USSR
0.45-0.73 m/sec Range occupied by first Jones Creek, Hourston and
100 ftsh in each channel Br1t. ColudJ1a MacKinnon
sect ton (1951)
0.39-0.64 m/sec Range occupied by first Jones Creek, Hourston and
spawning pair Brit. Columia MacKinnon
(1951)
Substrate 0.6-3.8 em i,-; Jones Crk. spawning channel Jones Creek, Hourston and
composition diameter primarily pink salmon Brit. Colullbia MacKinnon
• (1957)
N ...... Lucas (1959) • 2-10 em in Robertson Crk. spawning Robertson Crk.
diameter chanral Brit. Columia
2-250 mm in Range of substrate Terror River, Wilson et al.
diameter diameter utilized by Alaska (1981)
spawning pink sal~n
Water 0.2-0.3 m Valves for relatively Amur River Kuznetsov
depth fast and slow current USSR (1928)
velocity, respectively
0.5 m Uncrowded conditions Southern Dvinin (1952)
Sakhalin
USSR
0.1-1.2 m Crowded conditions Southern Dvinin (1952)
Sakhalin
USSR
-------------------Table 2 -AQUATIC HABITAT CRITERIA USED FOR CONCEPTU~l SUITABILITY INDEX CURVES
I
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cont•d REGARDING UPSTREAM PAS~AGE OF PINK SAlMON
Parameter
Water
depth
Observed Values
0.15-0.53 In
0.42 m
0.39 m
o.og-1.20 m
0.37-0.63 m
o.og-1.32 m
0.28-0.78 m
Remarks
Mean valve for sites
selected by first spawn-
ing pink salmon in each
of the 15 channel sections
Mean valves for sites
selected by first 100
within each of 15 channel
sections
Range of water depths
utilized by sp~ning pink
salmon
Preferred depths utilized
by spawning pink salmon
Range of water depths
utilized by spawning pink
salmon
Water depths utilized by
80% of spawning pink
location
various streams
in Washington
State
Reference
Collings
(1974)
Jones Creek Thurston and
spawning channel MacKinnon
Vancouver Is land
Brit. Columbia
Jones Creek Thurston and
spawning channel Mackinnon
Terror River,
kodiak Island,
Alaska
Terror A her.
Kouiak Island,
Alas~a
Skagit A her.
Washington
Skagit Rher,
Washington
Wilson,Trihey,
Baldridge,
Evans, Thiele,
Trudgen (1981)
llil son, Trihey,
Baldridge,
Evans, Th1 e 1 e,
Trudgen (1981)
GraybH 1 eta 1.
(1979)
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Table 3 • AQUATIC HABITAT CRITERIA USED FOR CONCEPTUAl SUITABiliTY INDEX CURVES
REGARDING UPSTREAM PASSAGE OF PINK SAlMON
Parameter Observed Values Relllilrks Location
Water I Survha 1-Water Laboratory controlled Egg source -
temperature Tempe 1rature env i romnent tests with Grace Creek
pink salmon eggs -near Ketchikan.
94-100% -~1ent % surviva 1 to Alaska
88-93% -4 5 c hatching stage. • o · 18-40% -3.00(
0% -2.0 c
Completion of gastrula
stage varied with water
temperature -1
Amb~ent -26th day,
4.50 C -45th day,
3.0 C -62nd day
Dissolved 6 mg/1 Hi nimum valve recomnende.d laboratory
oxygen for statsfactory develop-· tests using
ment of pink salmon eggs eggs fr0111
and alev1ns Sashin Creek,
Alaska
Reference
Battey & Evans
(1971)
Ba i1 ey , R 1 ce ,
Pella & Taylor
(1980)
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)(
Ill a
~
~ -_, -• 4
~ -:a •
1.0
,.... 2
1.0
0.1
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\
\
\
\
\
\
\
\
\
~I "I:.AII _.."......, •URATIOIII
OtrADULTI
See text for qualificathna
for use of these curves.
fNOT "ICOMMINDID II'Oa
A~~LICAT10N TO S~ICII'IC
WATiaSHID. WITHOUT II'IILD
YI:RI,ICATION}
' • 12 11 20 24
Water Te111perat .. :c
w .... , ...... ,.tw ........ ·-........... 5 -
~-------------------------t I
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0.2 o.4 0.1 a.a· 1.0 1.2 1.~
Wat•Deptb.•
s-..1. wat• d...._ tor .--•• of lltnk •••--
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)(
"' Q z -
>
~ _, -• c
~ -~
GD
1.0
0 5
Flgtn A
1.0
0.5
0
Fi~e. 5
1.0
0.5
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4 8 12 11 20
Watllr Temperat~n.-c
SPAWNING
See text for qualifications
tor use of these curves.
(NOT "ECOMMEN8ED FOR APPLICATION
TO SII'ECIFIC WATERSHEDS WITHOUT
II'IELD VEIIIIIfi'ICA TION)
0.2 1.4 0.1 0.1 1.0 1.2 1.4
c..r ... v.-.,. ml•c
cwr .... ladl ........ far_ ...... "* .-..on.
1
I \
I \
I \
I \
I \
I \
I \ I \ I
I \
1 2 ~ 4 5 I 1 I I
Suttatr .. Diameter. crw
Fltur• b SuDatrate condtuona auftable for Pink aalmon
IO&wnmg activity.
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• "' a z -
>
~ .... -• c
~ -::t
Ill
1 0
-
05
Ft ... 7
ALEVIN DEVELOPMENT
,....--?
PRE-GASTRULA STAGE
2 • a 10 12 14
Wat~ Te~noerature, •c
Eftlcta ot water t .. ~:~rature on Pink aalmon egg aurvtval
baaed uoon reaulta of Iaiiey and Evana.1871
1 o.,..-------1
05
0
Figure a
1.0
0.5
POST-GASTRULA STAGE
2 e • tO 12
Wa•r Temoerature. •c
Etfecta of water temoerature baaed on reaultl ot Bailey
and Evana. 1 871.
See text for qualifications
for uae of these curves.
(NOT RECOMMENDED POR APPLICATION
TO IPECIF'IC WATERSHEDS WITHOUT
PIELD YEAIF'ICATION}
1 2 3 4 5 I 7 8 9 10 11 12
OIIIOived Oxygen, mg/1
ftgute 9 Recommended dteaolved oxygen level• for aaUatactory
develoomentand aurvtval of Plnkaanon egga and alev1na.
IV. DEFICIENCY IN DATA BASE
Many measurements of aquatic habitat parameters could not be adapted
to habitat suitability curves. Most studies of current velocity,
water depth and substrate selection by spawning pink salmon did not
record the relative abundance (availability) of each habitat type.
The availability of these habitats, which vary within and among
streams, influences selection by spawning pink salmon.
Components of egg and alevin habitats were also usually not expressed
in a fonn which was usable for habitat suitability curves.
Intragravel flow, dissolved oxygen, substrate composition and other
aquatic habitat components which influence the survival and fitness of
pink salmon eggs and alevins are difficult to measure in the field.
Most aquatic habitat evaluations are conducted with the objective of
detennining upper and lower thresholds. Few studies examine the
relative growth rates, survival or overall habitat quality associated
with incremental chan~es in chemical and physical parameters between
threshold levels.
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V. RECOMMENDATIONS AND FURTHER STUDIES
Current velocity, water depth and substrate conditions selected by
female pink salmon for spawning should be measured and analyzed
throughout Alaska using standardized techniques. Past studies have
measured current velocity as mean column velocity or the velocity at
0.12 or 0.16 m above the redd. A focal point measurement (current
velocity at the fish's snout) combined with mean column veiocity at
the redd site would more realistically represent actual current
velocities.
There is a need for a standardized method of substrate classification
which evaluates substrate composition, predominate particle size,
s~ape, imbeddedness and angularity. Ideally the classification system
should be objective and not overly time consuming. Conmonly used
methods for substrate analysis usually require considerable time and
expense for collection, transportation and analysis.
Frequency analysis of pink salmon habitat preferences may be a useful
tool for constructing suitability curves; a frequency distribution is
comparable to a habitat suitability curve. Previously used frequency
distributions reflected stream or stream-reach habitat availability
(Wilson, Trihey, Baldrige, Evans, Thiele and Trudgen, 1981} rather
than habitat preferences of the fish.
Pink salmon spawning habitat is a composite of available current
velocity, water depth and substrate conditions. To better understand
constituents of pink salmon spawning habitat within a stream all
available habitat should be inventoried, whether ft is utilized or
nat.
A variety of lotic systems from small, clearwater streams to larger
clearwater and glacial streams should be examined in variou~
geographica 1 areas of Alaska to better determine aquatic habitat
preferences of spawning pink salmon. These investigations, ideally,
-34-
should encompass both •on• and 'off' stocks. Aquatic habitat-pink
salmon relationships will be very different among different papulation
densities.
Laboratory experiments investigating the effects of various substrate
compositions on intragravel flow, dissolved oxygen and survival of
pink salmn eggs and alevins should be initiated. Field studies
should compliment laboratory research in assessing the inf1uenc8l of
fines on pink salmon produ~tion. The contribution of fine s•;bstrate
material to pink salmon spawning streams by various land use
act ivities should. be detennined.
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VI. LITERATURE CITED
Alaska Department of Fish and Game. 1978. Alas.ka's Fisheries Atlas.
Alaska Dept. of Fish and Game, Vol. II., 196 p.
Bailey, J.E. 1969. Effects of salinity on intertidal pink salmon
survival. Alaska Dept. o.f Fish and Game Infonnational Leaflet
187., p. 12-15.
Bailey, J.E. and D.R. Evans. 1971. The low temperature threshold for
pink salmon eggs in relation to a proposed hydroelectric
installation . U.S. Fish and Wildlife Service, Fish Bulletin
69(3) :587-593.
Bailey, J.E., S. Rice, J. Pella and S. Taylor •. 1980. Effect,s of
seeding density of pink salmon, Oncorhynchus gorbuscha, eggs on
water chemistry, fry characteristics and fry survival in gravel
incubators. Fish Bulletin U.S. 78:649-658.
Banks, J.W. 1968. A review of the literature on the upstream
migration of adult salmonfds. J . Fish Biol. 1(2):85-136
Bell, M.C. 1973. Fisheries handbook of engineering requirements and
biological criteria . Useful factors in life history of most
conmon species. Submitted to Fish .-Eng . Res . Program, Corps of
Engineers, North Pac. Div., Portland, Oregon. (unpublished).
Bjorn, T.C. 1969. Embryo survival and emergence studies. Job No. 5,
Federal Aid in Fish Restoration . Job Completion Report. Proj.
F-49-R-7. Idaho Fish and Game Depart., Boise. 11 p.
Coble, D.W. 1961. Influence of water exchange and dissolved oxygen
in redds on survival of steelhead trout embryos . Trans . Amer.
Fish. Soc . 90:469· ~74
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VI. LITERATURE CITED
Alaska Department of Fish and Game. 1978. Alaska's Fisheries Atlas.
Alaska Dept. of Fish and Game, Vol. II., 196 p.
Bailey, J.E. 1969. Effects of salinity on intertidal pink salmon
survival. Alaska Dept. of Fish and Game Informational Leaflet
187., p. 12-15.
Bailey, J.E. and D.R. Evans. 1971. The low temperature threshold for
pink salmon eggs in relation to a proposed hydroelectric
installation. U.S. Fish ard Wildlife Service, Fish Bulletin
69{3}:587-593.
Bailey, J.E., S. Rice, J. Pella and S. Taylor .. 1980. Effects of
seeding density of pink salmon, Oncorhynchus gorbuscha, eggs on
water chemistry, fry characteristics and fry survival in gravel
incubators. Fish Bulletin U.S. 78:649-658.
Banks, J.W. 1968. A review of the literature on the upstream
migration of adult salmonids. J. Fish Biol. 1(2):85-136
Bell, M.C. 1973. Fisheries handbook of engineering requirements and
biological criteria. Useful factors in life history of most
common species. Submitted to Fish.-Eng. Res. Program, Corps of
Engineers, North Pac. Div., Portland, Oregon. (unpublished).
Bjorn, T.C. 1969. Embryo survival and emergence studies. Job No. 5,
Federal Aid in Fish Restoration. Job Completion Report. Proj.
F-49-R-7. Idaho Fish and Game Depart., Boise. 11 p.
Coble, D.W. 1961. Influence of water exchange and dissolved oxygen
in redds on survival of steelhead trout embryos. Trans. Amer.
Fish. Soc. 90:469-~74
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Collings, M.R. 1974. Generalization of spawning and rearing
discharges for several salmon species in western Washington.
U.S. Geological Survey. Open-file report. 39 p.
Combs, B.D.
eggs.
1965. Effect of temperature on the development of sal.on
Prog. Fish-Cult. 27:134-137.
Cordone, A.J. and D.W. Kelly. 1961. The influence of inorganic
sediment on the aquatic life of streams. Calif. Fish and Game.
47(2). 189-228.
Davidson, P and L. Vaughan. 1943. Factors affecting the upstream
migration of pink salmon. Ecology 24:149-168
D~inin, P.A. 1952. The salmon on South Sakhalin. Inz. TINRO.
37:69-108
Godin, T-6, 6 1980. Temporal aspects of juvenile pink salmon,
Onchrhynchus gorbuscha, (Walbaum) emergence from a simulated
gravel redd. Can. J. Zool. 58(5}:735-744
Graybill, J., R. Burgner, J. Gislason, P. Huffman, K. Wyman, R.
Gibbons, k. kurko, Q. Stober, T. Fagnan, A. Stayman and D.
Eggers. 1979. Assessment of the reservoir-related effects of
the Skagit project on downstream fishe~ resources of the Skagit
River, Washington. Final Report for City of Seattle, Dept. of
Lighting, Seattle, Washington. Univ. of Washington, College of
Fisheries. Fisheries Research Institute. p. 595
Hanavan, R.M. 6. and 8.1. Skud. 1954. Intertidal spawning of pink
salmon. U.S. Fish and Wildlife Service, Fish Bulletin 56:167-186
Hays, F.R., I.R. Wilmot, and D.A. Livingstone. 1951. Oxygen
consumption of the salmon egg in relatfon to development ana
activity. J. of Exp. Zool. 116:337-396
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Helle, J.H., R.S. Williamson and J .E. Bailey. 1964 . Intertidal
ecology and life history of pi nk salmon at Olsen Creek, Prince
William Sound, Alaska. U.S. Fish and Wildlife Serv i ce, Spec.
Sc i . Rep. Fish 483 . IV+ 26P.
Hoar, W.S. 1956 . The behavior of migrating pink aM chum salmon fry.
J. Fish. Res. Bd. Can. 13(3):309-325 .
Hourston, W.R. and D. MacKinnon. 1957. Use of an artificial spawning
channel by salmon . Trans . Amer. Fish. Soc :220-230.
Hunter, J.G. 1959. Survival and pr.oduction of pink and chum salmon
in a coastal stream . J. Fish . Res. Bd. Canada 16:835-886 .
Kirkwood, J.B. 1952 . Inshore-marine and f ·reshwater life history
phases of the pi nk salmon and chum salmon in Prince William
Sound, Alaska. Ph.D. Dissertation , Univ. of Louisvill, 300 p.
Koski, K.V . 1966. Survhal of coho salmon (Oncorhynchus kisutch)
from egg deposition to emergence in three Oregon coastal streams.
M.S. Thesis, Univ. of Or egon, Corvallis. 84 p.
Krok~in, E.M. and F.V. Kroquis. 1957. Study of the Bolshaia River
basin and its salmon spawning grounds. Izv. T1nro, 9, 156 p.
Kuznetsov, I .I. 1928. Some observations on the spawning of Amur and
Kamchatka salmon. Izv . Tinro. 2:195 p.
Luca s , K.C. 1960 . The .Robertson Creek spawning channel. Canadian
Fish . Cult. 26 :3-23.
Ma c:Donald, J. 1960. The behavior of Pacific salmon fry during their
downstreJm migration to freshwater and saltwater nursery areas.
J. Fi sh. Res . Bd. Canada, 17(5):655-676 .
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McCuddin, M.E . 1977 . Survival of salmon and trout embryos and fry in
gravel-sand mixtures. M.S. Thesis, Univ. Idaho, Moscow. 30 p.
McNeil, William J. 1962. Mortality of pink and chum salmon eggs and
larvae in southeast Alaska streams . Ph.D . Thesis, Univ. of
Washington, Seattle . 270 p.
McNeil, W.J. and J.E. Bailey. 1975. Salmon ranc.her's manual,
Northwest Fish Center, Auke Bay Fish. Lab. Processed Report. 95p.
Morrow, J.E. 1980. The freshwater fishes of Alaska. Alaska
Northwest Pub. Co., Anchorage. 248 p.
Neave, F. 1966. Pink salmon in British Columbia. A review of the
life history of North Pacific salrncr,. Int. Pac. Salmon Fish.
Comm ., Bull. 18:70-79.
Noerenburg, Wallace A. 1963. Salmon forecast studies on 1963 runs in
Prince William Sound. Alaska Oepartmnt of Fish and Game.
InfoMmational Leaflet No. 21. 28 p.
Phillips, R.W. and H.J. Campbell. 1961. The embryonic surv i val of
coho salmon and steelhead trout as influe~ced by some
environmental conditions in gravel beds . 14th Ann. Rep . Pac.
Mar. Fish. Comm., Portland:60-73.
Prichard, A.L. 1.948. Efficiency of natural propagation of the
pink salmon (Onchrynchus gorbuscha) in McClinton Creek, Masset
Inlet, British Columbia. J. Fish Res. Bd . Canada 7(5):224-236.
Reiser, D.W. and T.C. Bjornn. 1979. Habitat requirements of
anadromous. salmonids. USDA Forest Service. Gen. Tech. Rep.
PNW-96. 54 p.
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Scott, W.B. and E. J. Crossman. 1973. Freshwater f.tshes of Canada.
· Bull. Fish. Res. Bd. Canada No . 184. 966 p.
Semko, R.S. 1939. Kamchatka pink salmon, Izv. Tinro, 16 . 111 p.
Sheridan, W. 1962a. Waterflow through a salmon spawning riffle in
Southeastern Alas.ka. u.s. Fis,h Wildl. Serv., Spec. Sci. Rep.
Fish. No. 407. 20 p.
Sheridan, W. 1962b. Relation of stream temperatures in southeast
Alaska in. N.J. Williamovsky (editor). Symposium of pink
salmon. p. 101-117, H.R. MacMillan Lect. Fish., Univ. of British
Columbia, Vancouver
Thompson, K.E. 1972. Determining streamflows for fish life. In.
Proc. Instream Flow Requirement Workshop, Pacific N.W. River
Basins Comm. p. 85-103
Vaux, W.G. 1968. Intrag'ravel flow and interchange in a streambed .
U.S. Fish Wildl. Serv., Fish . Bull . 66:479-489 .
Webb, J. 1978a. Fisheries inventory of the Shaktoolik River, Alaska.
Fairbanks District, Bureau of Land Management . 13 p.
Webb, J. 1978b. Fisheries inventory of the Buckland River, Alaska.
Fairbanks Dis.trict, Bureau of Land Management. 10 p.
Webb, J. 1980. Fisheries inventory of the Inglutalik River, Alaska .
Fairbanks District, Bureau of Land Management. 10 p.
Wickett, W.P. 1958. Review of certain environmental factors
affecting the 9roduction of pink and chum salmon. J. Fish . Res.
Bd . Can. 15:1103-1123.
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Wilson W •• W. Trihey , J. ~ldrigC C. Evans, J. Thiele and D. Trudgen.
1981. An assessment of environmental effects of construction and
operation of the proposed. Terror Lake hydroelectric facility,
Kodiak, Alaska. Arctic Environmental Information and Data
Center, Anchorage, Alaska . 419 p.
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I FRESHWATER HABIT AT
1 RELATIONSHIPS
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BROAD WHITEFISH-COREGONUS NASUS
~
I ALASKA DEPARTMENT OF FISH & ~E
HABITAT PROTECTION SECTION I RESOURCE ASSESSMENT BRANCH
I APRIL, 1981
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FRESHWATER HABITAT RELATIONSHIPS
BROAD WHITEF ISH -COREGONUS NASUS
By
Stephen S. Hale
Alaska Department of Fi sh and Game
Habitat Division
Resource Assessment Branch
570 West S3rd Street
Anchorage, Alaska 99502
May 1981
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ACKNOWLEDGEMENTS
Many people from the Alaska Department of Fish and Ga~ and from the Auke
Bay Fisheries Laboratory of the National Marine Fisheries Service freely
gave their ti.e and assistance when contacted about this project and 1t is
a pleasure to thank them and fishe~ biologists from other agencies,
especially those who provided unpublished data and observations from their
own work. The librarians of the Alaska Resources Library and the U.S. Fish
and Wildlife Service were of great help.
This project was funded by the U.S. Fish and Wildlife Service, Western
Energy and Land Use Team, Habitat Evaluation Procedure Group, Fort Collins,
Colorado. Contract No. 14-16-0009-79-119.
TABLE OF CONTENTS
BROAD WI~ITE'FISH
Page
I. Introduction 1
A. Purpose 1
B. Distribution 2
c. Life History Summ~ry 4
D. Ecological and Economic Import-ance 8
II. Specific Habitat Requirements 9
A. Adults 9
1. Spawning migration 10
2. Spawning 10
3. Oventintering 11
B. Incubation of Embryos 11
c. Juvenile Rearing 12
III. Suitability Index Curves 13
IV. Deficiencies in Data Base and Recommendations 16
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I. INTRODUCTION
A. Purpose
This report presents available infonAition on the freshwater
habitrt requirements of the broad whitefish, Coregonus nasus
(Pallas) and evaluates the habitat parueters which are lalSt
important to the species or are most often critical to survival
or li•iting to production.
Because the range of the species includes coastal brackish waters
which are used as SUIIRr feeding areas or as overtrfintering areas,
information from such areas was also considered.
The emphasis of this report is on habitat requirements, primarily
those of a physical or chenical nature. Certain Mological
factors affecting the well-being of the population such as
feeding, predation, competition, and disease are not
comprehensively treated.
This report is intended to support habitat evaluation activities
by presenting a data base for the species and by pointing out
where more data are needed.
Although information has been examined from throughout the range
of the species, emphasis is placed upon Alaska. While there
appears to be wide differences in growth rates between different
populations within the state, there is insufficient information
to show that habitat requirements differ among the various
populations. There are some problems with the taxon~ of the
species. Information referring to Coregonus nasus (Pallas) sensu
Svardson in Scandinavia was not used because that species
evidently is different from the Coregonus nasus (Pallas) of
Siberia, Alaska, and northwest Canada.
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The life history of the broad whitefish, especially that of the
early life stages, is not well known. Even less is known of
habitat to 1 erances, preferences, and requirements. 'The b·road
whitefish appears to ha.ve fairly wide habitat tolerances.
Several reports on the life histories of related species are
presented in the Biology of Coregonid Fhhes, edited by Lindsey
and Woods {1970).
B. Distribution
The broa.d whitefish is a lacustrine-fluvial species, but is found
more ·often in rivers than i n lakes. It also occurs in brack1 sh
areas of c.oa s ta 1 waters.
It is distributed in North America in Bering Sea and Arctic Ocean .
drainages from the Kuskokwim River system in Alaska north and
east to the Perry River, Northwest Te·rritories. In the USSR, it
is distributed from the Pechora River· near the Ural Mountains
east in Arctic Ocean drainages and south in Bering Sea drainages
to the Bay of K,i'f, and in the Penzhina River on the northeastern
corner of the Sea of Okhotsk (Berg, 1948; McPha i1 and Lindsey,
1970; Scott and Crossman, 1973). The southern limit inland is
approximately 60° N.
In Alaska, the broad whitefish is found in most dra1nages north
of the Kus .kokwim Rh1er systeat, where it is conmon, and the Alaska
Range. (Fi ·gure 1). It occurs in the Yukon Riv·er system from the
mouth to the headwaters in British Columbia (Morrow. 1980),
including the Koyukuk River and Porcupine River drainages. It h
COIIIII)n in the !itinto Flats area of the Tanana River systen but
apparently unconmon further upstre.am. It is widespread in the
drainages emptying into the Be·ring Sea (Kuskokwim River and
north} , Chukchi Sea, and Beaufort Sea . The Sus i tna River a.nd the
Cop~er River drainages have no broad whitefish {Alt, 1971).
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Marn Study Sites
[!] Col v i lie Ri ver
[f) Sagavanirktok At vet
{]] M i nto Flats
I!J Yukon-K vsk~kwim Del to
ST l..-..ENCE
~ANO
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F.gura 1 Orsrrrbutron of broad whitefrsh in Alaska (R .. Baxter,
personal communecahon; Morrow, 1980} and main study sites.
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C. Life History Sunnary
The Hfe history of the broad whitefish is not well known.
Migrations occur between summer feeding areas, spawning areas,
and overwintering areas. In general, spawning migrations occur
in the late sua.r end fall as the fish move out of s.-.er
feeding areas and .ave to spawning areas. Baxter (1973) has
noted that, in the Kuskokwi• River area, the ripening fa-ales
move downst~ out of the tundra lakes, ponds, and streams in
August and September and begin a slow migration up the Kuskokwim
River. They are followed by the sexually developing males in
September and by the non-spawning adults of both sexes in late
September through October. I..ature fish leave the tundra in
October through Decemer. Broad whitefish apparently migrate
downstream out of the Minto Flats into the Tanana River in August
(Kepler, 1973; Townsend and Kepler, 1974). On the North Slope,
fish that had been feeding in coastal areas enter the
Sagavanirktok River in late August to migrate to the spawning
a rea s (Ben dock, 1977 ) . A s i zeab 1 e spawning run moves up the
Colville River in August (Bendock, 1979). Alt and Kogl (1973)
found that the Colville run is spread over several months and
peaks in late July.
Wynne-Edwards (1952, cited by McPhail and Lindsey, 1970) stated
that upstream spawning migration of broad whitefish occurred in
the lower Mackenzie River in July and August. Evidently, the
m1 gra t ion up the Meckenz i e peaks in the inner de 1 ta during
September and October (DeGraaf and Machniak, 1977). In the upper
Yukon River, broz.tl whitefish on spawning migrations have been
observed entering small tributaries in September; ripe fish have
been captured in the main river in early October (McPhai 1 and
Lindsey, 1970}. Spawning in the Mackenzie River apparently takes
place in back eddies during October (DeGraaf and Machniak, 1977).
Spawning in the USSR takes place in October and November (Berg,
1948).
C. Life History Sunnary
The Hfe history of the broad whitefish is not well known.
Migrations occur between summer feeding areas, spawning areas,
and overwintering areas. In general, spawning migrations occur
in the late sua.r end fall as the fish move out of s.-.er
feeding areas and .ave to spawning areas. Baxter (1973) has
noted that, in the Kuskokwi• River area, the ripening fa-ales
move downst~ out of the tundra lakes, ponds, and streams in
August and September and begin a slow migration up the Kuskokwim
River. They are followed by the sexually developing males in
September and by the non-spawning adults of both sexes in late
September through October. I..ature fish leave the tundra in
October through Decemer. Broad whitefish apparently migrate
downstream out of the Minto Flats into the Tanana River in August
(Kepler, 1973; Townsend and Kepler, 1974). On the North Slope,
fish that had been feeding in coastal areas enter the
Sagavanirktok River in late August to migrate to the spawning
a rea s (Ben dock, 1977 ) . A s i zeab 1 e spawning run moves up the
Colville River in August (Bendock, 1979). Alt and Kogl (1973)
found that the Colville run is spread over several months and
peaks in late July.
Wynne-Edwards (1952, cited by McPhail and Lindsey, 1970) stated
that upstream spawning migration of broad whitefish occurred in
the lower Mackenzie River in July and August. Evidently, the
m1 gra t ion up the Meckenz i e peaks in the inner de 1 ta during
September and October (DeGraaf and Machniak, 1977). In the upper
Yukon River, broz.tl whitefish on spawning migrations have been
observed entering small tributaries in September; ripe fish have
been captured in the main river in early October (McPhai 1 and
Lindsey, 1970}. Spawning in the Mackenzie River apparently takes
place in back eddies during October (DeGraaf and Machniak, 1977).
Spawning in the USSR takes place in October and November (Berg,
1948).
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Morrow (1980) states that spawning of the broad whitefish in
Alaska occurs in Septener and October and possibly into
November. Baxter (lg73) found that the spawning season in the
1 ower Kuskokwim River lasts from October to early December.
Adults do not spawn every year. Broad whitefish in the Colville
River apparently spawn in September in the upper section of the
river (Alt and Kogl, 1973). Cohen (1954) reported that broad
whitefish (Coregonus nasus kennicotti) spawn in Ikroavik Lake
near Barrow in July and possibly 1n June.
Spawr.ing usually takes place in rivers. In Allskil, there are
spawning populations of broad whitefish in Lake Minchumina
(Baxter. 1g73) and possibl .v also in the K111ick Lakes of the
North Slope (Terry Bendock, personal co111111nication). Cohen
(1954) reported a spawning population of broad whitefish in
Ikroavik Lake. Kuz'min (1969) reported that an attempt to raise
broad whitefish in likes from spawn taken from rivers was not
successful because the f~les did not COMe to sexual maturity.
Kuz'min suggested that running water is a requirement for
successful reproduction of these populations.
Broad whitefish broadcast their eggs over substrates ranging from
IIRJd and sand to gravel and cobble (Baxter, 1973; Kogl, 1971;
Morrow, 1980). Little else is known about their spawning habitat
needs.
The fecundity of a sample of 11 females from the Mackenzie River
ranged ~rom 26,922 to 65,798 eggs/feaale with a mean of 39,721
eggs/¥emale (DeGraaf and Machniak, 1977). Three fish from the
Kuskokwim River had fecundities ranging from an estimated 46,219
to 127,707 eggs/female (Baxter, 1973).
Apparently, in several stocks, there is a post-spawning
downstream migration of adults to overwintering ar'!as in deep
sections of rivers or in brackish water areas or lakes (Baxter,
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personal c~nicetion; Bendock, 1977; Berg, 1948; Morrow, 1980;
Scott and Cross11111n, 1973). In the Mackenzie River, this
downstream migration occurs in early November. Broad whitefish
ove~inter in the outer Mackenzie Delta and in lakes of the inner
Delta (DeGraaf and Machniak, 1977).
In the lower Kuskokwim River, the population which ove~inters in
the 11111in river migrates back to the summer feeding areas of the
tundra system in late May or early June after spring nmoff
(Baxter, lg73). A similar migration occurs in the Minto Flats
area. Broad whitefish move in June from the Tanana, Tolovana,
and Chatanika Rivers to feed in the lakes and sloughs of the
Flats (Kepler, 1973). On the North Slope, some broad whitefish
migrate out of the larger rivers such as the Colville, Canning
and Sagavanirktok when these rivers break up in early June and
move into shallow bays and lagoons of the Beaufort Sea for summer
feeding (Bendock, 1977). Other broad whitefish remain in the
rivers throughout the su.mer.
Young broad whitefish hatch in the spring and apparently migrate
to summer feeding areas which may include lakes (Baxter, 1973;
Berg, 1948; Morrow, 1980). There is no info~tion on whether or
not the young-of-the-year fish have rearing areas separate from
the immature and adult fish.
The broad whitefish is apparently a bottom feeder. The diet
inc 1 udes gastropods, pe 1 ecypods • chi ronom1 ds inc 1 udi ng midge
larvae and pupae, msquito larvae, other dipteran larvae and
adults, terrestrial insects, trichopterans, aq)hipods, mysids,
copepods, and phytoplankton (Bendock, 1977 and 1979; Berg, 1948;
Boughton and Clemens, 1966; DeGraaf and Machniak, 1977; Furn;ss,
1975; McPhail and Lindsey, 1970; Scott and Crossman, 1973}.
Apparently, little feeding, if any, occurs during overwintering.
Sto11111chs of fish captured wh i 1 e overwintering in the Co lv i 11 e
River {Bendock, personal communication; Kogl and Schell, 1975)
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and the Saaavanirktok River (Bendock, 1977) have been empty. In
the Kuskokwim River area, the fish do not feed from the time they
leave the tundra system in the fall until the time they return to
the tundra lakes and strea.s in the spring (Baxter, 1973). Stein
et al. (H73, cited by DeGraaf and Mact'l .. iak, 1977) collected
broad whitefish from lakes in the Mackenzie Delta during the
summer. They reported that a lower percentage of fish taken from
lakes had empty stcachs than those taken from the Mackenzie
River, indicating the i~rtance of lakes as feeding areas •
Most broad whitefish reach sexual maturity at about age five to
seven (Alt, 1976; Berg, 1948). At that time, they are around 35
-SO em fork length and weigh around 0.5-2 kg (Alt, 1976;
Bendock, 1979; Berg, 1948). Specimens have been captured in
Siberia weighing up to 12 kg and perhaps 16 kg• the longest fish
reported was 86 em long {Berg, 1948). The mexiiiiJIII age reported
to date is 22 years (Craig and Haldorson, 1980).
The broad whitefish is the largest of the Alaskan whitefish
(Morrow, 1980) except for the sheefish. The maximum length
reported for most populations of Alaskan broad whitefish is
around 65 em fork length (Alt, 1976; Bendock, 1979; Kogl, 1971);
the largest fish from the Kuskokwim River was 72.4 em fork length
and weighed 7.6 kg ( R. Baxter, personal c0111111nication). Most
Alaskan broad wtdtefish captured are between 25-60 em fort
length, between 0.5 -2.5 kg in body weight, and are four to
eight years of age (Alt, l976i Bendock, 1979; Furniss, 1975).
The average weight of fish in the commercial harvest in the lower
Kuskokwim is 3.3 kg (R. Baxter, personal communication). There
are considerable differences in growth rates and in size of broad
whitefish among different geographical areas.
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D. Ecological and Economic l!pOrtance
The broad whitefish is of excellent quality as a food fish and is
important in ca..ercial and subsistence fisheries. ln Alaska, a
limited con~~ercial s~~~~~ner fishery on the Co 1 ville River de 1 ta
takes about 3000 broad whitefish or about 7,000 kg per year (Alt
and kogl, 1973). The broad whitefish is the target species in
the Nechelik Channel of the Colville (Kogl and Schell, 1975).
Another small commercial fishery exists in the Yukon-Kuskokwim
Delta area. The broad whitefish is widely used throughout its
range in Alaska for subsistence food. ln the Kuskokwim River
delta area, it is second only to salmon in importance (Baxter,
1973). It is taken year round by gill nets; in the summer, fish
fences and dip nets are also used. Small numbers are taken each
year in Alaska by sportfishermen.
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II. SPECIFIC HABITAT REQUIREMENTS
A. Adults
Broad whitefish have been captured in Alaska and northwest Canada
in water temperatures ranging fro11 0 to l6°C (Bendock~ 1977•
Craig and Haldorson, 1980; Kogl, 1971; Muth, 1969). They nay
tolerate summer temperatures in shallow ponds of the Kuskokwim
River delta up to about zooc {R. ~axter, personal communication).
There is no information on preferred temperatures, but fish from
the Mackenzie River (0 to 15.5°C annual range} have a greater
growth rate than Coppe~ine River fish (0 to 10.0°C annual range)
which may suggest that summer temperatures above 10°C are more
favorable for growth (Muth, 1969). The longer growing season and
the greater food abundance in the Mackenzie could also be
factors. Alt (1976) suggests that the slower growth rate of
broad whitefish from the Sagavanirktok R1Yer and Ina~ruk Basin
(Seward Peninsula) as compared to other populations within Alaska
may be a result of the shorter ice-free period in those two
areas.
Broad whitefish in Alaska generally occur in streams where the
gradient is less than 0.75 m/km (Alaska Department of Fish and
Game, unpublished manuscript; Kogl, 1971). Current velocities
for several bodies of water on the North Slope where broad
whitefish occur range from 0 to around 180 em/sec. (Kogl~ 1971).
The broad whitefish occurs in Alaska in tundra ponds where the pH
can range down to pH 5.5 or 6.0 (K. Alt, personal communication;
R. Baxter, personal c~nication). They also occur in lakes
where the pH at the surface can range upward to 8.1 (Boughton and
Clemens, 1966). However, the exact range that they can tolerate
or that they prefer is unknown.
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Broad whitefish have been caught in waters with salinities oi:
11.4 to 40.8 parts per thousand {ppt} in April on the Colvil e
River Delta (Kogl and Schell, 1975), 0-30 ppt in coastal areas of
the North S1 ope (Craig and Ha 1 dorson, 1980). 4-28 ppt in Prudhoe
Bay (Bendock, 1977), and occa.sionally in brackish water of 5 ·6
ppt in coastal areas of Siberia (Berg, 1948). Alt (1976) states
that they are seldom taken in waters of greater than .20 ppt
salinity. They have been caught in waters of the North Slope
with turbidities of 2-1 ·46 NTU (Craig and Haldorson, 1980) a11d
5-10 ppm (Kogl, 1971).
1. Spawning Migration
In laboratory experiments at 12 -l3°C, Jones {197?) fcund
that the critical velocity for broad whitefish with a f orlc
length of about 35 em was approximately 65 em/sec. Jo1es
projected that adult broad whitefish could move 100 m i' 10
minutes against a current velocity of 40 em/sec. Upstream
migrations could be inhibited by higher current velocities
or stream reaches longer than 100 m with velocities in :he
range of 40 em/sec. However, the fish occur in the sunuer
in streams of the North Slope which have current velocities
of 80 to 180 em/sec (Kogl, 1971).
2. Spawning
Broad whitefish spawn at water temperatures close to ooc 'K.
Alt, personal conmunfcation; R. Baxter, personal
communication; Berg, 1948). Spawning takes place at abolt
the time of freezing of 1 akes and streams. The upper
temperature limit is unknown.
Spawning takes place over substrates ranging from mud an<
sand to gravel, cobble, and boulders with perhaps fine
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gravel being the most connan substrate (Alt, personal
communication; Baxter, personal communication; Kogl, 1971).
Running water prior to spawning may be a requirement of
certain stocks of broad whitefish for successful sexual
deve 1 opment of fema 1 es Kuz • min, 1969). However, there
appear to be strictly lacustrine stocks. Examples in Alaska
are the lake Minchumina population which evidently spawns on
gravel and cobble along the lake shore (R. Baxter, personal
conmunicat1on) and the lkroavik lake population (Cohen,
1954).
3. Overwintering
Broad whitefish overwinter at 0°C with no apparent ill
effects (Baxter, 1973; Bendock, 1977). Kogl and Schell
(1975) found broad whitefish in the Colville Delta in April
where at dissolved oxygen concentrations were 2.3 to 7.8
mg/1 and Bendock (1980) has found them overwintering in
holes of the Colville River where dissolved oxygen levels
were 1.4 to 4.6 mg/1. Fish found at the lower dissolved
oxygen level aopeared healthy. Baxter (1973) stated that 2
mg o2;1 is apparently the minimum oxygen level tolerable for
Kusko~im River broad whitefish.
B. Incubation of Embryos
Very little information is available for the embryonic stage of
the broad whitefish. Because of their geographic location and
time of spawning, must incubate at water temoeratures close to
0°C. However, Cohen (1954) st~tes that broad whitefish
{Coregonus nasus kennicotti) spawned in lkroavik Lake near Barrow
in ,)ul,v; the eggs developed at 6-l2°C and hatched in 30-60 days.
-11 ..
. --------~
C. Juvenile Rearing
Young of the year and juveniles apparently mix with the adults.
It is not presently known 1~ there are dif~erence~ in habitat
requ1 rements among these 1i fe stages.
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I!I. SUITABILITY INDEX CURVES
No su~tabil ity index curves were drawn for the brtJad ~thitefish because
there is not sufficient information to construct meaningful curves.
The ~~tide distribution of this species in habitats with differing
ranges of habitat parameters further c~ounds the problem. Tables I
and II provide a general overview of certain physical and chemical
parameters •
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I!I. SUITABILITY INDEX CURVES
No su~tabil ity index curves were drawn for the brtJad 'tthitefish because
there is not sufficient information to construct meaningful curves.
The ~~tide distribution of this species in habitats with differing
ranges of habitat parameters further c~ounds the problem. Tables I
and II provide a general overview of certain physical and chemical
parameters •
.. lJ ..
RROAO ~HITEFI~tl Tahle 1. Adults
--
f'arilml"ter Ob~erved Value~ Remarks location Reference
Temperature, 0 overr~interinq Kuskokwim River Raxter (1973}
"C area
0 -overwi nteri nq Colville and Oenrlock ( 1977 l
Saqavanirktok
Rivers
0 -15.5 annual ranqe Machnzie River Muth ( 1 969)
0 -10 .o annual range Copperm1ne River Muth (19fiCJ)
0 -14 temperature at t1~Tte of capture North Slope Craig and Haldorson
(1980)
14 -16 su111ner range North Slope Kogl ( 1971)
may go up to 20 tundra ponds Yukon-Kuskowim R. Baxter (personal
I Delta c Olllllln i r. a t1 on ) __,
t
01 ssolved 2.3-7.8 overwi n te ring Colville River Kogl and Schell
Oxygen, mg/1 Oelta ( 1975)
1.4 -4.6 overwintering Co lv 111 e R i ve r Bendock ( 1980)
7. minimum tolerable at ouc Kuslrokwim River Buter (1973)
Salinity, 11.4 -40.8 April Colville River l(ogl and Schell
ppt. Oelta (1975)
0 -30 Co as ta 1 areas of Cra i q ancl Ha 1 riorson
North Slope (1980)
4 -28 Prudhoe Bay 8endock (1977}
5 - 6
Siberia Berg ( 1948)
• ......
U'1 •
---------13RO/\O WIIJTfFJSII lrblr. II. Spawnin~
Parameter Ohsr?rvcrl Values Re1t1c1rks
Temperature, ~rounrl 0 oc
close to 0
0 - 4
- ----
toci'Uon
Kuskokwirn River
USSR
Alaska
----
Referertre
R. B~xter (person~l
comnunir.at.ion)
Rerq ( 1948)
K. Alt (personal
con11lJn i cat 1 on)
-
IV. DEFICIEtlC!ES IN [l.ATA BASE AND RECOt.f~ENDATIONS
Because so little is known about the habitat tolerances, preferences,
and needs of the broad whitefish, infonnation is needed on almost all
aspects. There is a need for basic descriptive life history studies~
extensive measurements of habitat parameters, and phys i ological
experil'lents in the laboratory.
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LITERAiURE CITED
Al~ska Departrne~t of Fish and Game (unpublished manuscript). Whitefish
investigations of the Yukon-Kuskokwim Delta. Cotrmercial Fisheries
Division~ Bethel.
Alt, Kenneth T. 1971. Distribution, movements, age and growth, and
taxonomic status of whitefish {Coregonus !2.:._) in the Tanana-Yukon
drainage and North Slope. Alaska Department of Fish and Game.
Federal Aid in Fish Restoration~ Project F-9-3, Study R-11, Job
R-11-F, Vol. 12: 19-31.
Alt, Kenneth T. 1976. Age and growth of Alaskan broad whitefish, Coregonus
nasus. Trans. Am. Fish Soc. 1D5:526-528.
Alt, Kenneth T., and Dennis R. Kogl~ 1973. Notes on the whitefish of the
Colville River, Alaska. J. Fish. Res. Bd. Can. 3D (4):554-556.
Baxter, Rae, 1973. Coregonus nasus (Pallas). Alaska Department of Fish
and Game, Commercial Fisheries Division, Bethel.
Bendock, Terrence N., 1977. Beaufort Sea estuarine fishery study. Final
Report DCS Contract No. D3-5-022-69. Alaska Department of Fish and
Game, Fairbanks.
Bendock, Terrence N., 1979. Inventory and cataloging of Arctic area
waters. Alaska Department of Fish and Game. Federal Aid in Fish
Restoration, Study No. Gl, Job No. G-1-l, Vol. 2D: 1-64.
Bendock, Terrence N., 198D. Inventory and cataloging of Arctic area
waters. Alaska Department of Fish and Game, Federal Aid in Fish
Restoration, Study No. Gl, Job No. G-1-l.
Berg, Leo S. 1948. Freshwater fishes of the USSR and adjacent countries.
(Translated from Russian). Vol. 1, Acad. Sci. ·ussR Zool. Inst.
Translated by Israel Program for Scientific Translations, 1962. 504
pp.
Boughton, R. V., and W. A. Clemens. 1966. The annual production of the
whitefishes Coregonus clupeafonnis and £:... nasus in Teslin Lake,
British Columbia. Northwest Science. 40 (4):147-154.
Cohen, Daniel M. 1954. Age and growth studies on two species of
whitefishes from Point Barrow, Alaska. Stanford Ichthyological
Bulletin 4(3): 168-187.
Craig, P.C., and L. Haldorson, 1980. Beaufort Sea Barrier Island-Lagoon
ecological process studies; final report, Simpson Lagoon studies.
Part 4. Fish. OCS Annual Report. Contract No. 03-6-022-35193, LGL
limited, Sidney, British Columbia. 259 pp.
DeGraaf, D., and K. Machniak, 1977. Fishe~ investigations along the Cross
delta pipeline route in the Mackenzie delta. Chapter IV, pages 1-169
in (P. McCart, ed.). Studies to determine the impact of gas pipeline
development on aquatic ecosystems. Arctic Gas. Biological Report
Series, Vol. 39. Prepared by Aquatic Environments L i1nited.
Furniss, R.A. 1975. Inventory and cataloging of Arctic area waters.
Alaska Department of Fish and Game, Federal Aid in Fish Restoration,
Project F-9-7, Study G-1-I. Vol. 16, 47 pp.
Jones, David R., (1971). An evaluation of the swimming perfonmance of
several fish species from the Mackenzie River. University of British
Columbia, Vancouver. 53 pp.
Kepler, P., 1973. Population studies of northern pike and whitefish in the
Minto Flats complex with emphasis on the Chatanika River. Alaska
Department of Fish and Game. Federal Aid in Fish Restoration. F-9-5,
Study G-Il, Jog G-II-J. Vol. 1~: 59-81.
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Kogl, Dennis R., 1971. Monitoring and evaluation of Arctic waters with
emphasis on the North Slope drainages: Colville River study. Alaska
Department of Fish and Game. Federal Aid in Fish Restoration, Project
F-9-3, Job G-Ill-A, Vol. 12:23-61.
Kogl, Dennis, and Donald Schell. 1975. Colville River Delta fisheries
research. Chapter 10, pages 483-504 in Environmental Studies of an
Arctic Estuarine System -Final Report. Prepared by Institute of
Marine Science, University of Alaska. U.S. Environmental Protection
Agency, Corvallis. EPA-660/3-75-026. 536 pp.
Kuz•min, A.N., 1969. Development of the reproductive system in female
broad whitefish (Coregonus nasus (Pallas)), reared in ponds and lakes
of northwest USSR. Problems of Ichthyology. 9(2): 1g7-205.
Lindsey, C. C., and C.S. Woods (eds.), 1970. Biology of coregonid fishes.
University of Manitoba Press, Winnipeg. 560 pp.
McPhail, J. D., and C. C. Lindsey, 1970. FreshWater fishes of Northwestern
Canada and Alaska. Fisheries Research Board of Canada Bulletin 173,
Ottawa. 3B1 pp.
Morrow, James E., 1980. The freshwater fishes of Alaska. Alaska Northwest
Publishing Co., Anchorage. 248 pp.
Muth, K.M., 1969. Age and growth of broad whitefish, Coregonus nasus, in
the Mackenzie and Coppermine Rivers, N.W.T. J. Fish. Res. Bd. Canada,
26 (8): 2252-2256.
~cott, W. B •• and E. J. Crossman. 1973. Freshwater fishes of Canada.
Fisheries Research Board of Canada Bulletin 184. Ottawa. 966 pp.
Townsend, Alan H., and Phil P. Kepler, 1974. Population studies of
northern pike and whitefish in the Minto Flats complex with emphasis
on the Chatanika River. Alaska Department of Fish and Game. Federal
Aid in Fish Restoration. Project F-9-6, Study G-II, Job G-11-J. Vol. r1
15 : 59-79 .
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FRESHWATER HABITAT
RELATIONSHIPS
ROUND WHITEFISH-PROSOPI UM CYLINORACE UM
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ALASKA DEPARTMENT OF FISH & GAME
HABITAT PROTECTION SECTION
RESOURCE ASSESSMENT BRANCH
APRIL 1981
I
FRESHWATER HABITAT RELATIONSHIPS
ROUND WHITEF'ISH (PROSOPIUM CYLINDRACfUM)
By
Stephen S. Hale
Alaska Department of Fish and Game
Habitat Division
Resource Assessment Branch
570 West 53rd Street
Anchorage, Alaska 99502
May 1981
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ACKNOWLEDGEMENTS
Many people from the Alaska Department of Fish and Game and from the Aul:e
Bay Fisheries Laboratory of the National Marine Fisheries Service freely
gave their time and assistance when contacted about this project and it 1s
a pleasure to thank thetn ar.d fishery biologists fro~~ other agf:ndes,
especially those who provided unpublished data and observ~tions f~ t~!ir
own work. The librarians of the Alaska Resources Library and the U.S. Fish
and Wildlife Service we re of great help.
This project was funded by the U.S. Fish and Wildlife Service, Western
Energy and Land Use Team, Habitat Evaluation Procedure Group, Fort Coll ·ns,
Colorado. Contract No. 14-16-0009-79-119.
TABLE OF CONTENTS
Round Whitefish
I. Introduction
A. Purpose
B. Distribution
c. Life History Summary
D. Ecological and Economic Importance
II. Specific Habitat Requ i remen .;s
A. Spawning
B. Incubation
c. Juvenile Rearing
o. Adults
III. Suitab i lity Index Curves
IV. Deficiencies in Data Base and Recormendati ons
Page
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I. INTRODUCTIO~
A. Purpose
Thh report presents available infonnati on on the freshwater
habitat requirements of the round whitefish, Prosop1um
cylindraceum (Pallas), and evaluates the habitat parameters which
are most important to the species or are most often critical to
survival or limiting to production. The emphasis of this report
1s on habitat requirements, primarily those of a physical or
chemical nature. Certain biological factors affecting the well
being of the population such as feeding, predation, competition,
parasites, and disease are not comprehensively treated.
This report is intended to support habitat evaluation activities
by presenting a data base for the species and by pointing out
where more data are needed. Although information has been gained
from throughout th£ range of the species, emphasis is placed upon
Alaska. While there appears to be wide differences in growth
rates between different populations within the State, there is
insufficient infonnation to show that habitat requirements differ
among the various populations.
The life history of the round whitefish is not well known. It is
one of the least studied coregonids (Jessop and Power, 1973).
Several good papers or. the life histories of related species are
presented in Biology of Coregonid Fishes edited by Lindsey and
Woods (1970). Most studies of round whitefish have dealt with
age and length; little is known of habitat tolerances,
preferences, and requirements. The round whitefish appears to
ha~e fairly wide habitat tolerances. They are widely distributed
in Alaska, except for the Aleutian Islands and Southeast, and
occur in a variety of habitats including lakes and rivers.
B. Distribution
The round tt"hitefish is distributed widely in Siberia and the
no~hern pa~ of No~h America. It is one of the most widespread
and conmon species of no~hern waters (McPha i 1 and lindsey,
1970). In Siberia, it occurs in Arctic Ocean drainages from the
Yenisei River east to the Bering Sea, south to !"'O~hern
Kamchatka, and is also found in drainages on the no~h side of
the Sea of Okhotok. In North America, it occurs in New England,
the Great Lakes (except for Lake Erie), in most of Canada (except
for the southern pa~ of the four western provinces and in the
region of the Manitoba -Ontario boundary where there is a
discontinuity) and in Alaska (McPh~11 and Lindsey, 1970; Scott
and Crossman, 1g7J).
The round whitefish occurs throughout Alaska except for the Yukon
-Kuskokwim delta, Aleutian Islands, Kodiak Island, and most of
the southeast part of the State; although it does occur in the
Chilkat, Alsek, and Taku drainages (R. Baxter, personal
communication; McPhail and Lindsey, 1970; Morrow, 1980} (Figure
1.) It is most abundant in gravelly mountain streams and
associated lakes {Alt, 1971; R. Baxter, personal communication;
Berg, 1~48; Krasikova, 1968; McCa~ et al., 1972).
C. Life History Sunnary
At least some populations of round whitefish apparently engage in
spawning migrations but they are not as strong or directed as
those of some of the other whitefish (Morrow, 1980). Movements
have been observed in tributaries of the Sagavanirktok River in
August and September prior to spawning season (McCa~ et al.,
1g12; Yoshirhara, 1972). In Newfound Lake, New Hampshire, fish
move to the spawning area in the fall; the males usually arrive
before the females (Nonnandeau, 196g). Craig and Wells {1975}
repo~ed aggregations of round whitefish in spring-fed sections
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Fagure 1.
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MAIN STUDY SITES
(I] Colville Atver
[!] Sagavanirktok River
~ Minto Flata
[!] Paxaon Lake, Summn ~ake
[!] Klbuck Mounta1na
COOK INUT • ~
Diatrlbutton of Round wftitettaft in Alaska (A. Baxter. personal
communtcatto. Morrow, 1980) and main study sites
-3-
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of the Chandalar River in the fall, which may indicate the
existence of a spawning migration.
Spawning takes place in the fall in shallow. gravelled areas of
lake shores or streams. Spawning in interior Alaska occurs in
late September through October (Morrow, 1980). It may occur in
northern Yukon Territory as early as late August ~Bryan, 1973,
cited by Bryan and Kato, 1975). November and December is the
spawning season in the southern parts of t..e range (Scott and
Crossman, 1973). Apparently there is little or no feeding during
prespawning or spawning activities (Craig and Wells, 1975;
Normandeau, 1969).
During spawning in Newfound lake, males and females swim in pairs
(Nonnandeau, 1969). Prior to spawning round whitefish on the
spawning grounds in Aishihik Lake, Yukon Territory, slowly swam
near the bottom in groups (Bryan and Kato, 1975). The eggs are
broadcast over the substrate and settle into crevices in the
gravel.
Fecundity ranges from about 2,000 -14,000 eggs/female with large
females producing up to 20,000 eggs; the average is around 5,000
-10,000 eggs (Bailey, 1963; Craig and Wells, 1975; Furniss,
1974; Krasikova, 1968; McPhail and Lindsey, 1970; Nonnandeau,
1969}. No p3rental care is provided for the eggs.
There are indications that in many areas, round whitefish, once
metu·re, spawn every year (Craig and Wells, 1975; Krasikova, 1968;
McCart et al., 1972); however, this is not the case in all areas
(Jessop and Powe,r, 1973).
In Newfound lake, eggs incubate about 140 days (at 2.2°C) and
hatch in April and May (Nonnandeau, 1969). The young hatch as
sac fry and the yolk is absorbed in two or three weeks. The fry
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remain upon the bottom after n .• ':ching and seek shelter in rubble
and boulders. They evidently leave the spawning grounds within a
month. In the northe~ Yukon Territory, fry have been found over
gravel areas of streams soon after spring breakup (Bryan~ 1973,
cited by Bryan and Kato, 1975).
There is 1 ittle infonn~tion on whether the young fish have
rearing areas separate from the adults. A survey of streams in
the Sagavanirktok River basin showed that fry were abundant in
the main river and lower Atigun River, but not in other areas
where juveniles and adults were present (McCart et al., 1972).
Ko9l (1g71) believed the Colville River delta to be an important
rearing area as young :ge groups were captured there. Rearing
also takes place in the summer in the tributaries of the Colville
River (Alt and Kogl, 1g73). Krasikovi (1968) stated that in
Siberia the young feed in the upper reaches of streams and along
the shores of lakes. Bailey (1963) stated that there may be a
segregation by age during the summer in the Isle Royale area of
Lake Superior.
The adult round whitefish is usually found in deep lakes in the
southe~ part of its range and more often in rivers and streams
in the northern parts (Scott and Crossman, 1973). In the Great
Lakes, it is considered to be a shallow water species. Koelz
(1929, cited by Scott and Crossman, 1973) stated that it is found
at water depths less than 45.7 m and Dryer (1966, cited by Scott
and Crossman, 1973) found them to be most abundant in western
Lake Superior at water depths less than 36.6 m. The greatest
water depth at which Dryer caught fish was 71.9 m. Scott and
Crossman (1973) stat£ that a U.S. Fish and Wildlife Service
vessel caught one specimen in a bottom gillnet set in eastern
Lake Superior at 218.9 m. In the Kuskokwim River area, round
whitefish occur in lakes at least as deep as the hypolimnion (R.
Baxter, personal communication).
-s-
The round whitefish has been ~ported to occur in brackish waters
a 1 of'lq the coast11 ne of Hudson Bay and off the mouths of the
Coppennine and Mackenzie rivers (Scott and Crossman, 1973).
Evidently, it does not occur in brackish water in Alaska (Alt,
1971); at least it has not yet been captured in such areas.
The round whitefish 1s a bottom feeder, consuming primarily
benthic invertebrates in shallow areas of streams and inshore
areas of lakes. Food items include: mayfly larvae, caddisfly
larvae and adults, chirono.id larvae and pupae, gastropod and
pelecypod molluscs, fhh eggs and small fish, innature Diptera
including blackfly and mosquito larvae, stonefly nymphs, and
cladocerans (Furniss, 1974; Krasikova, 1968; McCart et al., 1972;
Morrow, 1980; Nonnandeau, 196g; Scott and Crossman, 1973).
Krasikova (1968) found that the feeding rate of round whitefish
in July anc! August is considerably higher in streams than in
lakes. Movement of round whitefish into tributa~ streams of the
Sagavanirktok River apparently are sunner feeding excursions
(McCart et al., 1g72}.
The maximum length, weight, and age ~ported for the species is
56.1 em total length, 2.0 kg (perhaps 2.3 kg) and age 22 (Craig
and Wells, 1g75; Furniss, 1974; Morrow, 1980; Scott and Crossman,
1973). MacKay and Power (1g68) stated that the ultimate size for
the species was 40 -50 em fork length. Round whitefish in
Alaskan waters a~ usually less than 40 em fork length and
usually weigh less than 0.5 kg although specimens up to 52 em and
1.5 kg have been taken ( K. Al t, personal coiiii'IJnication; R.
Baxter, personal communication).
The fish reach sexual maturity at about age 4 or 5 in the
southern part of their range and at age 6, 7 or 8 in northern
parts (McPhan and lindsey, 1g7o; Morrow, 1980; Peck, 1964).
Similar diffe~nces exist between fast and slow growing
populations in Sibe~ia (Kras1kova, 1968).
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Gt"owth rates vary considet"ably tht"oughout the range (Bailey,
1963; Ct"aig and Wells, 1975; Falk and Dahlke, 1974; Jessop and
Power, 1973; Mraz, 1964; Peck, 1964; Whye and Peck, 1968). Lake
Michigan fish reach a total length of 50 em in 7 years; fish from
a lake in the Brooks Range take 12 years to reach this length
(Morrow, 1980). Krasikova {1968) believed that the differences
in gt"owth rates among areas resulted from differences in food
production.
D. Ecological and Economic Importance
The t"'und w~ftefish plays a minor role in lake ecosystems as a
predator' on the eggs of other fish but it primarily consumes
benthic invertebt"ates. The round whitefish is a prey item for
other' species of fish, including lake trout and pike.
The round whitefish is a high quality food fish. In past years,
it has supported a commercial fishery in the Great Lakes up to
about 180 metric tons annually (Scott and Ct"ossman, 1973). It is
of some conmercial impot"tance in the USSR. In Alaska,
subsistence fishermen take limited numbers of round whitefish.
The fish fs sought by sport fishermen fn New Brunswick (Scott and
Ct"'ssman, 1973) and a limited sport fishery exfsts in Alaska
(Alt, 1971).
-7-
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II. SPECIFIC HABITAT REQUIREMENTS
A. Spawning
Round whitefish in Aishihik Lake, Yukon Territory, spawned when
water temperatures were around 1 -2°C (Bryan and Kato, 1975).
Krasikova (1968) found them spawning in the Rybnaya stream of the
USSR in early October when the water temperature was around 0°C.
Lake Superior fish spawned at 4.5°C (Scott and Crossman, 1973}.
Round whitefish spawn in both lakes and streams. At the outlet
of Aishihik Lake, Bryan and Kato {1975) observed them spawning in
current velocities ranging from less than 31 em/sec to 63 em/sec.
A sampling grid in the spawning area revealed that eggs were most
dense in the faster water. However, they also spawn in still
water (Koelz, 1929, cited by Bryan and Kato, 1975; Nonmandeau,
1969).
Round whitefish spawned in the outlet of Aishihik Lake at water
depths ranging from 0.7 to 2.5 m (Bryan and Kato, 1975). Eggs
were most abundant at depths less than 1 m. In Newfound Lake,
New Hampshire, they were obsP:ved spawning at water depths of
0.15 to 2.0 m or more (perhaps down to 3.66 m) although most eggs
were found at depths between 0.15 and 0.60 m (No~ndeau, 1969}.
Spawning in the Great Lakes takes place at water depths of 4 to
15 m (Koelz~ 1929, cited by Scott and Crossman, 1973). The
substrate chosen by round whitefish for spawning in Newfound Lake
was a rocky reef covered with gravel-and rubble and a few large
boulders. The bottom was kept free of silt by wind driven waves.
Sandy areas were not used for spawning (No~ndeau, 1969}. Bryan
and Kato (1975) found that the round whitefish of Aishihik Lake
spawned at the outlet over a variety of substrates ranging from
111.1d to gravel and boulders but that the eggs were most dense over
gravel (particle size 2-250 mm). Other investigators have
reported them spawning over grave 1 and bou 1 ders ( Koe 1 z, 1929,
cited by Bryan and Kato, 1975), over gravel (AOF&G, unpublished
-8-•
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B.
MS ), and over cobble and gravel in widened stretches of small
streams (Krasikova, 1968).
Incubation
The water temperature of Newfound lake was 2.3°C during the
incubation period of round whitefish eggs (Nonnandeau, 1969).
Given that round whitefish in Alaska spawn in the fall at water
temperatures around 0°C, the eggs must incubate at temperatures
close to 0°C. Eggs of round whitefish which spawned in lakes .
would probably incubate at slightly higher temperatures.
No information is available on dissolved oxygen levels required
during incubation but the oxygen levels encountered must be
fairly high, given that the eggs are located on the surface of
substrate and incubate at low water temperatures.
C. Juvenile Rearing
Little infonnation is available on any ha.bitat requirements or
preferences that juveniles may have which are different from
adults . Baxter (personal co~~~nunication) has found that Age I and
II fish live in the same areas as adults but in shallower waters .
Peck (1964) compared the growth rates of round whitefish in Paxon
and Surrmit Lakes and fo,und that the young-of-the-year fish in
Paxson Lake have a greater growth rate. Peck suggested that this
was a result of an earlier warming of the waters in the spring
and a greater food productivity in Paxson Lake.
D. Adults
In the western part of the North Slope, round whitefish are found
in the sunmer in streams where the water temperature ranges from
3 -l6°C (Kogl, 1971). Fall migrations in the Sagavanirktok
River drainage have been observed at water temperatures ranging
-9-
from 0 -13°C (McCart et a 1 q 1972; Yoshi hara, 1972). ~ound
whitefish in Moosehead Lake, Maine, were distributed in August at
water temperatures ranging from 13.9-17,5°C (Cooper and Fuller,
1945, cited by Ferguson, 1958). Round whitefish have been found
overwintering in deep holes in the Colville, Kuparuk, and
Sagavanirktok Rivers at water temperatures of 0 -1°C (Be~dock,
1977, 1979 and 1980).
Round whitefish in streams of the Kilbuck Mountains are generally
found where the gradient is greater than 0.5 m/km {AOF&G,
unpublished MS}. They occur in North Slope streams where the
gradient ranges from about 1 -14 m/km and the current velocity
ranges from about 24-274 cmVsec (kogl, 1971}. They may not
actually be present in the higher current velocities, but do
occur in streams which have these velocities. Berg (1948} and
Krasikova (1968} stated that round whitefish in the USSR prefer
swift currents.
Jones (197?) determined experimentally that the critical velocity
for nine round whitefish with an average fork length of 30.4 em
was 42.5 em/sec.
Round whitefish are found on the North Slope in the summer over
substrates ranging from -..d to cobble anci boulders (Bendock,
personal communication; Kogl, 1971}. However, they seem to be
found most conmonly in streams with a gr•avel bottom (AOF&G,
unpublished MS; Berg, 1948; Krasikova, 1968).
The turbidity of streams of the westem North Slope where round
whitefish are found in the summer ranges from ~lear to 15 ppm
(Kogl, 1971}. They apparently have a preference for clear
streams (R. Baxter, personal communicationi T. Bendock, personal
communication; DeGraaf and Machniak, 1977).
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In the Colville River, healthy appearing fish ~ith food in their
stomachs were taken from ~aters ~ith dissolved oxygen
concentrations ranging from 2.6 • 5.6 ppm (Bendock, 1980). In
the Kuparuk and Sagavanirktok Rivers where round whitefish ~ere
taken, the d1ssol ved oxygen saturation ranged from 49 -!00~
(Bendock, 1977). StreUIS of the ICilbuck Mountains and Nulato
Hills where they are found are well oxygenated throughout the
year (ADF&G, unpublished HS).
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III. SUITABILITY INDEX CURVES
No suitability index curves were drawn for the round whitefish becau;e
it was beli<eved that there is not sufficient data to construct
meaningful curves. The wide distribution of the species in a varie·:y
of hab1tats wan broad ranges of habitat parameters further compoun•ls
the problem of describing opt1llllln h.a.bitat. Tables I and II provide a
general overview of certain parameters.
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Table 1: Round Whitefish Spawning
Parameter Observed Values RP.marks Location Reference
Temperature around 1 - 2
Aishihik lake, Bryan and Kato, 1975 oc Yukon Territory
around 0 Rybnaya River, Kras 1kova, 1968
USSR
4.5 lake Superior Scott and Cross~n.
1973
Current 0 Great lakes koelz, 1929 (cited b~ • Velocity, Bryan and kato , 1975 ....
Cf em/sec
0 Newfound lake, Nonnandeau, 1966
H. H.
<31 -63 eggs most abundant in outlet of Aish1hik Bryan and kato, 1975
faster water lake, Yukon Terr.
Water 0.7 -2.5 eggs most abundant at outlet of Aish1h1k Bryan and kato, 1975
Depth, water depths <I m lake, Yukon Terr.
m
0.15-2.0 + range Newfound lake, , Nomandeau, 1969
N. H.
0.15 -0.60 most eggs
4 -15 range Great lakes Koelz, 1929 (cited by
Scott and Crossman, 1973)
Table II: Round Whitefish, Adult
Parameter Observed Values Remarks location Reference
Temperature, 3 -16 have been captured at these Western North Kogl, 1971 oc temperatures in the summer Slope
0 -13 fall migrations Happy Valley Crk. McCart et al., 1972
(trib. of
Sagavanirktok A.)
0 -12 fall migrations lupine Rtver Yos:hihara, 1972
(trib. of
I Sagavanirktok A.) .....
t 0 - 1
overwintering Colville, Kuparuk Bendock, 1977 & 1980
and Sagavanirktok A.
1J .g-17.5 distribution in August Moosehead lake, Cooper and Fuller,
Maine 1945 (cited by
Ferguson, 1958)
Dissolved 2.6 -5.6 mg/1 overwintering Colvtlle River Bendock, 1980
Oxygen
49 -100% overwintering Kuparuk and Bendock, 1977
saturation Sagavanirktok R.
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IV. DEFICIENCIES IN DATA BASE AND RECOMMENDATIONS
Because of the paucity of information, it is not possible to elaborate
on specifics regarding the habitat tolerances, preferences and needs
of the round whitefish. If the habitat of the round whitefish is to
be adequately described in Alaska, more research 1s required. Studies
needed range from basic descriptive life history studies to extensive
measurements of habitat parameters to physiological exoeriments 1n the
laboratory.
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liTERATURE CITED
Alaska Department of F1sh and Game. (Unpublished manuscript). Whitefish
investigations of the Yukon-K.uskolcwim Delta. Co-rcial Fisheries
Division, Bethel.
Alt, Kenneth T. 1971. Distribution, ~ve.ents, age and growth, and
taxono~~ic status of whitefish (Coregonus sp.) in the Tanana-Yukon
drainage and North Slope. Alaska Dept. of Fish and Game. Federal Aid
in Fish Restoration, Project F-9-3, Study R-11, Job R-II-F. Vol.
12:19-31.
Alt, Kenneth T. and Dennis R. Kogl. 1973. Notes on the whitefish of the
Colville ~iver, Alaska. J. Fish. Res. Bd. Can. 30(4):554-556.
Bailey, Merryll M. 1963. Age, growth, and maturity of round whitefish of
the Apostle Islands and Isle Royale Regions, Lake Superior. Fish.
Bull. U.S. Fish and Wildl. Serv. 63(1):63-75.
Bendock, Terrence N. 1g77. Beaufort Sea estuarine fishery study. Final
Report. OCS Contract No. 03-5-022·69. Alaska Dept. of Fish and Game,
Fa1rbanks.
Bendock, Terrence N. 1979. Inventory and catalog1ng of Arctic area
~ters. Alaska Dept. of F;sh and Galli!. Federal Aid in Fish
Restoration, Study No. 6-1, Job No. 6-I-1. Vol. 20:1-64.
Bendock, Terrence N. 1980. Inventory and cataloging of Arctic area
waters. Alaska Dept. of Fish and Game. Federal Aid in Fish
Restoration, Study No. G-1, Job No. G-I-1.
Berg, Leo S. 1948. Freshwater fishes of the USSR and adjacent countries.
(Transl. from Russian). Vol 1. Acad. Sci. USSR Zool. Inst. Israel
Program for Scientific Translations, 1962. 504 pp.
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Bryan, J. E. and D. A. K?to. 1975. Spawning of lake whitefish 9 Coregonus
clupeaformis, and round whitefish, Prosopium cylindraceum, in Aishihik
Lake and East Aishihik River, Yukon Territory. J. Fish. Res. Bd. Can.
32(2):283-288.
Craig9 P. C. and J. Wells. 1975. Fisheries inveGtigations in the
Chandalar River Region9 Northeast Alaska. Chapter 1, pages 1-114 1!
P. C. Craig (ed.). Fisheries investigations in a coastal region of
the Beaufort Sea. Arctic Gas Biological Report Series. Vol. 34.
Prepared by Aquatic Environments Limited.
DeGraaf, D. and K. Machniat. 1977. Fishery investigations along the cross
d·elta pipeline route in the Mackenzie delta. Chapt. IV. Pages 1-169
-~ P. McCart. (ed.). Studies to determine the impact of gas pipeline
development on aquatic ecosystems. Arctic Gas Biological Report
Series. Vol. 39. Prepared by Aquatic Environments liartted.
Falk, H. R. and l. W. Dahlke. 1974. Data on the late and round whitefish,
lake cisco, northern pike and arctic grayling from Great Bear Lake,
N.W.T., 1971-1973. Canada Dept. of the Environment, Fisheries and
Marine Service. Data Report Series No. CEN/D-74-1.
Ferguson, R. 0. 1958. The preferred temperature of fish and their
midsummer distribution in temperate lakes and streams. J. Fish. Res.
Bd. Can. 15(4):607-624.
Furniss, Richard A. 1974. Inventory and cataloging of Arctic area waters.
Alaska Dept. of Fish and Game. Federal Aid in Fish Restoration,
Project F-9-6, Study No. G-1, Job No. G-I-1. Vol. 15:45 pp.
Jessop, B. H. and G. Power. 1973. Age, growth and maturity of round
whitefish (Prosopium cy11ndraceum) from the Leaf River, Ungava,
Quebec. J. Fish Res. Bd. Can. 30:299-304.
.,
Jones, David R. 197?. An evaluation of the swimming performance of 1
several fish species from the Mackenzie River. Univ. of British
Columbia, Vancouver. 53 pp.
Kogl, Dennis R. 1971. Monitoring and evaluation of Arctic waters with
emphasis on the North Slope drainages: Colville River study. Ala .ska
Dept. of fish and Glme. Federal Aid in Fish Restoration, Project
F-9-3, Job G-Ill-A, Vol. 12:23-61.
Krasikova, V. A. 1968. Materi.als on the blology of the round whitefish,
Co~gonus cyl i ndraceus ( Pa 1 l .u et Pennant), from the Nori 1' sk lake and
River system. Problems of Ichthyology. 8(2):301-303.
lindsey, C. C. and C. S. Woods . (eds.) 1970. Biology of coregonid fishes.
Univ. of Manitoba Press, Winnipeg. 560 pp.
McCart, P., P. Craig and H. Bain. 1972. Report. on fisheries
investigations in the Sagavanirktok River and neighboring drainages.
Report to Alyeska Pipeline Service Company. 83 pp.
MacKay, Isabel and G. Power. 1968. Age and growth of round whitefish
(Prosopium cylindraceum) from Ungava. Trans. Am. Fish Soc.
25(4):657-666.
McPhail, J. D. and C. C. Lindsey. 1970. Freshwater fishes of Northwestern
Canada and Alaska . Fisheries Research Board of Canada FiuHetin 173,
Ottawa . 381 pp.
Morrow, James E. 1980. The freshwater fishes of Alaska. Ala.ska Northwest
Publishing Co., Anchorage. 248 pp .
Mraz, Donald.
Michigan.
19M·. Age and growth of the round whitefish in Lake
Trans. Am. Fish. Soc. 93(1):46-52.
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Normandeau, Donald A. 1969. Life history and ecology of the round
whitefish, Prosopium cylindraceu• (Pallas), of Newfound Lake, Bristol,
New Hampshire. Trans. Am. Fish. Soc. 98(1):7-13.
Peck, James W. 1964. A comparative study of the growth of the round
whitefish, Prosopium cylindracew~, in the Copper River drainage,
Alaska. M.S. Thesis, Univ. of Michigan, Ann Arbor. 53 pp.
Scott, w. B. and E. J. Crossman. 1973. Freshwater fishes of Canada.
Fish. Res. Bd. Canada Bull. 184. Ottawa. 966 pp.
Whye, George L. and James W. Peck. 1968. A li~ological survey of Paxson
and Summit Lakes in interior Alaska. Alaska Dept. of rfsh and Same.
Info~tfonal Leaflet No. 124. 40 pp.
Yoshfhara, Harvey T. 1972. Monitoring and evaluation of Arctic waters
with emphasis on the North Slope drainag~s. Alaska Dept. of Fish and
Game. Federal Aid in Fish Restoration, Project F-9-4, Study No.
G-Ill, Job No. &-III-A. Vol. 13, 49 pp.
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0
FRESHWATER HABITAT
RELATIONSHIPS
ARCTIC GRAYLING-THYMALLUS ARTICUS
ALASKA DEPARTMENT Of ASH & GAME
HABITAT ~OTECTION SECTION
RESOURCE ASSESSMEN:T BRANCH
I A~IL, 1981
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FRESHWATER HABITAT RElATIONSHIPS
ARCTIC GRAYLING (THYHALLUS ARCTICUSJ
By
Steven W. Krueger
Alaska Department of Fish and Game
Habitat 01v1s1on
Resource Assessment Branch
570 West 53rd Avenue
Anc~orage, Alaska 99502
May 1981
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ACKNOWLEDGEMENTS
Many people from the Alaska Department of Ffsh and Game and from the Auke
Bay F·f sheri es Laboratory of the Nat i or. a 1 Marine Fisheries Service free 1 y
gave their time and assistance ~hen contacted about this project and it is
a pleasure to thank them and fishery biologists from other agencies,
especially those ~ho provided unpublished data and observations from their
o~n work. The librarians of the Alaska Resources Library and the U.S. Fish
and Wildlife Service were of great help.
This project was funded by the U.S. Fish and Wildlife Service, Western
Energy and Land Use Team, Hab 1 tat Evaluation Procedure Group, Fort Collins.
Colorado. Contract No. 14~16·0009·79-119.
TABLE OF CONTeNTS
Arctic Grayling
Page
(. Introduction 1
A. Pur;:lose 1
8. Distribution 2
D. Life History Summary 4
E. Economic Importance 12
II. Specific Habita~ Requirements 13
A. Lake Inlets/Outlets 13
1. Upstream Migration 13
2. Spawning 15
3 . Post-Spawning Movements 19
4. Development of Eggs and Alevins 19
5. Sunner Rearing 20
6. Migration to Overwinterfn9 Areas 22
7. Overw i ntering 22
B. Bog Streams 22
1 . Upstream Migration 22
2. Spawning 25
3. Post-Spawning Movements 28
4. Oevelor~nt of Eggs and Alev i ns 30
5. Sunmer · Rearing 31
6. Migration to Overwintering Areas 32
7. Winter rearing 33
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III.
IV.
v.
VI.
Page
c. Mountain St~ams 34
l. Upstream Migration 34
2. Spawning 35
3. Egg and Alevin Development 36
4. Swnaer Rearing 36
5. Migration to Overwintering Areas 40
6. Winter Re1ring 40
D. Spring Streams 42
1. Upstream Migration 42
2. Spawning 43
3. SUJIIIIer Rearing 43
4. Migration to Overwintering A~as 43
5. Winter Rearing 43
Habitat-Arctic Grayling Relationships 44
Deficiencies in Data Base 53
Recommendations and Further Studies 55
literature Cited 58
I. INTRODUCTION
A. Puroose
The purpose of this project is to describe how selected physical
and chemical features of lotic habitat within Alaska influence
the survival and behavior of the various 11fe stages of Arctic
grayling. Thymallus arctfcus (Pallas).
Objectives of this project are:
1) To gather data from published and unpublished sources within
Alaska. Canada and Montana and from conversations with
fishery biologists from the above areas concerning the
relationships between lotic aquatic habitat and Arctic
grayling survival and behavior .
2) To develop an Alaska data base catnposed of narrative and
tables of observed physical parameters to better understand
habitat-Arctic grayling relationships; and
3} To identify areas where data are lacking and to recmnmend
studies to fill gaps 1n the data.
The following "Life 'iistory Sunmary• and "Specific Habitat
Relationships" sections will identify the lotic habitat
relationships of the various life history and seasonal behavior
stages of the Arctic grayling which include:
upstrea~ spawning migration;
spawning;
post-spawning movements;
incubation;
su~~~~~er rearing;
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migration to overwintering areas; and
winter rearing
B. Distribution
The Arctic grayling is a holar"ctic species of the genus
Thymallus. Within North America it occu~ from Hudson Bay west
through northern Manitoba, Saskatchewan, Alberta and British
Columbia, the Northwest Territories (excluding most islands of
the Arctic archipelago), \.he Yukon and most of Alaska. In
Eurasia it is found as far" west as the Kara and Ob Rivers and
south to Northern Mongolia and North korea.
Several isolated, relict populations exist in North America. One
is located in a fraction of its original r"ange in the extreme
headwaters of the Missouri River" drainage fn Montana (Nelson,
1953). Two other" relict populations are found in Canada, one in
southeast British Columbia and the other in southwest Alberta
(Scott and Crossman, 1973).
An Ar"ctic grayling population ir the Great Lakes region vas
eliminated fn the 1930s. Possible causes were log drives during
the spawning season, intense angling effort and general habitat
degradation (Creaser" and Creaser", 1935). Attempts to restock
these waters failed. Arctic grayling have been introduced to
Colorado, Utah, Idaho and Yennont (Scott and Crossman, 1973).
Arct ·lc gr"ayltng are distributed over 111ch of Alaska (Figure 1}
(McClean and Delaney, 1978). Ofstr"fbution of Arctic grayling
within southeast Alaska is primarily li~fted to stocked lakes.
Occasionally fish drift downstream in large stream systems, such
as the Stikine River. These rive~ often support substantial
populations of grayling fn their headwater" reaches (McClean and
Delaney, 1978).
-2-
1 ....
l\
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Insert Figure 1
' •
MA IN STUCY SIT ES
OJ TANANA RIVER
ll) KAVIK RIVER
[l) GUUCANA RIVER-
(1] TYEE LAICE
(3] COlVII.LE RIVER
AL!UTIAN ISlANDS • • ~ • Ah 4f?.:. ~tll.;li'
FIGURE l. OlSTRIBUT lON OF ARCTIC GRAYLING IN ALASKA (FROM
ALASKA DEPARTMENT OF FISH AND GAME, 1978)
AND MAIN STUDY SITES .
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Numerous clearwater tributa~ies and lakes within the upDer Coppe~
and Susitna River drainages contain Arctic grayling. G~ayling a~e
found in the Gullcana and Oshetna Rivers. Arctic grayling are
extremely limited within the Prince WilliAm Sound area and lower
Copper Rfver drainage and somewhat lfmi ted within the lower
Susftna River drainage. Both of these large rivers are glacial
and support relatively small populations. However, a few
cl eanfater tributaries of. the 1 ower Copper River and 11111ny
cleanfater trtbutar1es of the lower Susitna River, such as the
Talachulitna and Chunflna River. and Lake Creek contain Arctic
grayling. Arctic grayling are not found on the west side of Cook
Inlet south of Tyonek (McClean and Delaney, 1978).
Nume~us lakes within the Kenai Peninsula were stocked and now
suoport reproducing populations of Arctic grayling. These
include Crescent. Upper and Lower Paradise, Bench and Twin Lakes
(McClean and Delaney, 1978). Selected lakes on Kodiak and
Afognak Islands contain Arctic grayling.
Grayling are also found in clear water stream of the Alaska
Peninsula and Bristol Bay drainages. Especially large
individuals are found in the Ugashik and Becha~f Lake and Togiak
River drainages (McClean and Delaney, 1978).
Arctic grayling are widely distributed in the remaining Arctic
and sub-Arctic areas of Alaska, with the exception of the
Yukon-Kuskokwim River delta area (McClean and Delaney. 1978).
( . l f fe Hf story SUIIIIIIry
Arctic grayling usually migrate to spawning sites just prior to
or during spring breakup. Several factors influence the upstream
migration of this fish including distance separating
ovenfintering and spawning sites, streamflow and water
tenperatu res. Tack (1971) reported upstream 110vement of fish
-4-
with f n the i ce cove red Chen a River about Z • 5 weeks prior to
breakup condftfons. Tripp and McCart (1974) reported upstream
movement of fish in the ice covered Mackenzie River to ice-free
tributaries.
Fish passage can be prevented by ice jams, beaver dams or
waterfalls. The swfnnfng perfonnances of adult and juvenile
Arctic grayling are influenced by fish size, water temperature,
current velocity and the size and extent of barriers. Sex and
spawning condition also influence the migration of adult fish
(MacPhee and Watts, 1976).
The importance of the spawning migration to juvenile fish is not
clear. Tack (1980) related this phenomenon to homing. Juvenile
fish may become imprinted on visual, olfactory or other
conditions and recognize the spawning area upon maturation.
The establis•nt of 1111le spawning territories IMY initiate
spawning activity (Kruse, 1g59; Bishop, 1971; Tact, 1971). Males
select territorial sites far various physical conditions where
spawning will eventually tate place.
Grayling territories vary fn size with respect to stream width,
water depth, current veloc1 ty, channel configuration, spawner
densfty and, possibly, other conditions. Kruse (1959) measured
territories of 0.15 by 0.61 m (0.5 by 2.0 ft) in a 1.5 m (5 ft)
wide reach of Northwest Creek, an inlet of Grebe lake, Wyoming.
However, 1n a wider (2.4-3.0 m, 8-10 ft) reach of the same
stream, territory dimensions were approxfJMtely 1.2 by 1.2 m {4
by 4 ft). Tact {1g71) noted that male fish territories in the
outlet of Mineral Lake, Alaska were about 2.4 by 2.4 m (8 by 8
ft).
Ripe males establish spawning territories and vigorously defend
them from other males. Short head-on thrusts usually repel
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sub-adults. Adult males are repelled by lateral displays.
sometimes foll~ed by direct attacks (Tack, 1971}.
Grayling spawn in areas with surface current velocities less than
1.4 Ill/sec (4.5 ft/sec}, varying vater depths and relatively
small, un1mbedded gravels about 2.5 em (1 in) in diameter.
Fertilization occurs after the females leave their holding areas
and pass through male territories. The females are pursued by
.everal males who attempt to court her. The successfu1 male
places his dorsal fin over her back, initiating simultaneous body
arching and vibrating. The male may drive the posterior third of
the female's body fnto the substrate where eggs and sperm are
released.
The fertilized eggs sink to the bottom of the stream and adhere
to the substrate. No actual redd is constructed but the eggs may
be covered by as much as 5 em (2 in.) of dislodged substrate.
There 1 s no parent a 1 care of the eggs. After spawning , the
female resumes her former resting position before possibly
spawning again. Both sexes may spawn more than once with various
partners. They are capable of spawning annually (Brown, 1938;
Kruse, 1959; Bishop, 1971; Tack, 1971; Scott and Crossman, 1973).
It is not kn~n whether juvenile and adult Arctic grayling return
to the same spawning stream.
The duration of Arctic grayling spawning activity may range from
four days to two weeks (Warner, 1955; Tack, 1971; McCart, Craig
and Bain, 1972; Tripp and Mccart, 1974}.
Age of 1111turity fs variable and generally is greater in the
northern reaches of thfs fish's range. Grayling have been found
to reach sexual maturity at age 2 or 3 in Michigan {Creaser and
Creaser, 1935) and 1n Montana (Kruse, 1959) streams. Most fish
in the North Sl~pe (of the Brooks Range, Alaska) and northern
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Yukon and the Northwest Territories mature between ages four to
nine (Bishop, 1971; McCart, Craig and Bain, 1972; deBruyn and
McCart, 1974;. Craig and Poulin, 1974). Fish in the lower
Kuskowkwim River, Seward Peninsula and Tanana River, Alaska reach
maturity at ages five to six (Alt, 1966 and 1978; Wojcik, 1955).
The life span of Arctic grayling is variable; northern
populations generally live longer than southern populations.
Maxii!Rim ages of Arctic grayling from Montana range from seven to
eleven years (Brown, 1938; Nelson, 1953). Several fish from
selected Beaufort Sea drainages in the Yukon Territories and the
Chandalar River drainage in north-central Alaska ranged in age
from 15 to 22 years (de Bruyn and McCart, 1974). Maximum ages of
fish from various Tanana Rfver, Alaska drainages are about 11
years (Tack, 1973). Regional differences in grayling life spans
may result from varying environmental conditions over their range
(Craig and Poulin, 1974) or from differences in aging techniques
(scale versus otolith).
Female fecundity varies with fish size and stock. Brown {1938)
reported egg fecundities ranging from 1,650 among Grebe Lake,
Wyoming individuals (length and weight unknown) to over 1Z,900
eggs among large (0.91 kg (2 lb)) females from Georgetown Lake,
Montana . Mean fecundity va 1 ues among fema 1 es from Grebe Lake
varied from 1,900 eggs 1n individuals less than 280 nm (11 in)
fork length (fl) to 2,800 in fish exceeding 305 nm (12 in) fl.
Female f1sh ranging 1n fork length (fl) from 331 to 373 1m1
(x • 353 nn) from Weir Cr-eek, a small tributary of the Kav '1k
River, Alaska contained from 4,580 to 14, 730 eggs (x • 8,482
eggs) (Craig and Poulfn, 1974}. Ten females (295 to 395 mm fl)
from the upper Chandalar R1ver, Alaska drainage contained 2,330
to 9,150 eggs (x • 4,937 eggs) (:raig and Wells, 1974).
Deve 1 opment. of A ret 1 c gray 1 i ng eggs to hatching occurs very
rapidly {13-32 days) a .nd is influenced primarily by water
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temperatures (Henshall, 1907; Ward, 1951; Bishop, 1971; Kratt and
Smith, 1971; Tryon, 197~). The hatched fry, or alevins, with
attached yolk sac, are about eight mm long {Scott and Cross~n,
1973). The yolk sac is c~mplet~ly absorbed in one to two weeks.
Kruse (1959) examined survival of emergent grayling in Grebe
Lake, Wyoming. He estimated that fis~ survi~al through the fry
stage was about six percent in one of several inlet streams.
Prabab 1 e causes of th 1s high mort a 1f ty were egg d 1s 1 adglll!n t ,
predation and low fertilization. Water vel'lcity could also
influence egg and alevfn survhal. No redds were const"'cted
during spawning and fertilized eggs may nat have been covered
with gravel.
Alevins hatching within the gravels probably have higher survival
than those hatching an the exposed substrate (Kratt and Smith,
1977). The gravel provided cover, decreased the chances of
dislodgement and lessened swimming stresses in the early stages.
Growth of Arctic grayling varies considerably aver its range, but
fish from northern regions generally grow more slowly than fish
frt'nl southern areas (C.-afg and McCart, 1974b). The largest
grayling recorded from Alaska weighed 2.13 kg {4 lb, 11 oz), and
was 54.6 em (21.5 in) long {Ugashik Narrows, Alaska Peninsula,
1975). The Canadian record grayling weighed 2.71 kg (5 lb, 7 oz)
and measured 53.3 em (21 in) ( Northwest Territories, 1967).
Growth rates of young of the year (yoy) Arctic grayling can be
extremely variable among drainages due to differences in length
of open water (growing) seasons, temperatures and food supplies.
For example, 49 fish within the outlet of Chick Lake (along the
Donnelly River, a tributary of the Mackenzie River, Northwest
Territories, Canada} attained a mean fo.-k length (fl) of 49 mm ~
4 mm by 8 July, 1973. In contrast 38 individuals from a small
inlet to Chick Lake were only 20 mm ~ 4 mm by July (Tripp and
-8-
McCart, 1974). The Chick Lake inlet had lower water temperatures
and lower benthic invertebrate standing crops than those of the
Chick Lake outlet. Elliott (1980) noted substantial differences
in growth rates amng yay Arc:ti c grayling amng sma 11 bog and
mountain streams within Alaska. He ascribed those differences to
food availabfl ity, water temperatures and durations of the
growing seasons.
Arctic grayling are opportunistic feeders and consume more and
larger prey as they grow. Young of the year fish have been
observed feeding prior to total yolk sac absorption (Brown and
Buck, 1939; Kruse, 1959). Fish inhabiting lakes may consume
Oaphn i a and chi ronomi d 1 a rv ae and pupae. Elliott ( 1980 )
investigated the sumner food habits of fish in selected spring,
rapid-runoff and bog streams cro~sed by the Trans-Alaska Pipeline
System (TAPS). Early yay fish (less than 3.5 mm fl) consumed
about three different aquatic and terrestrial invertebrate taxa
whereas larger yoy fish (equal to or greater than 3.5 11111 fl)
consumed up to eight taxa. Immature chironomids were the most
frequently eaten taxon.
Larger fish consume dri ftfng flllllilture and mature aquatic
invertebrates, mature terrestrial invertebrates and occasionally
leaches, fishes, fish eggs, shrews and lemmings (Rawson, 1950;
Kruse, 1959; Bishop, 1971; Scott and Crossman, 1973}. Mature
fish apparently feed infrequently or not at all during the
upstream spawning migration.
Arctic grayling may feed during the winter. Fish, captured by
gill net under the ice wf th i n poo 1s of the Saga va n1 rkto k and
Colville Rivers, Alaska, contained ephemeropteran and plecopteran
nymphs (Alt and Furniss, 1976; Bendock, 1980}.
Predation on Arctic grayling eggs and alevins by other fishes
could significantly reduce fish production. Tack (1971) reported
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whitefish preying upon Arctic grayl fng eggs at the outlet of
Mineral Lake, Alaska. Rainbow trout (Salmo gairdneri
Richardson}, Arctic char {Salvelinus alpinus (Linneaus}}, round
whitefish (Prosopfum cylindraceum (Pallus)}, northern pike (Esox
lucius Lfnnaeus}, longnose suckers (Catostomus catostomus
{Forster}). and other fishes 11111y also consume Arctic grayling
eggs and alevfns (Bishop, 1971; MacPhee and Watts, 1976; Alt,
1977).
Spawned-out adult fish may remain within spawning areas or
migrate considerable distances to sunmer feeding areas within
lakes or streams. A spawned-out fish tagged in late June 1972 ;n
a lake outlet entering the Mackenzie River near Nonnan Wells,
Northwest Territories was recovered within a month in the Great
Bear River, 159 km (99 mi) distant (Jessop et al., 1974). Tagged
adults have been shown to leave Poplar Grove Creek, Alaska, a
smell bog stream, within several weeks after spawning and move to
other areas for feeding (MacPhee and Watts, 1976; W11lfam5 and
Morgan, 1974).
Movement of juvenile fish out of spawning streams can ocr.ur
during or slightly after adult fish emigration. Decreased flows
and lower food availability influence both adult and juvenile
fish movements. Some juvenile fish may remain near spawning
areas through the summer.
Studies examining the summer microhabitat selection by juvenile
salmonids in various Pacific Northwest streaiiS indicate that
larger individuals progressively move to faster and deeper stream
reaches for increased cover and food availability (Everest and
Chapman, 1972; Lister and Genae, 1970). Most of the la;"Qer
juveniles were found in relatively fast water with some cover and
areas of low current velocity. Everest a.ld Chapman (1972)
speculated that fish hold in areas of low current velocities and
feed in areas of faster velocity with higher prey densities.
-10-
Fry (yoy) may remain ~ithin their natal streams or migrate to
other systems ~here they feed and g-row during the relatively
short Arctic and sub·Arctic open water season (McCart and Bain,
1972; MacPhee and Watts, 1976). The movement of' laf"(jer, older
fish out of spawning streams may lessen competition among age
classes. Rearing fish are segregated by size (age) ~ith yoy fish
generally occupying areas of lower current velocities. and more
shallow water. Yearling and older fish generally occupy deeper.
slightly faster areas (Chislett and Stuart, 1979). Larger fish
have been observed fn pools upstream of smaller fish; areas which
probably contain higher densities of prey (Vascotto, 1971).
Fish appear to return to the same su111ner rearing areas. Many
tagged individuals have been recovered the following year in the
same areas (Tack, 1980).
Limited studies monitoring ffsh movements fn sNll bog and
mountain streams have detected a late sun~ner to early fall out
migration of juvenile and yoy fish (McCart, Craig and Bain. 1972;
MacPhee and Watts, 1972). Downstream movement of juvenile fish
generally occurs slightly before migration of yoy fish (Craig,
McCart and Bai n, 1972). Arctic grayling IIIJSt migrate to their
overwintering grou .nds before the streams become impassable from
low flows or ice buildup. Decreasing water temperatures and
flows associated ~ith the onset of winter probably influence the
timing of migration to overwintering areas .
The winter dfst.rfbutfon of Arctic grayling fs more restricted
than the s~r distribution. Most bog and many s~ll mountain
and lake inlet and outlet streaas become dewatered or freeze
solid during the fall and winter months. Fish over.inter in
lakes. open pools, spring and glacial streams and in spring fed
mountain streams. Fish have been found in pools of la"9e
interior Alaska mountain streams, such as the Chena River.
Spring streams in the Tanana River drainage fn interior Alaska
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witn seemingly suitable conditions for overwintering fish, do not
appear to support overwintering of Arctic grayling (Reed, 1964;
Pearse, 1964; Van Hyning, 1978). Spring fed streams along the
north slope of the Brooks Range, Alaska, are often the only areas
with flowing water and arP. important fish overwintering areas
(Craig and Poulin, 1974).
0. Economic Importance
Arctic grayling are the basis of an important summer recreational
fishery. The broad food habits of this fish allow anglers to use
a variety of t!ehniques, including fly casting.
Roadside angling is popular during the summer on streams and
lakes along the Alaska, Steese, Elliot, Taylor, Glenn, Parks,
Richardson and Nome-Taylor highways. Fly-in and float fishing
trips are also popular during the summer.
-12-
II. SPECIFIC HABITAT REQUIREMENTS.
A. Lake Inlet/Outlets
1. Upstream Migration
a. Water T!!!erature
Water temperatures associated with the upstream
migration of Arctic grayling to spawning areas withir
inlets and outlets of takes may t~nge from 0°C (32°F:
to about 4°C (39.2°F}. Warner (1~5) stated that fisn
began entering a selected inlet of Fielding Lake as
soon as there was flowing water. Water t~eratures ,,f
the inlet during the initial phase of the migration
were not given but water temperatures during the las:
t.o days of the migration were 0.6°C and 1.1°C (33°F
and 35°F).
"'any arctic streams may be impassable to Arctic
gray 11 ng prior to spring breakup because of ice
conditions or dewatering. Fish have been observed
moving upstream through narrow furrows in anchor ice
created by meltwater (Wojcik, 1955). However, maximum
numbers of fish usually migrate within these streams at
or near peak flow conditions (MacPhee and Watts, 1976;
Tack, 1980). Ripe Arctic grayling have been detected
moving upriver within fee-covered spawning streams such
as Trail Creek, a tributary of the Mackenzie River,
Northwest Territories, Canada (Jessop, Chang-Kue,
Lilley and Percy, 1974). Arctic grayling nay ascend
rapid runoff and bog streams. and inlets and outlets of
1 a kes as soon as f1 ow conditions permi t pas sage to
spawning sites.
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Tack {1972} reported Arctic grayling at the mouth of
the Mineral Lake outlet approximately nine days after
water temperature iJf the same outlet reached 1 °C
(33.8°F). The first large catch of fish at the same
location occurred three days later, probably a fun~tion
of the length of time requ fred to move from their
ove~interfng habitat.
KMUse (1gsg} noted that Arctic grayling fn Grebe Lake,
Wyoming, begin spawning migrations into four inlet
strea~ and the outlet stream (the Gibbon Riveri vhen
water temperatures range from 5.6° to 7.8°C (42°-46°F)
and 2.2° to 4.6°C {36°-40°F), respectively. Average
daily water temperatures at ~he conclusion of the
spawning migrations in the inlet and outlet streams
were 7.2° to 8.3°C (45°-47°F) and ~.2° to 8.9°C
(36°-48°F). These temperatures are noticeably higher
than those associated with spawning migrations in
Fielding and Mineral Lakes.
Tack (1980) found no correlation between momentary
temperature reductions below 1.0°C and upstream
movement of fish fn the Chena River. Dfel water
temperature patterns may 1 nfl uence upstream fish
movement in other strea~s (Wojcik, 1g54, Warner, 1g55;
MacPhee and Watts, 1976).
b. Current Velocity
Arctic grayling usually begin spawning migrations to
inlets and outlets of lakes during breakup conditions.
Wojcik ( 1g54) ff rst observed fish in one of severa 1
inlet streams of Fielding Lake, Alaska as it began
flowing in mid-May, 1954. Warner (1955) observed the
first fish in the mouth of the same stream the
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Arctic grayling spawned in four inlet tributaries of
Tyee Lake, near Ketchikan, Alaska during May and June
of 1980 in water temperatures ranging from 6° to 11°C
(42.8°·51.8°F) (Cuccarease, Floyd, Kelly and LaBelle,
1980). Water temperatures of the streaRIS during
initial fish spawning were not reported.
Fish spawning occurred in four inlet streams of Grebe
Lake, Wyoming in water temperatures ranging from 4.4°C
to 10°C (40°·50°F). Water temperatures in the outlet
stream during fish spawning ranged from 2.2 °C to 10°C
(36°F-socr) (Kruse, 1959).
Brown (1938) observed several fish spawning in a small,
beaver dammed inlet tributary to Agnes Lake, Montana in
water temperatures of 10°C (50°F).
b. Current Velocity
Arctic grayling spawn in a wide range of current
velocities in inlets and outlets of lakes. Wojcik
(1954) reported fis:1 spawning in "slow, shallow
backwaters, and not in riffles as had been supposed•• in
an inlet stream to Fielding Lake. The following spring
the fish spawned in surface current velocities of about
1.2 m/sec (3.9 ft/sec) (Warner, 1955). Observations
were 1 imited by ice and snow cover during 1954 and
1955.
Surface current velocities in territories of 22 males
along the outlet to Mineral Lake, Alaska ranged from
0.34 m/sec to 1.46 m/sec (1.1 ft/sec-4.8 ft/sec) with a
mean value of 0.79 m/sec. (2.6 ft/sec) (Tack, 1971).
-16-
c. Substrate
Arctic grayling have been reported spawning over gravel
substrates of inlets and outlets of lakes within Alaska
and Montana. Warner {1955) obse.rved fish spawning over
fine (about 1 em) gravel. Much of the stream was
covered by ice and snow, therefore observations were
made a long an 0.18 km (200 yd) open reach near the
mouth of the stream and fn smaller open areas upstream.
Tack (1971) described the spawning substr·ate in the
outlet of Mineral Lake as being ~pea size .~
Fish spawning has been observed within riffles and runs
of four inlet tributaries to Tyee Lake near Ketchikan,
Alaska. Spawning substrate ranged from sand to small
cobble. Coarse sand and gravel to about 2.5 em (1 in)
in diameter was co11100nly used by most ffsh (Cuccarease,
Floyd, Kelly and LaBelle, 1980).
Arctic grayling were observed spawning over a
sand-gravel subs t rate in an inlet stream to Agnes Lake,
Montana by Brown (1938). He discussed the 1 imited
variety of substrate and other habitat conditions
within the stream and the need to better determine the
characteristic.s of optimum Arctic grayling spawning
habitat in Montana streams.
Kruse (1959) ranked sand ( .3 em), gravel ( .3-7 em) and
rubble (7 .6-30.5 em) in descending order as suitable
substrate material for Arctic grayling spawning.
Riffles were utilized more often than pools for
spawning.
Fish were reported spawning over relatively fine
gravel, not exceeding 3.8 em (1.5 in) in diameter with
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d.
most material not exceeding 1.25 em (0.5 in) in
diameter, within the outlet of Bench Lake, Alaska
{personal communications, Stephen Hamma~trom, 1981).
Similar size substrate is used for spawning in the
outlet of Crescent Lake, Alaska (personal
communication, Ted McHenry, 1981).
Water Depth
Arctic grayling spawn in a range of Wlter depths.
Selection of spawning sites is more strongly influenced
by current velocities and substrate conditions. Fish
in an inlet to Fielding Lake spawned in Nshallow back
waters 11 (Wojcik, 1954} and in depths of 16 em (6 in)
{Warner, 1955).
Water depths measured in 22 fish territories fn the
Mineral Lake, Alaska outlet stream ranged from 0.18 to
0.73 m (0.6 to 2.4 ft) with a mean value of 0.30 m (1.0
ft.).
Cuccarese, Floyd, Kelly and LaBelle (1980) observed
fish spawning in various inlet streams to Tyee Lake,
Alaska in water depths ranging from 0.15 to 0. 91 m
(0.5-3.0 ft.) in the largest and rnost intensively
utilized stream and from 0.05 to 0.46 m (0.17-1.5 ft)
fn several smalier, shallower streams with
substantially fewer spawners.
e. Light
Grayling spawning occurs during daylight hours and
probably stops during the evening (Scott and Crossman,
1973). Few observations of Arctic grayling spawning
have been made during the evening (K~se, 1959).
Netting at the Bench Lake outlet found few Arctic
grayling spawning during evening and early morning
hours (personal comt~.mication, Stephen Ha11111arstrom,
1981).
3. Post-Spawning Movements
Spawned-out Arctic grayling (fish having completed spawning)
c0111110nly vacate spawning sites within lake inlets and
outlets and return to lakes or to other areas (Wojcik, 1954;
Warner, 1955; Tack, 1980). Fish spawning within the outlet
to Mineral Lake leave the stream upon completion of spawning
and migrate downstream and then up to the Little Tok River
or Trail Creek. Food availability probably influences
post-spawning fish movement and distribution (Tack, 1980).
Small lake inlets may become dewatered by mid to late
summer. Adult spawned-out fish typically leave these
intermittent streams after spawning and enter lak~s (Kruse,
1959).
4. Development of Eggs and Alevins
a. Water Temperature
Strean1 water temperatures influence Arctic grayling
development rates within lake inlets and outlets.
Kruse (1959) observed that eggs hatched 19 days after
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fertilizati " in an inlet stream to Grebe Lake at water
temperatures from 3.9° to 9.2°C (39.0°·48.5°F).
Fertilized eggs from an inlet to Fielding Lake, Alaska
re~ .red 18 days to hatch in water temperatures ranging
from 6.1° to 9.4°C (43°-49°F) during the spring of 1954
and 1955. Fertilized eggs required only eight days to
hatch in water temperatures of 15. 5°C (60°F) at an
Alaskan hatchery.
Henshall (1907) recommended minimum water temperatures
of 5.5°C (42°F) for successful development of Arctic
grayling in Montana hatcheries.
Water temperatures characteristically rise during the
incubation period; therefore, eggs are not usually
exposed to freezing. However, no upper or lower lethal
temperature data for Arctic grayling eggs were found in
the literature.
5. Summer Rearing
a. Current Velocity
Low flows during incubation could result in desiccation
or freezing of developing eggs and alevins. Wojcik
(1954) noted significant diel flow fluctuations along
an inlet of Fielding Lake, Alaska and discussed the
possibility of recently fertilized eggs becoming
exposed, desiccated or frozen.
Downstream migration of yoy fish within inlets of lakes
is probably a response to more suitable current
velocities and an abundance of food items in lakes.
Newly emerged yoy fish held positions in .. quiet water
coves and eddies" during the day along an inlet to
-20-
G~be Lake. Wyoming (Kruse. 1959). At night the yoy
fish vacated a~as of low cur~nt velocities and
actively migrated downstream to Grebe Lake.
Fry have also been found in shallow margins of Tyee
Lake. Alaska and in small, shallow, pools in the delta
a~a of inlet st~am of Tyee Lake. High aquatic
invertebrate production in littoral areas provided
ample food (Cucca~se, Floyd, Kelly and LaBelle, 1980).
Fry have not been observed in mainstem reaches of the
inlet streams.
Arctic grayling fry within lake outlets typically
occupy areas of low current velocities. Yoy fish have
been observed in stream margins with shallow depths and
low current velocities (personal communication, Stephen
Hammerston and Ted McHenry, 1981).
b. Water Depth
Water depths occupied by yoy fish in lotic and lentic
areas may vary considerably. Depths are probably
selected for the associated current velocities and food
availability. Fry within several inl2ts to Grebe Lake,
Wycxning occupied shallow, slow habitats prior to
migrating downst~am to Grebe Lake (Kruse. 1959). Fry
within Tyee Lake, Alaska occupy shallow littoral
~aches ranging in depth from 2 to 46 em (1-18 in).
They also occupy shallow, q•.iiet pools in delta ~gions
of the inlet streams rather than the mainstem reaches
(Cuccarease, Floyd, Kelly and LaBelle, 1980). Yoy fish
have been observed in shallow ~rgins of the outlets of
Bench Lake and Crescent Lake, Alaska (personal
communication. Stephen Hammarston and Ted McHenry,
1981).
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5. Migration to Overwintering Areas
No infonnation was found in tne 1 iterature concerning
movements of Arctic grayling witnin lake inlets and outlets
to overwintering areas. Fisn probably overwinter in lakes
that are relatively deep and do not freeze to the bottom.
7. Overwintering
No information was available in the literature concerning
overwintering nabitat of fish witnin 1 akes. Tne winter
ecology of Arctic grayling within lakes is poorly understood
(personal communication, Fred Williams, 1981).
B. Bog Streams
1. Upstream Migration
a. Water Temperature
Adult grayling usually migrate upstream before
juveniles. Water temperatures are lower at tnis time.
Upstream migration of yearling, older juvenne and
adult grayling witnin Poplar Grove Creek usually
commenced wnen mean water temperatures ranged from 2°
to 4°C (36°·39°F) during early to mid-May of 1973, 1974
and 1975. Upstream movement usually ceased when water
temperatures approacned 12° to 14°C (54-57°F) during
late Ma.v to early June. Mean water temperatures during
peak upstream migration of yearling fish (6°-12°C) were
consistently nigner tnan temperatures during adult
upstream migration (3°-7°C). Oiel variations in water
temperatures never exceeded 2°C during May and June.
-22-
Mature 'green' (non-ripe) Arctic grayling entered Weir
Creek, a tributary of the Kavik River, Alask.a when
water temperatures were about 5°C. The migration
ceased when water temperatures approached 12°C (Craig
and Poulin, 1974).
Upstream migration of adults in Weir Creek was similar:
mature ffsh were first observed in water of about 4°C
(39°F}. Migration ceased when temperatures reached
l5°C (60°F) (Craig and P'oulin, 1974). Maximum water
temperatures at the tennination of the juvenile
upstream fish migration reached zooc (67°F).
b. Current Velocity and Discharge
The upstream migration. of juvenile and a~ul t Arctic
grayling in bog streams usually coincides with high
flows resulting from snow melt and surface run-off
during spring breakup. The first mature 'green' fish
were taken nine days after breakup in Weir Creek.
Alaska during early June (Craig and Poulin, 1974). The
time between initiation of flow in Weir Creek a.nd
a ·rrival of fish was probably due to the distance from
overwintering areas to the creek (probably the
Shaviovik River, about 85 km distant). Wojcik (1954)
captured mature 'green' fish at the mouth of Shaw Creek
and Little Salcha Creek ;n the Tanana. River, Alaska in
early April 1953 a.nd 1954 while the streams were frozen
and impassable to fish. However, as melt water scoured
furrows in the ice, the fi.sl'l bega.n migrating upstream.
MacPhee and Watts (1976) trapped adult and juvenile
fish in Poplar Grove Creek, a tributary of the Gulkana
River, at peak and decreasing stream flows associated
with spring breakup.
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The upstream fish migration may span several weeks .
Streamflow can be substantially reduced by the time the
upstream migration of adult and juvenile fish is
completed. For example, flows wit;1in Poplar Grove
Creek, Alaska decreased during the upstream migration
of adult and juvenile Arctic grayling during 1973, 1974
and 1975 (MacPhee and Watts, 1976). Adult and juve.nile
fish generally began moving upstream in mid-May durinf'}
the initial stages of the ope.n-water season. The
relatively high discharge at this time ranged from
about 1.3 to 4.0 m3/sec {46-141 cfs). However, the
peak of the juvenne fish migration consistently
occurred at lowe·r flows, about five to ten days after
the peak of the adult fish mig ·ration. Juvenile
migration continued for several days after the adult
migration. Discharge at the end of t .he adult and
juvenile migrations were generally less than 1.1 m3tsec
(38 cfs). The yearling fish lagged several days behind
the older juvenile fish .
Upstream migration of ju11enile and adult Arctic
grayling in Weir Creek, a tributary of the Kavik River,
Alaska, resembled migrations in Poplar Grove Cree.k
(McCart, Craig and Bain, 1974). Adult fish migrated
upstream in Weir Creek in 1971 between earl.v and late
June, 1971 as discharge decreased from 20 m3 /sec to
about 2 m3/sec. Juvenile fish moved upstream from mid
to 1 ate June, about two weeks after the peak of the
adult fish run.
Juvenile an.d adult Arctic grayling migrated upstream in
Nota Creek, a tributary of the Mackenzie River,
Northwest Territories, Canada, during spring breakup in
May. During this time dishcarges decreased from
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1.67 m3/sec to 0.38 m3tsec (58 to 13 cfs) (personal
communication, Derrick Tripp, 1981).
Current velocities may influence the timing of juvenile
and adult fish passage in bog streams. MacPhee and
Watts (1976) demonstrated that large Arctic grayling
could negotiate faster water than smaller fish.
Decreasing flows may enhance the abn i ty of juveni 1 e
fish to pass upstream and could be responsible for ti.e
lag between adult and juvenile fish. Other factors,
such as increasing water temperatures~ probably
influence the timing of the upstream migration of
juvenile and adult fish (MacPhee and Watts, 1976).
2. Spawning
The influences of current velocity, water depth and
substrate on fish spawning in bog streams are not well
documented. Flood stage flows and yellow or brown stained
water limit observations.
Spawning data carrel a ted to water temperature and flow
conditions are available. Selected studies using weirs and
seines noted the spawning condition of fish ('green', 'ripe'
or 'spawned out'}~ water temperature, actual or relative
flow conditions and direction of fish movement.
a. Water Temperature
Water temperatures of bog streams can be considerably
higher than those of lake inlets and outlets during
spawning. Minimum water temperatures may approach 4°C,
the water temperature which apparently triggers
spawning in lake inlets and outlets (Tack~ 1980), and
may approach or exceed 10°C (50.0°F). Arctic grayling
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have spawned at the outlet of ~·ea Lake, Alaska in wate ..
temperatures of 7°C (44°F) (McCart, Craig and Bain,
1972). The Tea Lake system drains a flat, marshv arec
and is a bog stream.
Bishop (1971) reported that water temperatures of about
8° to 10°C (47°·50°F) appeared to stimulate fish
spawning in Providence Creek, Northwest icrritories.
Maximum water temparatures in Weir Creek during the
Arctic grayling spawning period ranged from 4° to 16cc
(39°-61°F) (Craig and Poulin, 1974). Maximum water
temperatures in Nota Creek whfm Arctic gray1ing wen
"ripe" ranged from 3,5° to ll°C (38°-5l°F}. MaximurT
water temperatures at the peak of spawning ranged fr >m
4.5" to uoc (40"-52°F) {personal conmunication,
Derrick Tripp, 1981).
Water temperatures in Happy Valley Creek during Arct c
grayling spawning ranged from 4° to l2°C (39°-53°F)
{McCart, Craig and Bain, 1972).
b. Current Velocity and Discharge
Observations of Arctic grayling spawning with respect
to current velocities are limited. Several pairs of
spawning fish were observed in shallow riffles along
Mainline Spring Creek, Alaska (personal communication,
George Elliott, 1980).
Flows within bog streams are typically high but usuall)
decrease during the Arctic grayling spawning period.
Flows in Happy Valley Creek, Alaska decreased
substantially over the 10 day spawning period. Arctic
grayling spawned in Weir Creek, Alaska for 10 days as
-26-
flows decreased. Arctic grayi i ng in Poplar Grove
!:reek, Alaska spawned in late .May to early June as
streamflows decreased from peaks of about 1.1 m3/sec
(23.2 cfs) in 1973 and 4.0 m3/sec {84.8 cfs) in 1971
(MacPhee and Watts, 1976). Bishop {1971) reported fish
spawning in Providence Creek, tributary of the
Mackenzie River, during breakup conditions.
c. Substrate
Arctic grayling appear to use a wide range of substrate
sizes in bog streams fJr spawning. They spawned over
gravel ranging from 2.5 to 3.75 em (1-li in) diameter
in Mainline Springs Creek near Atigun Pass, Alaska
(personal communication, George Elliott, 1980).
Spawning has also occurred in the outlet of Tea Lake,
Alaska, near the Trans-Alaska Pipeline, over sand and
fine gravel substrate. about 0.6 em (i in.) in diameter
(Craig, McCart and Bain, 1972).
Substr~te used for spawning in Providence Cn!ek.
Northwest Territories, Canada was gravel mixed with
sand. Fish did not spawn over pure mud, sand or clay
(Bishop, 1971}.
Fish spawned in organic detritus in Mi 11 ion Do 11 ar
Creek, Alaska, along the Trans-Alaska Pipeline
(persona 1 cormrun i cation, George Elliott, 1980). The
substrate in Million Dollar Cn!ek ~s silt and fine sand
overlain by organic muck (Elliot, 1980).
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d. Water Depth
e.
Observations of water depths used far spawning in bog
streams are limited.
Several pairs of fish spawned in riffles 5 em (2 in)
deep in Mainline Spring Creek, Alaska (personal
communication, George Elliott, 1980).
Fish apparently spawn only during the day. as observed
by Bishop (1971) in Providence Creek.
3. Post-Spawning Movements
Arctic grayling may migrate downstream il'llllediately after
spawning. In 1973, post-spawners left Weir Creek, Alaska
within two weeks after spawning. Large juvenile Arctic
grayling also emigrated within two weeks. Tagged adult fish
from Weir Creek, Alaska were captured later in the Kavik
River and the Shaviovik River (Craig and Poulin, 1974).
Rapidly decreasing streamflaws probably influenced fish
movements.
Tagged adult post-spawners from Happy VallPy Creek displayed
similar downstream movement following spawning. No adult
fish were found upstream of the weir (McCart, Craig and
Bain, 1972).
An outmigration of adult and juvenile Arctic grayling
occurred in Poplar Grove Creek during late May and early
June after spawning and when streamflows were steadily
declining. Emigration of spawned-out adults extended from
-28-
mid·May through mid-June 1973 as flows steadily declined
from 1.4 m3tsec (49 cfs) to 0.3 m3tsec (11 cfs). Large
juvenile Arctic grayling outmigrated after the adults during
mid-~une. Nat all adult and large juvenile Arctic grayling
left Poplar Grove Creek; of the 1,085 adults and 1,973 large
juvenile fish found migrating upstream, only 779 and 937
were detected passing downstream. Many of these fish
migrate to the Gulkana River drainage (Williams and ~organ.
1974; Williams, 1975 and 1976). Weir data suggest that
adults may remain in Poplar Grave Creek until the stream
freezes in fillll.
Adult grayling ususally leave Nota Creek, a bog stream
entering the Mackenzie River, Northwest Territories, Canada,
within two weeks after spawning. Same spawned-out adults
may return to food-rich Nata Lake for short periods of time.
Emigration of large five and six year old juveniles was
followed by younger, smaller individuals through early July.
By mid·July young of the year Arctic grayling and some
yearling and two year olds occupied Nota Creek (persondl
communication, Derrick Tripp, 1981). Decreased living space
and food availability associated with low flaws are probably
important factors influencing fish movements.
Tack (198() suggested that the outmigratian of juvenile and
spawned-out adult fish may allow yay fish to rear and feed
in natal streams without competition. Adult and juven:le
fish may rear in other stream systems that are rich in food,
such as spring streams.
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4. Development of Eggs and Alevins
a. Water Temper~
Extremely limited fnformation is available concerning
egg incubation and wate~ temperature relationships in
bog streams. Bishop (1971) subjected fertilized eggs
to a range of water tempe~atures. He determined that
eggs hatched in approximately 14 days at a mean water
temperature of 8.8°C {48°F).
A~ctic grayling eggs required 18 to 21 days to hatch in
Nota Creek in water tempe~atures ranging from 5.5° to
13°C (42°-55.5°F) and a mean wate~ temperature of 9.6°
to 10.3°C (49°-50.5°F) (personal communication~ Derrick
Tripp, 1981).
b. Cu~rent Velocity and Discharge
Spates could dislodge and destroy eggs and severely
reduced flows could lead to desiccation.
Aquatic habitat selected by rearing yay fish in bog
st~eams may have current velocities of from 0 to 0.15
m/sec:. Larger fish generally select faster water.
Elliott (1980) measured mean column velocities at
~aldfng positions of •early• yay (~ 35 11111 fl} and
'late' jOY (> 35 mm fl) fish in selected bog stre~s
during June and Augu~t 1980. The mean column current
velocities associated with 'early' yay fish were 0.02,
0.07 and 0.03 m/sec in Million Dollar Creek (n ~ 198},
Pamplin's Potholes (n = 175), a~d the Tea Lake inlet,
Alaska (n =57), respectively.
-30-
Larger 'late' yoy fish were found in slightly faster
mean current velocities: 0.08, 0.09~ 0.14 and 0.1
m/sec. in Pamplin's Potholes {n = 87), Tea Lake
inlet/outlet (n = 71), North Fork Fish Creek (n = 33),
Mainline Spring Creek, Alaska (n = 18), respectively.
5. Summer Rearing
a. Current Velocity
Newly emerged yay fry select protected stream areas
where current velocities are extremely lo~ {personal
conmunication, George Elliott, 1980; de Bruyn and
McCart, 1974; McCart, Craig and Bafn, 1972). Typical
emergent fry rearing areas include shallow backwaters
and flooded stream margins and side channels.
Juvenile fish (age 1 and older, measuring 50-250 mm f1)
have been observed in bog streams with slightly greater
current velocities than yoy ffsh. The average mean
current velocities occupied by juvenile fish fn the Tea
Lake inlet (n ~ 9) were 0.175 m/sec and in Main1ine
Springs Creek, Alaska (n = 16), 0.196 m/sec. Limited
observations of adult Arctic grayling (250 mm fl) from
bog streams showed adult fish holding in mean current
velocities of 0.262 m/sec (Elliot, 1980).
b. Substrate
Rearing fish of all ages were associated with a variety
of substrates fnclusing detritus, silt, sand, and
grave 1s. Arctic gray 1 i ng showed 1 itt 1 e movement in
small bog streams during July and August following fish
spawning and movement to sumer rearing areas and
before movement of fish to overwintering areas (McCart,
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Craig anaBain, 1972; Craig and Paulin, 1974; MacPhee
and Watts, 1976).
c. ',.Ja ter Depth
Water depths occupied by rearing fish vary
considerably. Newly emerged yay fish have been found
in extremely shallow, slow water, flood channels and
backwater sloughs (Personal communication, George
Elliott; de Bruyn and McCart, 1974; McCart, Craig and
Bain, 1972).
Early yay fish occupied small bog streams with depths
from about 0.09 to 0.85 m (0.3-2.8 ft). Late yay
occupied depths (within the same streams) ranging from
0.15 to 1.07 m (0.5-3.8 ft) (Elliott, 1980). Juvenile
and adult fish in bog streams along the Trans-Alaska
Pipeline System were found in water depths from 0.21 to
1.07 m (0.7-3.8 ft).
6. Migration to Overwintering Areas
Significant downstream movement of fish has been observed in
bog streams during late summer, apparently in response to
declining water temperatures and flows associated wi:h the
onset of winter. Emigration of yay and juvenile fish may
also occur during the summer.
-32-
a. Water iemperatur~
Decreasing water temperatures may influence the
downstream movement of Arctic grayling. Significant
numbers of yoy and juvenile fish moved downstream in
Weir Creek~ Alaska during September 1973. Minimum
wdter temperatures during early, mid and late September
were about 1°, 4° and 0°C, respectively. Downstream
movement of juvenile fish occurred about one week
before yoy fish in Weir Creek. No apparent
relationship could be demonstrated betwe~n downstream
movement of juvenile or yoy fish and water temperatures
(Craig and Poulin, 1974).
Similar downstream movements of yoy fish occurred in
Poplar Grove Creek, Alaska. Of the 65,536 yay fish
observed be tween Ju 1 y 17 and October 18, 1973, 96%
( 62,680} were observed in the 1 ower reaches during
October (MacPhee and Watts, 1976). In Poplar Grove
Creek, downstream migration of yoy fish may be related
to stream temperatures.
b. Current Velocity and Discharge
No relationship could be fcund between the downstream
movement of juvenile or yoy fish and stream flows in
Weir and Poplar Grove Creeks.
7. Winter Rearing
Winter rearing areas for Arctic grayling are limited in bog
streams because they often become dewatered or freeze solid
during the w1nter. Winter rearing areas such as deep lakes,
deep pools of mountain streams or spring fed streams, may be
quite distant from summer rearing areas. Fish overwintering
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areas in the Shaviovik River and su11111er rearing habitat
within Weir Creek are about 85 km (53 mi.) distant (Craig
and Poulin, 1974).
C. Mountain Streams
1. Upstream Migration
a. Water Temperature
Low water temperatures are prevalent during the
upstream mi gra ti on of adult and juven11 e A ret i c
grayling. Upst~eam migrants were taken in Vermillion
Creek, Northwest Territory. Canada about one week after
breakup when water temperature~ ranged from 0° to 3°C
(32°-37°F} (personal corrmunfcation, Derrick Tripp,
1981). Tack (1980) also found that water temperatures
of at least 1.0°C (34°F) stimulate upstream movement of
Arctic grayling in large mountain streams like the
Chena River near Fairbanks, Alaska.
b. Current Velocity anci Discharge
Upstream migration of adult and juvenile Arctic
grayling usually occurs during high flows at spring
breakup. A weir placed in Vermillion Creek captured
upst'!"ea.n migrating adult and juvenile Arctic grayling
for ten days after peak flows fn May of 1973 and 1975.
The M~ckenz1e River was covered with ice for up to ten
days a~ter breakup occurred in Vermillion Creek during
1973 and 1975 (personal communication. Derrick Tripp,
1981), and observations of fish migration were not
made.
-34-
Observations made under the ice along the Chena River
indicate that Arctic grayling initiate upstream
movement prior to breakup (Tack., 1980). The fish were
probably moving to upstream reaches of tne Chena River
or its tributaries. The relative importance of
streamflow and water temperature in relation to
upstream fish migration is poorly understood.
2. Spawning
a. Water Temperature
Limited data is available concerning the relationship
between Arctic grayling spawning and stream water
t~peratures. Spawning in the Chena River drainage has
been observed in water temperatures of 5°C (Reed, 1964;
personal communication, Jerome Hallberg). Tack {1g8Q)
discussed the possibility of tne 4°C isotherm
influencing the distribution of spawning fish in large
s:reams 1 ike the Chena Ri'ler.
b. Current V~locity ard Discharge
Fish spawn in mountain streams in a wide range of
current velocities. They have been observed ~pawning
in an overflow slough in tne Cnena River, Alask.a at
relatively low current velocities {Reed, lg64) and in
riffles of the East Fork Chena River, Alask.a where
surface current velocities approach 1.4 m/sec (4.5
ftlsec) {pel"'ional conmunication, Jerome Hallberg,
1~91). Bendock. (197g) reported spawning in pools of
the Colville River, Alaska with negligible current.
Nelson (lg54) observed fisn spawning activity along Red
Rock Creek , Montana in the ends of riffles. Fish
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3.
c.
spawning occur~ed in simila~ low flow a~eas along the
East Fork Chena River, Alaska (personal communication,
Jerome Hallberg, 1981).
Substrate
Arctic grayling use a variety of substrates for
spawning including mud, silt and g~avel up to 4 em (1.5
in) in diameter. Bendock (1979) observed fish spawning
on silt overlaying gravel in the mainstem Colville
River, Alaska and its tributaries. Spawning subst~ate
used by Arctic grayling in the E3st Fork Chena River,
Alaska consists of fine gravels from 0.75 to 4 em
(0.4~1.5 in) in diameter (personal communication,
Jerome Hallberg, 1981). Spawning has also been
observed in muddy sloughs along the Chena River (Reed,
1964).
Arctic grayling in Red Rock Creek, Montana spawned in
gravel-rubble substrate of unknown size but not in pure
silt or sand substrates.
Egg and Alevin Development
No information was found in the literature which discussed
egg and alevin development in mountain streams.
4. Summer Rearing
a. Water Temperature
Results of st~ndard, 96 hour bioassays (at te~t wa~er
temperatures of 5, 10, 15, 20 and 24.5 or 21.5°C)
indicate that Arctic grayling can tolerate a wide range
of temperatures. Fish from the Chena River near
Fairbanks, Alaska were used for this study. Results
were exoressed as median tol era nee 1 imit (TLM), the
temperature at which 50 percent of the individuals in a
test die (LaPerrier and Carlson, 1973).
Resu 1 ts i ntli cated that young of the year fish are
apparently more tolerant of relatively high water
te· .peratures than older fish. The TLM of yoy fish
exc~eds 24.5°C, the highest test water temperature.
Individuals of 10 em fl had TLM values of 20.0 to
24.0°C and fish of 20 em fl has TLM values of 22.5 to
24.s~c. The small fish were acclimated at 4°C and the
larger fish at 8°C.
Bioassay results indicate that water temperature
differences of 2°C at relatively high water
~emperatures can cause very dffferent survival rates of
Arctic grayling. For example, survival of 20 em fl
fish was 100 percent at 22.5°C (72.5°F) and 0 percent
at 24.5°C (75°F).
These bioassay results may not apply to actual stream
conditions because fish could avoid warm water
temperatures by moving to cooler areas.
b. Current Velocity and Discharge
Recently emerged yoy fry generally occupy areas with
low current velocities. The sma'l newly emerged fry
(about 20 ITin tot a 1 1 ength at 14 cays) have 1 i mi ~ed
swimming abilities. Chislett and Stuart (1979) noted
that newly emerged fry clustered in shallow, protected
reaches of flood channels, backwater sloughs and
sidechannel pools of the Sekunka River, British
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Columbia.
the E,sst
These fish are found in similar habitats in
Fork Chena River, Alaska (personal
communication, Je~ome Hallberg, 1981).
Nelson (1954) noted that recently emerged yoy fish were
dist~ibuted in "back.waters and protected areas ••. ,
away from strong currents" within Red Rock Creek,
Montana.
01 der yoy fi 5h co:-cupy progressively faster waters.
'Early' yoy fish (~ 35 mm fl) occupied a mean current
velocity of about 0.07 m/sec (0.22 ft/sec) (n = 183) in
selected headwater areas of the Gulkana River, Alaska
in early July. Larger yoy fish (> 35 mm fl) inhabited
slightly greater current velocities, 0.16 m/sec {0.52
ft/sec) (n = 157} (Elliot, 1980).
Chislett and Stuart (1979) found yoy fish (~ 35 mm fl)
occupying slow current a~eas of backwater and side
channels in the Sekunka River and Martin Creek, B~itish
Columbia during July 1978. Side channels contained
flowing water and were less ephemeral than backwater
channels. All yoy fish were found in low current
velocities.
Most of the backwater hauitats dewatered during August
1 ow flows and yoy fish (:> 35 mm fl ) inhabited
s idechanne 1 riffle a~eas and mo.rgi ns of mains tem
riffles.
By September and October yoy fish occupied sidechannel
riffles and margins of mainstem riffles where current
velocities approached 0.8 m/s~:. Yay fish at this time
ranged from 50 to 96 mm fl (Chlslett and Stuart, 1979).
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Summer distributions of yearling and older fish were
limited to mainstem and side channel pools in the
Sekunka River, British Columbia. Older fish, age 4 to
8+ (oldest aged fish), occupied larger, deeper pools
than the younger fish.
The distribution of juvenile and adult Arctic grayl ~119
in selected Alaskan mountain streams is similar to the
the distribution in the Sekunka River, British Columtia
where aduJt and juvenile fish are generally restrict~d
to pools and sloughs (Alt. 1978; personal
conmunication, Jerome Hallberg, Joe Webb, Terence
Ben dock, 1981}.
Fish will mo'e tnto shallower, faster riffle areas for
more food, such as s~lmon anc whitefish eggs (Bnndo(K,
1979; Alt, 1978).
c. Water Depth
Yoy fish generally occupy shallow lotic habitats with
low current velocities. Fry in the Sekunka River,
British Columbia selected shallow areas in sidechannels
and backchannels (CPiislett and Stuart, 1979). Yoy fi~;h
have been observed in backwater sloughs and sha~low
pockets of protected water in the East Fork Chena
River, Alaska (personal communication, Steven Grabacki,
Jerome Ha 11 berg and Sandra Sonn i ch sen, 1981) .
Older fish generally select deeper pools (Chislett et
al.; personal conmunications, Steven Grabacki, Jerome
Hallberg and Sandra Sonnichsen, lg81}.
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d. Cover
Recent 1 y emerged yoy fish seek various fonns of
instream cover when disturbed. Young (17-45 days old)
fry in shallow, siltbottomed back.channels of the
Sekunka River moved into deeper water with various
types of instream cover. Similar aged yay fish in
sidechannels used substraie interstices and shadows of
boulders for cover. Nelson {:954) noted that 14 to 21
day old fish in Red Rock. Creek., Montana made little
movement when disturbed and appeared to be "relatively
helpless."
Older fish commonly use logs, boulders and turbulence
for instream cover (personal communication, Jerome
Hallberg, 1981 ).
5. Migration to Overwintering Areas
6.
Little is known about Arctic grayling migration to
overwintering areas. Tack (1980) observed a slow downstream
movement of Arctic grayling in the Chena River, Alask.a and
compared it to the faster emigration of fish in North Slope
mountain streams where winter conditions occur very early
{Yoshihara, 1972). Yoshihara {1972) observed many fish
moving downstream in the Lupine River inmediately aft~r
water temperatures reached freezing. Age distribution of • emigrants in the Lupine River is not known.
Winter Rearing
The distribution of overwintering Arctic grayling is more
limited than the summer distribution. Streamflows are low,
much or all of the stream is ice-covered and stream reaches
-40-
can be frozen solid during the harsh Arctic and sub-Arctic
winters. Ovenr.rintering areas in mountain streams include
pools of intermittent or flowing streams (such as Colville
and Chena River, Alaska, respectively) or spring fed streams
which remain open during winter months (the lower Shaviovik
River, Alaska).
a. Current Velocity
Current velocities in overwintering sites are probably
very low. Conventional current velocity meters do not
function at air temperatures below freezing. Fish
overwinter in ·intermittent pools of the Colville River
where current velocity is negligible. In the Hulahula
River oveY"Jo~intering sites had current velocities of
0.15 m/sec (0.5 ft;sec).
b. Water Depth
Fish have been observed under the ice in pools of at
ll!ast 1.4 m {4.6 ft.) depth in the Colville, Chena and
East F~rk Chena Rivers. Bendock (lgao) found fish in
intermittent pools deeper than 1.5 m (4.8 ft) in a
reach of the Colville River, Alaska. Fish are
restricted to pools in the East Fork Chena River and
the mainstem Chana River, Alaska (Tack, 1980) during
the winter months (Hallberg, personal communication,
1981).
Arctic grayling have been taken by Kaktovik, Alaska
villagers in the Hulahula P.iver in late April (Furniss,
1975) and through the ice in the East Fork Chandalar
River near Arctic Village, Alaska (McCart, 1974).
Maximum water depths were about 0.6 m (2 ft) in open
water of the Hulahula River. Water depth of the East
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For( Chandala~ River was 1.5 m (5 ft). These sites are
thought to be important overwintering areas for Arctic
grayling.
Spring fed mountain streams are often tne only sites fn
tne North Slope whe~e water remains flowing throughout
the winter. Tnese streams are important overwintering
sites for Arctic grayling. Alt and Fumiss (1976)
captured adult ~·:":tic grayling in a spring fed pool in
the Franklin Bluff's area of the Sagavanirktok River,
Alaska on May 6, 1975. The app~ximate depth of' the
pool was about 1.2 m (3.9 ft).
c. Dissolved Oxygen
Bendock (1980) measured dissolved oxygen levels ranging
from 0.6 to 4.5 mg/1 in Arctic grayling overwintering
sites in the Colville Rive~. Dissolved oxygen levels
were about 4.8 mg/1 in the Sagavanirktok River at the
Franklins Bluff site on April 10, 1975 (Alt and
Fum iss, 1975).
D. Spring Streams
1. Upstream Migration
lfmfted investigations indicate that Arctic grayling may
enter springfed streams after spawning (Reed. 1964; Pearse,
1974; Tack, 1980).
~42-
Arctic grayling apparently do n~t spawn in springfed streams
where 1ow water temperatures may adversely influence egg and
alevin development (VanHyning, 1978).
3. Summer Rearing
Arctic grayling rear in springfed streams. Reed (1964)
reported that adult fish enter the Delta Clearwater River in
early June and younger juvenile fish enter fn late July.
Pearse (1974) found similar trends in the Delta Clearwater
in 1973; although adults tended to remain in the headwater
reaches and immatures remained downstream.
4. Migration to Overwintering Areas
Reed (1964) stated that irnnature, catchable (by rod and
reel) Arctic grayling moved downstream early in 1963 fn the
Delta Clearwater River. Larger adult fish remained in the
river through most of September. Some tagged adult fish
were found at the mouth of the Delta Clearwater River fn
late October. I
5. Winter Rearing I
Arctic grayling apparently do not rear in interior Alaska I
springfed streams {Van Hynfng, 1978), but have been found in
spri ngfed streams fn the North 51 ope (Craig and Poulin,
1974).
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III. HABITAT-ARCTIC GRAYLING QELATIONSHIPS
Tables r through VI summarize the reported water temperature levels
associated with various life stages and activities of Arctic grayling.
Table VII 1 ists the reported current velocities (or discharges)
associated with different life history stages and Tab·le VIII, the
substrate types used for spawning.
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Table 1: Observed water temperatures associated wtth upstream migration of
Arctic grayling to lake inlets/outlets.
Parameter
Water
Temperature
Observed Values
5.6 -7,8°C
Remarks
Ma•fmum water temperatures
during last 2 days of fish
migration
Ftrst mature fish captured
during fish migration
Water temperatures of
several inlets to Grebe Lk.
during tnittal fish spawning
migration a'ttvfty
)gSJ. )g54
Water temperatures of outlet
(Gibbon A.) to Grebe lake.
Wyoming during tnfttal fish
spawning mtgrattton activity
location Reference
Inlet to Fteldfng Warner (lg55)
lake, Alaska
Outlet to Mineral Tack (1972)
lake. Ale. ;.ka
Four inlets to Kruse (lg5g)
Grebe lake,
Wyoming
Gibbon River.
outlet to Grebe
Lake. Wyoming
Kruse ( 195g}
-------------------
Table II: Observed water temperatures associated with upstream migration of
Arctic grayling within bog (tundra) streams.
Parameter Observed Values Remarks Location Reference
Water sa -l2°C Maximum water temperatures Weir Creek, Craig ancf Poulin
Temperature during early to late stage Tributary to (1974)
of adult fish spawning migra-Kav1k R •• Alaska
tton. Incomplete ffsh sampling
due to high flows. 1973.
J.go -15.6°C Maximum water temperatures Happy Valley Crk, McCart, Craig and
beginning and end of adult tributary t!l Bain (1~72)
fish upstream migration, 1g11. Sagavanerktok
Weir placed tn stream after River, Alaska
fish migration began.
7.2° -19.4°C Maximum water temperatures Happy Valley Crk, McCart, Craig and
I from start to near tennin~-Sagavanerlctok Ba1n (1972) ....
(J'I tton of juvenile fish River, Alaska I
upstream migration, 1971.
2° -4°C Intttal average water tempera-Poplar Grove ~cPhee and Watts
tures of Poplar Grove Creek Crk, tributary (1976)
during start of adult, sub-to Gu 1 kana R. ,
adult, ju~entle fish upstream Alaska
migration durfng May }g73-1975.
12° -l4°C Average water temperatures Poplar Grove MacPhee and Watts
at end of adult, sub-adult, Crk, tributary (1976)
juvenile fish migration tn to Gulkana R.,
June 1973-1975. Ahska
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Table Ill: Observed water temperatures associated ~ith
Arctic grayling spa~ning in mountain streams.
Parameter Observed Values Remarks
Water 5.6°C Instantaneous water tempera-
Temperature ture of slough where fish
~ere observed spawning
Instantaneous water tempera-
ture during fish spawning
act1v1ty
lnst~ntaneou~ water tempera-
ture during fish spa~ning
activity
Instantaneous water tempera-
ture during fish spa~ning
activity
Instantaneous ~ater tempera-
ture during ftsh spawning
act 1vity
Location Reference
Slough along Reed ( 1964)
Chena River I
Ahska
Riffle I E. Fork Ha 1l:J,erg
Chena River, (personal
Alaska co11mun i cation}
Seabee 1 Rainy, BE.ndock
F ass il Crks, (1970)
tributaries
Colville R., Alaska
Nuka Rtver, tr1b-Bendock
utary I Colville (1979)
River, Alaska
Aniak R, trib-Alt (1977)
utary, Kuskokwim
River, Alaska
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Table IV: Observed water temperatures associated with
Arctic grayling spawning in lake inlets/outlets.
Parameter
Water
Temperature
Observed Values
3.3"C
2o -Joe
6° -11°C
4° -tooc
2.2 .. -10°C
Remarks
Initial fish spawning
activity occurred
Fish spawning activity was
completed at this temperature
3-4 days after commencement
of spawning activity.
First spawning activity
observed
Ftsh spawning ce~sed as
water temperature dropped
below 4"C
Fish spawning 1cttvtty
resumed and lasted 4
additional days
Fish actively spawning
fish actively spawning
fish activ~ly spawning
fish actively spawning
-----~--
location Reference
Inlet to fielding Wojcik (1954)
Lake, Alaska
Inlet to fielding Wojrik (1954)
lake, Aldska
Outlet to Mineral Tack {1972)
lake, Alaska
Outlet to Mineral Tack (197?.)
Lake, Alaska
Outlet to Mineral
lake, Alaska
Severa 1 in 1 ets
to T.vea Lake ,
Alaska
Four inlets to
Grebe lk, Wyoming
Outlet to Grebe
Lake, Wyoming
Gibbon River
Inlet to Agnes
lake. Montana
Tack (1977}
Cuccarease, Floyd
Ke 11 y , l aBe 11 e
(1980)
Kruse (1959)
Kruse (1959)
Brown (1938)
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Table V: Observed water te.peratures associated with
spawning of Arctic grayling within bog streams.
Parameter
Water
TeiiiPerature
Observed Values Remarks
Direct spawning observation,
one temperature reading
ftsh spawning activity seemed
to be related to these water
temperatures.
Maximum water temperatures
based on occurre"ce of rtpe
and spawned·out Arctic gray.
ling captured by we1r
Maximum water temperatures
during Arctic grar11ng
spawning acttvfty based on
condition of ftsh in weir
Location
Outlet Tyee lit,
A las lea
Reference
McCart, Craig and
Ba1n (1972)
Providence Crlt, Bishop (1971)
tributary to
Mackenzie River
Northwest Territory
Wetr Creek, Craig and Poulin
Alaska (1g74)
Nota Creek,
Alaska
Trtpp (perc;onal
cOIIIIIJntcation.
}g81)
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Table VI: Observed water temperaturPs associated with
Arctic grayling egg and alevin development.
Parameter
Water
Temperature
Observed Values
6.)0
-9,4°C
i = 7.7°C
Remarks
Eggs hatched in 19 days
Eggs hatched in 18 days
in 1954 and 1955
Eggs at hatchery facility
hatched in 8 days
location
Inlet to Grebe
lake. Wvomtng
Inlet to Fielding
lake. Alaska
Somewhere fn
Alaska
RP.ference
Kruse (1959)
Wo.ic1k ( 1954)i
Warner (195 5 )
Woj c i k ( 1954 )
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Table VII: Relationship of current velocity (or discharge)
to specific life history stages.
------------------------------------------__ L_ --
Activfty Current Rate or Flow Reference
-----------------------------------------------------------------
Spawning
Early Development
Juvenile Rearing
Adult Summer Habitat
Adult Winter Habitat
Upstream Nigration
slo~. shallow backwater
1.2 m/s
.34 m/s to 1.46 (i A,79)
sha11~w riffles
1.1 misec
4.0 m /sec
low flow and riffles of 1.4 m/s
neglig1b le
.02, .07, .03 m/s
shallow, protected areas
shallow pools
shallow pools
.08 to .195 m/s
.8 m/s
,26 m/s
open areas
at breakug
1. 33 - 4 m /s 3 2 m ts 3 ~ 20 m /s
1.67 m /s
at breakup
at breakup
high flow
Wojcik, 1954
Warner, 1955
Tack, 1971
E 11 i ott. 1980
MacPhee and Watts, 1976
MacPhee and Watts, 1976
Hallberg, 1981
Bendock, 1979
Elliott, 1980
Chtslett and Stuart, 1979
Cuccarese et al., 1980
Hanmarston, 1981
Elliott, 1980
Chi5lett and Stuart, 1979
[ 111 0 t t • 1980
Chislett and Stuart, 1979
MacPhee and Watts, 1976
MacPhee and Watts, 1976
Mctart et al., 1974
Tr1pp, 1981
Woj ci k , 1954
Warner, 1955
Tack, 1972
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Table VIII: Reported substrate types used for spawning.
Substrate
Fine gravel (1 em)
"pea-size"
sand to small cobble
sand -gravel
san~. gravel, rubble
fine gravel ( <3.8 em)
fine gravel
gravel, 2.5 -3.75 em
sand, fine gravel
sand, gravel
organic detritus
sand, muck
mud, silt, grav~l( ~ 4 em)
gravel, .75 - 4 em
Reference
Warner, 1955
Tack., 1971
Cuccarease et al., 1980
Brown, 1938
Kruse, 1959
Hammarstron, 1981
McHenry, 1981
Elliott, 1980
Craig et al., 1972
Bishop, 1971
Elliott, 1980
Elliott, 1980
Bendock, 1979
Ha 11 berg, 1981
11. )EFICIE NCIES I ~ DATA 9ASE
Factors influencing the migration of adult and juvenile Arctic
grayling from overwintering areas to spawning streams is apparently
influenced by flow and water temperature conditions. The timing of
adult and juvenile fish migrations are not understood; the juvenile
fish run lags sevP.ral days to several weeks behind the adult fish in
certain areas.
Information about selection of si t!s in mountain, lake inlet and
outlet, bog and spring streams in relation to current velocity. water
depth and substrate is limited. Most of the observations were made in
lake inlets and outlets and mountain streams.
Habitat selection by spawning Arctic grayling is influenced by at
least three variables -substrate, water depth !nd current velocity -
which collec~ively determine the habitat quality. For example, Arctic
grayling may be excluded from spawning areas by excessive current
velocitiPs despite acceptable substrates and water depths. There is
limited information on the interaction of various physical parameter~.
Factors influencing the survival and development of eggs and alevins
are not well understood . Studies indicate that egg dislodgement by
other spawning fish and spates may be a major cause of morta 1 i ty.
Minimum water t~mperatures required for successful egg development are
not known; however, water temperatures above 6,C {42~F) have been
reco11111ended.
The movement of Arctic grayling to overwintering ar~as appears to be
influenced by flow and water temperatures associated with the onset of
winter. This e~~~igration ntay be of short duration or extend over
several weeks. There is very little information about Arctic grayling
overwintering areas. Fish overwintering in the North Slope ue
limited to open water areas or to streams which do not completely
freeze. Fish overwintering in interior Alaska and Canada may remain
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in springfed and glacier streams. It is not known whether young of
the year Arctic grayling burrow into interstices within cobble and
rubble as water temperatures approach freezing.
The age structure of Arctic grayling populations during the open-water
season may be significantly different among various lake inlet and
outlet. bog. mountain and spring streams. Some streams appear to
function as nursery areas for young of the year and older juvenile
fish, and other streams may support only large juvenile and adult
fish. Fish may be either sedentary or nomadic during the open-water
rearing season. Explanations for fish emigration are speculat~ve but
living space and food availability are probably influential. Studies
of juvenile and adult Arctic grayling habitats focused on water depth~
current velocities and substrate in small bog and mountain streams.
Methods of describing water depth, current ve 1 ocity and substrate
characteristics varied among studie~. Some fnvest'gators measured
current velocity at the site of spawning Arctic grayling; others
estimated surface current velocities. Substrate sfze classification
systems also varied; few researchers evaluated substrate imbeddedness
at spawning sites.
The effects of water temperatures and current velocities on a
grayling's ability to ascend culverts has been studied for juvenile
and adult fish. Relatively few tests were conducted with yearling
grayling.
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V. RECOMMENDATIONS AND FURTHER STUDIES
In depth investigations are needed to detcnnine the relationships
between specific chemical and physical features of aquatic habitats
and grayling growth and behavior. For example, investigations should
be designed and conducted to assess the factors which influence
development and survival of Arctic grayling eggs and alevins in bog,
lake inlet and outlet, spring and mountain st.reams. Investigations
should consider egg dislodgement, predation, desiccation, and diel
drift of emergent fry.
Water temperatures and ice conditions during grayling egg and a 1 evin
de'leloprnent may be easier to sample than the lower water temperatures
and thicker ice cover characteristic of Pacific salmon and char
incubation periods.
Laboratory studies should assess the effects of various durations of
low water temperatures on the development and survival of eggs and
alevins. Water temperatures at or below threshold levels could cause
morphological deformities. slow de.v.elopment rates and high mortality
among eggs and alevins. These investigations could explain the
apparent avoidance of spring streams by spawning Arctic grayling.
Weirs could be used to sample grayling from bog, mountain, spring and
lake inlet and outlet streams to relate residence time and migration
to stream flow. water temperature and other physica 1 and cnemica 1
stream factors.
Unique tags on upstream migrating adult and juvenile fish would
provide specific migration data. Data from consecutive years could be
used to determine Arctic grayling homing to specific spawning and
rearing streams.
Arctic grayling spawning sites should be studied more extensively in
north~rn lati tudes. Characteristics of spawning habitat in terms of
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water depths, current velocities and substrate conditions should be
compared with habitat availability for specific stream systems. Water
temperatures and fish spawning activities should be monitored to
detect spawning activity cycles.
Internal radio transmitters could be used to monitor grayling movement
to and within open-water rearing areas and overwintering sites. Radio
telemetry could be used to study fish migration rates and patterns.
Surgical implantation of radio transmitters is probably the best
method for spawned-out adult and large juvenile fish. Surgical
implantation has less affect on fish equilibrium than esophageal
insertion (Winter, Kvechle, Siniff and Tester, 1978). Rates of
healing, condition of internal organs and the occurrence of infection
among the fish should be determined for various water temperatures.
Criteria for radio transmitter selection should include size of
transmitter and antennae, transmitter various temperatures, and signal
receptions at various depths.
The feasibility of marking juvenile Arctic grayling with fluorescent
pigment should also be determined. A variety of color combinations
could be used to identify specific stream locations and dates of
marking. lack of scale development may prevent pigment retention by
yay Arctic grayling less than 11 months old.
Comprehensive sampling of fry would determine movement of yay Arctic
grayling. Weirs of small mesh screen could be used to monitor yay
Arctic grayling movements. However smaller mesh size also
necessitates more frequent cleaning of the weirs.
Weirs should remain within streams as long as possible prior to
freeze-up to monitor Arctic grayling movements and physi ca 1 and
chemical habitat components. Weirs could remain in spring streams
which remain ice-free to monitor the presence of fish.
-56-
Investigations shoul,., be conducted to detennine open-water, lotfc
microhabitat selection by juvenile and adult Arctic grayling using
techniques similar to those described by Everest and Chapman (1972).
These investigations should describe ~ater depths, current velocities,
substrate, proximity to nearest fish and instream cover. Snorkeling,
which has been used extensively in the Pacific Northwest and
elsewhere, can be used where bank observations of fish are difficult.
Microhabitat investigations could complement fish movement data from
weirs and radio telemetry studies. Microhabitat studies could also
explain the apparent segregation of various age classes of Arctic
grayli"g in certain rivers such as the Chena River.
Food availability and various physical and chemical habitat influence
the s~mmer, open-water distribution of Arctic grayling within streams.
Drift and substrate sampling of invertebrates can be used to assess
food availability and its relationship to the distribution of Arctic
grayling.
Studies of juvenile and adult Arctic grayling overwintering habitat
should continue. Gillnets, SCUBA and other technfquP.s could be used
to investigate these overwintering habitats.
Fish passage studies should be conducted to assess the abfl ity of
juvenile and adult Arctic grayling to ascend culverts and other high
current velocity areas. MacPhee and Watts (1976) detennined that the
ability to ascend culverts was a function of culvert length and
spawning condition. Studies of yoy and older juveniles would identify
fish movement during the summer rearing season. The possfbflfty that
excessive current velocities associated with culverts prevent young
grayling from reaching small rearing strP.ams should be investigated.
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VI. LITERATURE CITED
Alaska Department of Fish and Game. 1978. Alaska•s Fisheries Atlas.
Alaska Dept. of Fi!h and Game, Vol. II. 196 p.
Alt, K. 1977. Inventory and Cataloging of Arctic Area Waters. Alaska
Dept. of Fish and Game. Federal Aid in Fish Restoration, Annual
Report of Progress, 1g76·1977. 18(G-I·P):l-l£3.
Alt, K. 1978. Inventory and Cataloging of Sport Fish and Sport Fish
Waters of Western Alaska. Alaska Dept. of Fish and Game. Federal Aid
in Fish Restoration, Annual Report of Progress, 1977-1978.
19(G-I·P):36-60.
Alt, K. and R. Furniss. 1976. Inventory and Cataloging of North Slope
Water!. Alaska Dept. of Fish and Gane. Federal Aid in Fish
Restoration, Annual Report of Progress. 17{F-9-8) p. 129-150.
Bams, R. 1967. Differences in performance of naturally and artffically
propagated sockeye salmon migrant fry, as measured with swimming and
predation tests. J. Fish. Res. Board (An. 2415):1117·1153.
Bendock, T. 1979. Inventory and Cataloging of Arctic Area Waters. Alaska
Dept. of Fish and Game. Federal Aid in Fish Restoration, Annual
Report of Progress, 1978-1979. 20(G-I-I):1-64.
Bendock, T. 1980. Inventory and Cataloging of Arctic Area Waters. Alaska
Dept. of Fish and &alii!. Federal Aid fn F1sh Restoration, Annual
Report of Progress, 1979-1980. 21(&-I·I):l-31.
Bishop, F. 1971. Observations on spawning habfts and fecundity of the
Arctic grayling. Prog. Fish Cult. 27:12-19.
-58-
Srown, C. 1938. Observations on the life history and breeding habits of
the Montana grayling. Copeia (3}:132-13E.
Brown, C. and G. Buck. 1939. When do trout and grayling fry begin to take
food? J. Wildlife Mn9t. 3(2):134-140.
Chislett, G. and K. Stuart. 1g79, Aspects of the life history of A~tic
grayling in the .)ekunka River drainage, British Colunmia. British
Columbia Fish and Wildlife Branch. 110 p.
Craig, P. and P. McCart. 1974a. Classification of stream types in
Beaufort Sea drainage between Prudhoe Bay, Alaska and the Mackenzie
delta .!!!. Classification of streams in Beaufort Sea drainages and
distribution of fish in Arctic and sub-Arctic drainages. P.J. McCart,
ed. Canadian Arctic Gas Study Co. Biological Report Series.
7(1):1-47.
Craig, P. and P. McCart. 1974b Fall spawning and overwintering areas of
fish populations along routes of proposed pipeline between Prudhoe Bay
and the Mackenzie delta, 1972-73 ~Fisheries Resea~h Associated with
Proposed Gas Pipeline Routes fn Alaska, Yukon. and the Northwest
Territories. P.J. Mccan:, ed. Canadian A~tic Gas Study Ltd./Alaska
Arctic Gas Study Co. Biological Report Series. 15(3):1-36.
Craig, P. and V. Poulin. 1974. Life history and movement of Arctic
grayling (Thymallus arctfcus) and juvenile Arctic char (Satvelinus
alpinus) in small tundra stream tributary to the Kavik River, Alaska
~Life Histories of Anadromaus and Freshwater Fishes in the Western
Arctic. P.J. McCart, ed. canadian A~tfc Gas ltd./Alaskan Arctic Gas
Study Co. Sfologfcal Report Series. 20(2):1-53.
-59-
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Craig, P. C. and J. Wells. 1975. Fisheries fnvesti9at1ons in the
Chandalar River region, northeast Alaska ~ Fisheries Investigations
in a Coastal Region of the Beaufort Sea. P.C. Craig, ed. Canadian
Arctic Gas Study Ltd./Alaskan Arctic Gas Study Co. Biological Report
Series. 34(1):1·114.
Creaser, C. and E. Creaser. 1g35, The grayling of Michigan. Pap. Mich.
Acad. Sci., Arts & Letters. 20:599-611.
Cuccarease, S., M. Floyd, M. ~elly and J. LaBelle. 1980. An assessment of
environmental effects of construction and operation of the proposed
Tyee Lake hydroe 1 ectri c project Petersburg and Wrange 11 , A 1 ask a,
Arctic Environmental l'1formatfon and Data Center, University of
Alaska, Anchorage, Alaska.
deBruyn, M. and P. McCart. 1974. Life history of the grayling (Thymallus
arcticus) in Beaufort Sea drainages in the Yukon Territory .!!!.
F is he ri es Research AHoci a ted wi th Proposed Gas p 1 pe 1 i ne routes 1 n
Alaska, Yukon and Nort~est Territory. P.J. McCart, ed. Canadian
Arctic Gas Study Ltd./Alaskan Arctic Gas Study Co. Biological Report
Series. 15(2):1·110.
Elliott, G. 198D. First interim report on the evaluation of stream
crossings and effects of channel modifications on fishery resources
along the route of the trans-Alaska pipeline. U.S. Fish and Wildlife
Service, Special Studies. Anchorage, Alaska. 77 p.
Everest, F. and D. Chapman. 1g12. Habitat selection and spatial
interaction by juven1le chinook salmon and steelhead trout in two
Idaho streams. J. Fish. Res. Board. Can. zg:g1-100.
Henshall, J. 1907. Culture of the Montana grayling. U.S. Fisheries
Station. Bozeman, Montana.
-6D-
Jessop, Chang-Kue, Lilley and Percy. 1974. A further evaluation of the
resou rc:es of the Mackenzie River va 11 ey as related to :l i pe 1 i ne
development. Report No. 747. Canada F1st1eries and Marine Service,
Dept. of the Environment. 95 p.
Kratt, L. and J. Smith. 1977. A post-t1atching sub-gravel stage in the
life t1istory of the Arctic grayling, Tt1ymallus arcticus. Trans. Am.
F1st1. Soc. 106(3):241-243.
Kruse, T. 1959. Grayling of Grebe Lake, Yellowstone National Park,
Wyoming. F1st1. Bull. 149. U.S. Fish and Wildlife Serv. 59:305-351.
LaPerrier, J. a~d R. Carlson. 1973. Thermal tolerances on interior
Alaska Arctic grayling. Institute of Water Resources. University of
Alaska, Fairbanks. Report No. IWR-46. 36 p.
Lister, 0. and H. Genae. 1970. Stream t1abitat utilization by cot1abiting
underyearlings of chinook salmon (.Q.. tshawytscha) and cot1o (.Q..
kisutch) salmon in the Big Qualicum River, British Columbia. J. Fish.
Res. Bd. Canada, 27:1215-1224.
McCart, P.
Alaska
Arctic
1974. Late winter surveys of lakes and streams in Canada and
along tt1e gas pipeline routes under consideration by Canadian
Gas Study Limited. 1972-1973. in Fisheries Researct1
Associ a ted witt1 Proposed Gas Pipe 1 i ne Routes 1 n A 1 a ska, Yukon and
Northwest Territories. P. McCart, ed. Canadian Arctic Gas Study
Ltd./Alaskan Arctic Gas Study Co. Biological Report Series.
15(1): 1-183.
McCart, P., P. Craig and H. Bain. 1972.
investigations in tt1e Sagavanirktok
Alyeska Pipeline Service Co. 170 p.
-61-
Report on fisheries
River and neighboring drainages.
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MacPhee, C. and F. Watts. 1976. Swimming performance of Arctic grayling
in highway culverts. Final Report to U.S. Fish and Wildlife Service.
Anchorage, Alaska. 41 p.
~elson, P. 1953. Life history and management of
(Thymallus signifer tricolor) in Montana.
18(3):324-342.
the American grayling
J. Wildlife Mgt.
Netsch, N. 1976. Fishery resources of waters along the route of t~e
trans-Alaska pipeline between Yukon River and Atigun Pass in North
Central Alaska. U.S. Fish and Wildlife Service. Resource Publication
124. 45 p.
Pearse, G. 1974. A study of a typical spring-fed stream of interior
Alaska. Alaska Dept. of Fish and Game. Federal Aid in Fish
Restoration, Annual Report of Progress. 1973-1974. Project F-9-6.
Phinney, 0. 19 __ . Mass-marking small fish with fluorescent pigment by
means of compressed air. University of Washington Fish. Res. Inst.
Cirt. 66-6. 4 p.
Phinney, 0., 0. Miller and M. Dahlbert. 1967. Mass-marking young
salmonids with fluorescent pigment. Trans Amer. Fish. Soc.
96(2):157-162.
Rawson, D. 1950. The grayling {Thymallus signifer) in northern
Saskatchewan. Canadian Fish Cult. B p.
Reed, R. 1964. Life history and migration patterns of Arctic grayling.
Alaska Dept. of Fish and Game. Research Report No. 2. 30 p.
Reed, R. 1966. Observation of fishes associat•'" with spawning salmon.
Trans. Am. Fish. Soc. 96(1}:62-66.
qoguski, G. and P. Winslow. 1970. Monitoring and evaluation of Arctic
waters with emphasis on the North Slope drainage. Alaska Dept. of
Fish and Game. Federal Aid in Fish Restora~ion. Annual Report of
Progress, 1969-1970.
Schallock, E. and F. Lotspeich. 1974. Low winter dissolved oxygen in some
A 1 askan rivers. U.S. Env i ronmenta 1 Protect; on Agency. Eco 1 ogi ca 1
Report Series Report. 33 p,
Scott, w. and E. Crossman. 1973. Fresnwater fishes of Canada. Fish. Res.
Board. Can. Pub. 173. 996 p.
Tack, S. 1971. Distribution, abundance and natural his~ory of the Arctic
grayling fn the Tanana River drainage. Alaska Dept. of Fish and Game.
Federal Aid fn Fish Restoration, Annual Report of Progress, 1970-lg71.
12(F-9-3}. 35 p.
Tack, S. 1972. Distribution, abundance and natural histo~ of the Arctic
grayling in the Tanana River drainage. Alaska Dept. of Fish and Game.
Federal Aid in Fish Restoration, Annual Report of Progress, 1971-1972.
lJ(F-9-5). 34 p.
Tack, S. 1973. Distribution, abundance and natural history of the Arctic
grayling in the Tanana River drainage. Alaska Dept. of Fish and Game.
Federal Aid in Fish Restoration, Annual Report of Progress, 1972-1973.
14(F-9-6). 34 p.
Tack, S. 1980. Distribution, abundance and natural history of the Arctic
grayling in the Tanana Rfver drainage. Alaska Dept. of Fish and Game. {
Federal Aid in Fish Restoration, Annual Report of Progress, 1979-1980.
2l(F-90-12). 32 p.
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Tripp, D. B and P. J. McCart. 1974. Life histories of grayling
(Thymallus arcticus) and longnose suckers (Catostomus catostomus) in
the Donnelly River system, Northwest Territories~ Life Hi$tories of
Ana.dromous and Freshwater Fishes 1n the Western Arctic. P.J. McCart,
ed. Canadian Arctic Gas Study Ltd./Alaskan Arctic Gas Study Co .
Biological Report Series. 20(1):1·91.
Tryon, C. 1947. The Montana grayling. Prog. Fish. Cult. 9{3):136-142 .
Van Hyning, J. 1978.
River drainages.
Fall and winter fish studies on the upper Tanana
Northwest Alaskan Pipeline Co. 77 p.
Vascotto, G. 1970. Summer ecology and behavior of the Arctic grayling of
McManus Creek, Alaska. M.S. Thesis. University of Alaska, Fairbanks .
132 p.
Warner, G. 1955. Spawning habits of grayling in interior Alaska. U.S.
Fish and Wildlife Service . Federal Aid in Fish Restoration, Quarterly
Progress Report. (F·1-R-5). 10 p.
Williams, F.T. and C. Morgan. 1974. lnventory and cataloging of sport
fish and sport fish waters of the Copper River and Pri nee Wil 1 i am
Sound drainages and the Upper Susitna River drainage. Alaska Dept. of
Fish and Game. Federal Aid in Fish Restoration, Annual Report of
Progress, 1973-1974. Project F-9-6, 15{G-I-F):24 p.
Williams, F.T. 1975. Inventory and cataloging of sport fish and sport
fish waters of the Copp~r River and Prince William Sound drainages and
the Upper Susitna Ri ·1er drainage. Alaska Dept. of Fish and Game.
Federal Aid in Fish Restoration, Annual Report of Progress, 1974-1975.
Project F-9-7, 16(G-I-F):23 p.
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~il liams, F.T. 1976 . Inventory and cataloging of sport fish and sport
f ish waters of the Copper River and Prince William Sound drai"ages and
the Upper Susitna River drainage . Alaska Dept. of F i sh and Game.
Federal Aid in Fish Restoration, Annual Report of Progress, 1975-1976
Project F-·9-8, l7(G-I-F) :22 p.
Wilson, W., E. Buck, G. Player and L. Dreyer . 1977. Winter water
availability and use conflicts as related to fish and wildlife in
arctic Alaska. A synthesis of information. Arctic Environmental
Infonnation and Data Center. Univerisity of Alaska, Ancnorage.
222 p.
Wilson, W .• E. Trihey, J. Baldrige, L. Evans, J. Thiele a nd 0. Trudgen.
1981. An assessment of environmenta 1 effects on opl!ration of the
proposed Terror Lake hydroelectric f~~i l ity, Kodiak, Alaska. Arctic
Environmental [nfonnation and Data Center. University of Alaska,
Anchorage. 419 p.
Winter, J.D., V. Kuechle, 0. Siniff and J.R. Tester. 1978. Equipment and
methods for radio tracking freshwater fish. Mi scel l aneous Report
152-1978. Agricultural Experiment Station. University of Minnesota.
18 p.
Wojcik, F.
Plan o.
1954 . Spawning habits of grayling in interior Alaska. Work
Job No. 1., Alaska Game Conwnission. U.S. Depart of the
Interior, Fish and Wildlife Service. Quarterly Report. Report No. 2.
Wojcik. F. 1955. Lffe history and management of the grayling in interior
Alaska. Unpub . M.S. thesis. Univ . of Alask, Fairbanks. 54 p.
Yoshihara, H. 1972. Monitoring and evaluation of A~ctic waters with
emphasis on the North Slope drainages. Alaska Dept. of Fish and Game.
Federal Aid in Fish Restoration, Annual Report of P~gress, 1971-1972.
Project F-9-4, 13(G-lll-A):49 p.
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FRESHWATER HABITAT
RELATIONSHIPS
DOLLY VARDEN-SALVELINUS MALMA (WALBAUM)
ALASKA DEPARTMENT OF FISH & GAME
HABITAT PROTECTION SECTION
RESOURCE ASSESSMENT BRANCH
JULY, 1982
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FRESHWATER HABITAT RELATIONSHIPS
DOLLY VARDEN CHAR (SALVELINUS MALHA (WALBAUM))
By
e Steven W. Kn1~er
Alaska Department of Fish and Game
Habitat Division
Resource Assessment Branch
570 West 53rd Street
Anchorage, Alaska 99502
May 1981
Acknowledgements
Fisheries biologists from the Alaska Department of Fish and Game, the U.S.
Forest Service, U.S. Fish and Wildlife Service, National Marine Fisheries
Service-Auke Bay Laboratory, Bureau of Land Management, and the Arctic
Environmental Information and Data Center provided insight into this
project and deserve recogn1 t1 on. The ass 1 stance of 11 bra ri ans at the
Alaska Resource Library, the U.S. Fish and Wildlife Service
Library-Anchorage a~d the Commercial Fisheries Library-Alaska Department of
Fish and Same-Juneau is heartily appreciated.
This project was funded by the U.S. Fish and Wi 1 dli fe Serv1 ce, Western
Energy and Land Use Team, Habitat Evaluation Procedure Group, Fort Collins,
Colorado. Contract No. 14-16-0009~79-119.
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TABLE OF CONTENTS
I Page
I. Introduction 1
I A. Purpose 1
B. Distribution 2
I c. Life History Summary 2
D. Economic Importance 14
II.
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Specific Habitat Requirements 15
A. Upstream Migration 15
1. Stream Flow 15
I 2. Water Temperature 15
3. Light 16
4. Current Velocity 16
I B. Spawning 16
1. Current Velocity 16
2. Substrate 16
I 3. WatPr Depth 17
4. Cover 17
I c. Inmigrant Migration to
Overwintering Areas 17
I 0. Inmigrant Overwintering Areas 18
E. Egg and Alevfn Development 18
1. Water Temperature 18
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F. Summer Juvenile Rearing 18
1. Water Depth 18
I 2. Current Velocity 18
3. Instream Cover 19
4. Substrate 19
I G. Juvenile Migration to
Overwintering Areas 20
1. Water Temperature 20
I 2. Stream Flow 20
H. Juvenile Overwintering 'Areas 20
I 1. Water Temperature 20
2. Stream' Flow 21
3. Substrate 21
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I. Juvenile Migration to
Summer Rearing Areas 22
1. Water Temperature 22
2. Stream Flow 22
J. Inmigrant Migration to Sea 22
1. Water Temperature 22
2. Stream Flow 22
3. Light 23
K. Smolt Migration to Sea 23
1. Water Temperatures 23
2. Stream Flow 24
3. Light 24
r r 1. Conceptual Suitability Index Curves 25
rv. Deficiencies in Data Base 26
v. Recommendations and Further Studies 29
VI. Li te ra tu re Cited 33
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I. INTRODUCTION
A. Purpose
The purpose of th1s project is to describe how selected physical
and chemical features of lotic habitat within Ala.ika influence
the survival and behavior of the various life stages of
anadromous Dolly Varden char {Salvelinus malma (Walbaum)).
Objectives of this project are:
1. to gather data from published and unpublished sources within
Alaska and from conversations with Alaskan fishery
biologists concerning the relationships between lotic
aquatic habitat. and anadromous Dolly Varden survival and
behavior.
2. to develop an Alaska data base for habitat-anadromous Dolly
Varden char relationships. Because there are not sufficient
data for the relationships betlllleen anadromcus Dolly Varden
char and conditions of the habitat, habitat suitability
index relationships were not developed.
J. to identify data gaps and recommend appropriate projects to
alleviate these gaps.
The following Life History Su11111ary and Specific Habitat
Relationships/Requirements sections w111 fdentffy the lotfc
habitat relationships of the various life history and seasonal
behavior stages of the anadromous Dolly Varden char which
include:
upstream soawn1ng migration,
spawning,
inmigrant migratio~ to overwintering areas,
inmigrant overwintering areas,
egg and alevin development,
summer juvenile rearing,
juvenile migration to overwintering areas,
juvenile overwintering areas,
juvenile •fgration to summer rearing areas,
inmigrant wngratfon to sea, and
smolt migration to sea
B. Distribution
The taxon~ of the Dolly Varden char, Salvelinus m~lma (Walbaum)
is quite complex and a topic of debate. Morrow (1980) recognizes
a northern and a sou them fonn of Do 11 y Varden char, wf th the
northern form equivalent to the anadromous Arctic char,
Sa 1 ve 1 1 nus a 1 pi nus ( L1 nnaeus) , and the southern fonn compr1 sing
the Dolly Varden char. Dolly Varden char is separate from the
bull char, Salvelinus confluentus, which occurs fn British
Columbia, Washington, Oregon, Idaho and Montana (Behnke, 1980}.
For purposes of this paper, Dolly Varden char are defined as
those fish which occur south of the Arctic char and north of the
bull char. Anadromous and nan-anadromous Dolly Varden char are
distributed within Alaska south of the Alaska Range.
C. L ffe History SUIIIIIry
Migration of ature and i-ture Dolly Varden char frc. the ocean
to southeastern Alaska •Y be concu,..nt and can extend frc. July
through October. Ia.ture individuals m~y stay in streams as
1 ong as spawners (two to three months} a 1 though non-spawning
fnmature Dolly Varden char seldom reNin mere than one month
prior to earigrating to other stream and lake systems, probably to
overwinter. Atout 80S of the non-spawning immigrants left Hood
Bay Creek, AT aska before the spawning fish (Armstrong and
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:.linslow, 1968). Dur~.ng 1966, tagged irrmatur-e fish remained in
Hood Bay Creek for 14 days (Armstrong, 1967), and mature fish for-
85 days.
Age at maturity is variable; mos:: Dolly Varden in southeast
AJ aska reach maturf ty by age fotJ .-or five. Ma 1 es may matur-e
before females (Armstrong and Blackett, lg65).
Tagging studies in southeast Alaska {Armstrong. 1965a and 1974;
Blackett, 1968; Heiser, 1966) indicated that mature, anadromous
Dolly Varden char use their natal streams to spawn and lakes to
overwinter. Imatu re fish ori gina t f ng in streams without 1 a kes
may enter several streams prior to finding a lake for
verwintering. Immature fish of lake-stream origin probably
re·enter the same system to overwinter. These tagging studies
i nd 1 ca te that 1 nma tu re and spawned -out ana d romou s Do 11 y Varden
char from numerous stream systems may use the same lake for
overwintering.
The Dolly Varden, like other chars, usually spawns between
Septenme,. and Noveamer (Scott and Crossman, 1973). Blackett
(1968) determined that the peak of spawning activity in Hood Bay
Creek occurred be~~een late October and early November. Spawning
occurs in other southeastern Alaska streams from mid-September to
mid-October (Blackett, 1968; Blackett and Armstrong, 1965). Both
sexes display spawning coloration, although males are often
scarlet on the ventral side and have black snouts.
Selection of spawning s1tes by anadromous Do1 ly Varden char
appean to be influenced by various physical factors including
curre;,t velocity, water depth and substrate composition. Fish
have been reported spawning in sidechannel and mainstem
riffle/n.m and pool reaches of streams on Kodiak Island and in
southeast Alaska (Blackett, 1968; Wilson, Trfhey, Baldrige,
Evans, Thiele and Trudgen, 1981).
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Limited observations indicate tha._ ... fish's spawning behavior
is similar to other chars (Blackett, 1g68; Scott and Crossman,
1973). Fish are usually paired, although more than one male may
accompany a female (the largest male is usually dominant;. The
female is solely responsible for excavation of the redd (a
depression in the stream substrate where fish spawn and deposit
fertilized eggs). She forms the redd by turning on her side and
thrashing the substrate with her caudal fin. The completed redd
is typically oblong shaped. Dimensions of the redd vary with the
size of the female and substrate and current velocities; redds
are generally 30 to 61 em (12-24 in) long and may be as deep as
30 em (12 in). The male spawner actively defends the redd from
male intruders and will nip and bite other males, sometimes
grasping another male in the caudal peduncle far up to six
seconds. Female spawners are not aggressive. (Blackett, 1968)
Fecundity of Dolly Varden char is variable among anadrorrmus
Alaska stocks ~nd is greater with increasing female fish age and
length (Blackett, 1968}. For exaq,le, fema 1 es about 300 mm fl
(fork length} from Hood Bay Creek, Alaska contained less than
1000 eggs and females exceeding 450 ~ f1 supported at least 2000
eggs. Ripe eggs are usually 0.45 to D.6 en in diameter
(Blackett, 1968).
The spawning pa 1 r descend 1 nto the redd and press a ga 1 ns t each
other laterally. After the pair completes spawning the female
may dig at the upstream end of the redd and displace gravel over
the fertilized eggs. This gravel layer probably protects the
eggs from sunlight and predation, reduces mechanical disturbance
by ice and other objects while allowing water to transport oxygen
to and metabolic wastes from the developing eggs.
Fish lillY spawn again with the same or a different partner and,
unlike salmon, are capable in subsequent year5. Males are less
likely to survive spawning than females (Annstrong and Kissner.
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1969). Armstrong and Kissner (1969) estimated that post-spawning
mortality in Hood Bay Creek9 Alaska was about 61% for male fish
and 4~ for female fish in 1967 and about 49% for male fish and 9~
for female fish in 1968. This differential mortality is not
understood, but the aggressive behav1or of male spawners is
pr~bably a factor .
Emigration of spawned-out Dolly Varden char to the sea or to
over.tintering area.s usually occurs w1 thin two weeks after
completion of spawning. Fish surveys in Hood Bay Creek indicated
that a 11 spawned-out adu 1 ts 1 eft the stream by 1 ate November
(Armstrong anu Winslow, 1968}. These fish may have entered the
ocean to feed for severa 1 months or moved to overwintering areas
(Armstrong, 1974).
Longevity of Dolly Varden char is variable but fish in southeast
Alaska may live nine to twelve years (Heiser, 1966; Armstrong,
1963).
Development of Dolly Varden eggs and alevins to the emergent fry
stage requires about 210 days (Blackett, 1968). Hatchin] has
been documented from 129 to 136 days at 8.5°C. The 1.5 to 2.0 em
long alevin typically remains in the gravel for an additional 60
to 70 days.
Limited research has focused on the survival of anadromous Dolly
Varden eggs and alevins. Blackett (1968) estimated that egg to
alevin survival within an area of Hood Bay Creek, Alaska was
about 41 percent.
Intragravel flow, dissolved oxygen and sediment composition can
influence the development and survival of salmonid eggs and ·
alevfns. Little work has focused on egg and alevin survival in
relation to physical and chemical factors. Inferences can be
made from work on other salmonfd species. For example, transport
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of dissolved oxygen to and metabolic wastes from develop1ng eggs
and alevins by intragravel flow is crucial for survival of
salroonid eggs and alevins (Vaux, 1962; Wickett, 1958).
Relatively low intragravel dissolved oxygen levels during the egg
development stage of various salmonfds may increase egg
roortality, delay or hasten egg development or reduce the fitness
of alev1ns (Alderdice, Wickett and Brett, 1958; Silver, Warren
and Duodoroff, 1963). Coble (1961) and other investigators have
detemined that salmonid egg survival 1s enhanced by increased
1ntragravel flow despite sufficient 1ntragravel dissolved oxygen
1 evel s. Factors wh"ich caul d reduce the quantity or qua 1 i ty of
intragravel water include reduced streamflow, sedimentation, and
accumulation of organic debris (McNeil and Ahnell, 1964; Koski,
1966; Reiser and Bjornn, 197g).
Severt streamflow alterations can hann developing salmonid eggs
and embryos. Mechanical scouring of the redd could dislodge
substrate and destroy developing eggs and embryos. McNeil (1966)
observed damaged pink and chum salmon redds and displaced eggs of
both s pee i es 1 n seve ra 1 sou thea stern A 1 as ka streams f o 11 owi ng
autumn spates. Koski {1966} reported low survival of coho salmon
eggs subjected to freshets several days after redd construction.
Low flows and cold winter temperatures could cause redds to
desiccate or to freeze. McNeil (1966) noted low survival of pink
and chum salmon eggs in streams with low winter flows during
harsh winter conditions in southeast Alaska.
Koski (1966) determined that significant accU111.1lat1ons of fine
sediments within chum salmon redds can retard or prevent the
emergence of fry. Deposition of fine sediments in anadromous
Dolly Varden char spawning areas could retard or prevent fry from ·
emerging.
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Water temperature affects natching rates of salmonid eggs; warmer
than no~l water temperatures can accelerate hatching and result
in earlier than no~l fry emergence (Sheridan. 1962). Blackett
{1968) detenn1ned that Dolly Varden began hatching after 129 days
in water with a mean temperature of 8.5°C. No upper or lower
temperature tolerance limits of Dolly Varden char eggs or alevins
were found in the literature.
Upon emergence, anadi"OIIIJus Dolly Varden char occupy relatively
quiet stream reaches. Blackett (1968) and Annstrong and E111ot
{1972) noted yoy fish in shallow stream margins of Hood Bay
Creek, Alaska. Annstrong and Elliott also !ound yay fish in
rivulets along Hood Bay Creek during early summer. These stream
margins were ofte" only 1 m wide and 4 em deep. No evidence of
yoy ffsh entrapal!nt fJ'i dewatering within these stream marg1 ns was
detected in the literature. Yoy Dolly Varden, which feed
primarily from the benthos. characteristically remain on or near
the substrate, sometimes remaining motionless and occupying
gravel interstices. Yoy coho salmon, however, feed primarily
from the surface within these sane areas (Blackett, 1968).
Earlier research indicates that juvenile salmonids occupy areas
(Wickham, 1967) with relatively slow current velocities often
adjacent to areas with faster current velocities and with higher
densities of drifting invertebrates (Everest and Chapman, 1972).
A fish moves periodically into the areas of greater food
availability to forage and returns to resting positions in areas
of slower current. This •fnilllfzes energy expenditures and
maximizes f -eding success (Cha~n and Bjornn, 1969}. As fish
grow they o /ten occupy deeper and faster areas of the stream
(Everest and ChaJ~~Bn, 1972). This behavior probab 1 y app 1 ies to
juvenile De lly Varden char. Leggett (1969) suggested that the
cylindrical body shape of bull char, which 1s morphologically
similar to Dolly Varden, probably enables these fish to occupy
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iH"eas of faster" current velocities than most other salmonids.
Other salmonids typically have lateraily compnessed bodies.
Habitat selection by older pre-smolt Dolly Varden char is not
well documented, although the char" ane distributed in deeper,
sometimes faster habitat than yay fish. Heiser (1966) noted that
yearling and older pre-smolt Dolly Varden char occupied both
still and flowing reaches of several inlet tributaries of Eva
Lake. This 1 ake was character"f zed by gravel and s 11 t substr·ates
with varying amounts of vegetation. Researchers studying Hooc
Bay Creek classified it into ten habitat types ranging frou
sloughs, undercut bank margins, pools and riffles. Baited minn~1
traps were placed in each habitat type, and length and fnequenc'
of Pr"!-smolt coho salmon and Dolly Varden char" were ;ompared fron
each habitat type during July and August 1971. Pre-smolt Dolly
Varden char and coho sal1110n were taken from all habitat types.
The smallest Dolly Varden char (41-50 mm) were taken in sloughs
and sidechannel undercut bank areas, and the largest Dolly Vard£1
.ere found in r1 ffl es.
Yearling and older pre-smolt Dolly Varden char occupy 11 pools
quiet sfdechannels and sloughs and tributaries off the mainstem!
of both ••• • the Terror" and Kfzhuyak Riven, Alaska, altho11<;~
juven11 e fhh are occasfona lly found behind boulders in faster
water (Wilson, Trihey, Baldrige, Thiele and Trudgen, 1981).
Minnow traps were found to give a biased indication of habitat
occupancy by fish because the bait lillY attr"act fish from a
consfderab 1 e d1 stance.
actually residing ll1illy
Habitat conditions where the fish are
be quite df fferent fro~~ cond1ti ans
immediately around the trap.
Reed and Anns t rang (19 71 ) noted that j uven 1 1 e coho sa 1 mon and
Dolly Varden char" were capable of entering and exiting baited
minnow traps fished for 24 hours. The p lacl!t!l!nt of two i engths
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of wire across the entrance to each trap resu .1ted in a higher
fish retention rate.
Distribution and abundance of pre-smolt Dolly Varden char may be
influenced by intra and interspecific fish interactions.
Observations of juvenile coho salmon~ Oncorhynchus kisutch
(Walbaum)~ and Dc1iy Varden char in Hood Bay Creek and in
quari ums (Annstrong and Elliott, 1972) revea 1 ed that Dolly
Varden fry are aggressive among themselves and in association
w1 th coho fry. Dolly Varden were frequently attacked by coho
sa 1 mon fry but were never observed at tacking coho fry. More
Dolly Varden fry remained ·near the substrate when associated with
coho fry than when they were alone. Coho fry occupied the upper
half of the aquaria when alone and with Dolly v~rden char fry.
Aquaria tests with older, pre-smolt Dolly Varden char and coho
salmon indicated that Dolly Varden established and defended
territories when alone and when with ·coho fingerlings. Dolly
Varden fingerlings generally occupied positions within the
aquaria at or near the bottom, but when alone, they occupied more
mid and shallow depth positions. Coho fingerlings were
consistently found in the upper strata of aquaria a..,d seldom
attacked Dolly Varden char fingerlings (Armstrong and Elliott,
1972).
Juvenile, anadromous Dolly Varden char grow relatively slowly
during the three to four years prior to emigrating to the Pacific
Ocean. Young of the year fish from Hood Bay Creek, A 1 ask a grew
about 10 mm between July 7 and October 1, 1965, reaching a w.ean
fork 1 ength of about 38 mm ( B 1 acket t, 1968) . Growth rates of
pre-smolt fish may vary and length ranges of yay and older age
classes often overlap. Pre-smelt fish fn Alaska generally grow ·
10 to 30 mm annually, primarily during the summer months
(Armstrong, 1963; Heiser, 1966; Blackett, 1968).
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The summer diet of stream rearing pre~smclt Dolly Varden char is
influenced by food availability, fish size and stream habitat
selection (Annstrong and Elliott, 1972). Gut analysis of
pre-smolt fish from Hood Bay Creek during the sunmer rearing
period {April to November) showed that substantial nunmers of
iii'IMture and adult aquatic insects were eaten throughout this
period. Emergent and emigrant yay salmon consumed invertebrates
from April to June and salmon eggs from July to November.
Relatively large pre-smelt Dolly Var~en char ate more and larger
food items than smll er fish. Pre-smo 1 t fish occupy1 ng stream
reaches characterized b} overhanging vegetation and relatively
low current velocities (such as sloughs and stream margins)
generally consumed more terrestrial and surface floating insects
than fish occupying m1d-channe1 areas with moderate to fast
current velocities and with greater invertebrate drift. ~o drift
samples were taken to compure dr1ft compos 1ti on with fish gut
contents. Pre-smolt Dolly Varden char occupy areas at or very
near the substrate in streams with coho sall!lln. The char may
browse along the substrate or conswne drifting invertebrates
(Amstrong and Elliott, 1972).
Land practices which result in removal of vegetation along stream
margins and deposition of fine sedinEnts in the stream channel
could possibly reduce the abundance and fitness of pre-smolt
Dolly Varden char (Amstrong dRd Elliott, 1972; Elliott and
Oinneford, 1976).
The season~l distribution of pre-smolt Dolly Verden char is
apparently influenced by fl uc:tuati ng flows and decli n1 ng water
temperatures during the late sunner and fall. Fish appear-ed to
be distributed evenly throughout Hood Bay Creek, Alaska from Juiy
through September. By November there were considerably fewer
fish in the duwnstream reaches. Significantly more pre-sl!lllt
Dolly Varden char were captured in the upper stream reaches
during this time. Fish were observed schooling in mid-stream in
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October, behavior which had not been noted previously. Water
temperatures du ri ng July through September ranged from 5o to 9 o C
(41"-48°r), and water temperatures during October and November
were substantially lower (Blackett, 1968}. Armstrong and Elliott
{lg72) found substantial nuntlers of pre-sroolt Dolly Varden char
in the upper reaches of Hood Bay Creek in late winter where water
temperatures were consistently 611 C wanner than in downstream
reaches. Armstrong and Elliott (1972) concluded that warm,
ice-free reaches of Hood Bay Creek attract overwintering
pre-sroolt Dolly Varden char and that survival rates are higher in
the warmer regions. Downstream reaches of Hood Bay Creek become
frozen during the winter.
Elliott and Reed (1974) and Elliott (1975) determined that
pre-srool t Dolly Varden char leave Starrigavan Creek, Southeast
Alaska and enter spring-fed tributaries during autumn. The
trfbutarie~ are characterized by relatively warm winter water
temperatJres and somewhat stable flows. Immigration of pre-smolt
Dolly Varden char and coho salroon to overwintering areas usually
cormll!nced in September, peaked in early October, and ceased by
December. Spates and decreasing water temperatures within
Starrigavin Creek appeared to stimJlate movement of fish into
these streams.
Selection of lotic overwintering habitat by juvenile Dolly Varden
char 1s not well documented.
Elliott and Reed (1974} noted that juvenile Dolly Varden char in
Spring ?ond Creek burrowed into logging slash and other debris
when water temperatures decreased to 211 C. Fish reappeared when
water temperatures rose above about 2°C.
Other researchers have noted roovemen! of juvenile salroonids when
stream wa":er temperatures decrease in the autumn. As water
temperatures decrease, fish activity levels and digestion rates
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drop (Reimers, 1957; Chapman and Bj ornn, 1969) . Chapman (1966)
stated that the distribution of winter ~aring juvenile salmonids
in the Pacific Northwest and other temperate areas is probably
space related. Fish reduce feeding and seek overwintering areas
when water temperatures decrease to or below 5°C. Winter stream
conditions, including reduced flows, partial or complete
ice-cover and water temperatures at or near freezing do not
constitute suitable aquatic habitat for ~aring salmonids.
Juvenile salmonids reduce the risk of mechanical injury and
displacement by avoiding shallow, cold stream reaches by moving
to warmer and deeper stream reaches, burrowing into substrate
interstices, or associating with subnerged logs and root masses.
Factors which could adversely affect the winter survival of
juvenile (pre-s~,.,lt) anadromous Dolly Varden include freezing
c~uring streamflow reductions and displacement and injury from
dis 1 odged substrate materia 1 during spates. The movement of
juvenile Dolly Varden to more suitable overwintering habitat in
response to ~duced winter streamflows has not been documented.
Bustard (1973) reported the movement of yay steelhead trout
(Salmo gaf~neri (Rfcha~son)) to overwintering areas in response
to altered flows in Carnation Creek, British Columia. Busta~
(1973) speculated that yoy steelhead trout overwintering within
'~small rubble, often less that 15 em in diameter11 could be
susceptible to injury from substrate movement during spates in
Carnation Creek, British Columbia. Hartmann (1968) found that
s~able ·Submerged log jams provide excellent winter habitat for
fish, although loose logging debris which is susceptible to
displacement by flood~; ts not suitable for salmnfd overwintering
habitat.
Renlval of submerged logging debris, natur~lly occurring fallen
trees and root masses, and destruction of bankside vegetation and
associated submerged roots could significantly reduce the
abundance of juvenile Dolly Varden char in streams where these
materials are used for overwintering habitat. Bustard (1973)
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emphasized the need to identify and preserve quality fish
overwinteiing habitat, especially along small permanent and
intermittent streams that may be overlooked as valuable fish
habitat during timber harvest operations.
Food consumption by pre-smolt Dolly Varden char in overwintering
areas is much reduced from summer levels (Anmstrong and Elliott,
1972; Elliott and Reed, 1974). Annstrong and Elliott (lg72)
ascribed the difficulty in obtaining juvenile Dolly Varden char
by baited minnow traps in the headwaters of Hood Bay Creek to the
relatively low water temperatures, 5°C tc 6.1°C, which influence
fish activity.
Emigration of pre-smolt Dolly Varden char from winter tc surrmer
rearing areas appears to be influenced by water temperature and
flow conditions (Elliott. 1975 and 1976). Rising water
temperatures were assc.ciated with the emigration of pre-smolt
Dolly Varden from Spring Pond Creek~ Alaska. Fish emigrated from
March or April through June. Floods appear to retard fish
emigration within Spring Pond Creek.
Emigration of inmature and mature anadromous Dolly Varden from
lakes usually occurs after ice breakup in lakes. Factors, other
than the breakup of ice, which could influenc~ the timing of fish
emigration from lakes include water temperature and streamflow
{Anmstrong, 1965b).
Behavioral and physiological changes, collectively termed
smoltification. and subsequent seaward migration of age 2 to 4
and sometimes older juvenile anadromous Dolly Varden typically
occurs in southeast Alaska streams fl"'lrr. April to June. Non--lake
systems may support an additional autumn smolt migration
(Annstrong, 1965 and 1g7o; Armstrong and Kissner. lg6g).
Physiological changes for salinity tolerance, probably begin
before ~eaward migration (Conte and Wagner. 1g65). Factors
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affecting timing of smoltification are speculative but fish size
appears to be influential (Armstrong, 1965a). For example,
Armstrong (1965) suggested that fish whicl", reach migratory size
several months after spring leave streams without lakes, such as
Hood Bay C~k. and enter stream! with lakes, such Js Eva Creek,
where they overwinter until the following spring. Fish have not
been found to migrate sealillla rd from 1 ake-stream systems, such as
Eva Lake during autumn. Annstron~ (1965) speculated that fish
that reach migratory size in lakes during the autumn probably
overwinter and migrate seaward the fo11owirg spring. Dolly
Varden smelts may range in length from about 100 to 180 mm fork
length (Heiser, 1966). Annstrong (1970} determined that spring
smolts from Hood Bay Creek, x = 129-134 mm (fl) were considerably
smaller than autumn smolts, x • 141-146 mm (fl) during 1967, 1968
and 1969.
D. Economic Importance
Anadromous Do 11 y Varden char are an important and sought after
sport fish (Morrow, 1980).
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II. SPECIFIC HA~1TAT REQUIREMENTS
A. Upstream Migration
Adult and immature anadromous Dolly Varden char leave the Pacific
Ocean and enter various 1 a ke and non-1 a ke stream sys terns from
July through December. Various studies have indicated that lake
and non-lake streams may support spawning anadromcus Dolly Varden
char although almost all fish {both spawning and non-spawning)
entering non-lake streams such as Hood Bay Creek, Alaska, leave
these streams and enter streams with lakes where they overwinter
(Armstrong, 1963; Armstrong, 1965b; Armstrong and Winslow, 1968;
Armstrong and Kissner, 1969). This section will discuss the
upstream migration of anadromous fish in non-lake streams.
1. Stream Flow
Adu 1 t and immature anadromous Do 11 y Varden may migrate up
non-1 alc:e streams tor varying distances from July through
November. Peak numers of fish have been reported during
spates in August and Septemer in Hood Bay Creek (Armstrong,
1967; Armstrong and W~nslow, 1968; Armstrong and Kissner,
1969).
Peak numbers of fish have been recorded entering selected
streams during periods of high water in August and September
( Anns trong and Wins 1 ow, 1968 i Armstrong, 1969) • Upstream
fish migration may be hindered by high current velocities
resulting from rapids and culverts. Low flows and shallow
water depths could also prevent upstream fish passage.
2. Water Temperature
Water temperatures coinciding with the conmencement, peak
and termination of the inmigr~tion of anadromcus Dolly
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Varden in Hood Bay Creek. Alaska during 1967, 1968 and 1969
were about 4.4° to 12.6°C, 6.1° to 11.1°C and 3,3° to 4.4°C
(Armstrong, 1967; Armstrong and Winslow, 1968; Armstrong and
Kissner, 1969). Water temperatures at the end of the
inmigration are slightly lower than those during the
beginning.
3. Light
Most Dolly Varden move upstream in Hood Bay Creek at night
(Armstrong and Kissner, 1969).
4. Currer.t Velocity
No information regarding the upstream swirrming ability of
lnadromous Dolly ~arden was found in the literature.
B. Spawning
1. Current Velocity
There are on 1 y 1 i m1ted observations of anadromous Dolly
Varden spawning habitat with respect to current velocity.
Bla:kett (1968) reported fish spawning fn a reach of Hood
Bay Creek, Alaska which had current velocities ranging from
0.3 to 1.2 m/sec (1.0-3.8 ft/sec). Blackett and Annstrong
(1965} noted fish (presumably spawning) in a reach of Rodman
Creek, southeast Alaska, with a current velocity estimated
to be about 0.63 m/sec (2 ft/sec).
2. Substrate
Anadromous Dolly Varden typically spawn in small gravels.
Blackett (1968) found fish spawning primarily in small
gravels, 6 to 50 mm in diameter in Hood Bay Creek. Blackett
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and Armstrong (1965) observed what appeared to be fish
spawning in Rodman Creek, southeast Alaska, in substrate
composed of "2 5t sand and 751: rubb 1 e. •• No substrate
classification scheme was presented. Spawning anadromous
fish use gravels ranging from 2 to 32 mm in diameter in the
Terror and Kizhuyak River~. KodiaK Island, Alaska (Wilson et
al., 1981).
3. Water Depth
The relationship between spawning habitat and water depth is
speculative. Blackett {1968) observed spawning fish at
depths exceeding 0.3 m whereas Blackett and Armstrong (1965)
noted probable spawning activity in a different southeastern
Alaska stream in water depths of about 1.25 m.
4. Cover
There is little available infonnation on tne influence of
stream cover on selection of spawning habitat, however,
cover may be a requirement.
C. Inmigrant Migration to Overwintering Areas
Inmigrant, immature Dolly Varden in non-lake streams such as Hood
Bay Creek, A, 1 aska usually 1 eave within severa 1 weeks; however,
spawners may remain for up to three months (Armstrong, 1967).
Peri ods of h 1 gh water may enha nee outm1 gra t ion of 1 nma tu re and
spawned-out Dolly Varden fn Hood Bay Cree~ (Armstrong and
Kissner, 196g).
Inmature and spawned-out adult Dolly Varden inmigrate to -
overwintering areas of Eva Lake at different times. Immature
individuals entered primarily during July, August and September,
and spawned-out adults entered in late October and November
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(Blackett and Armstrong, 1965). Most fish passed upstream during
periods of darkness.
D. Inmigrant Overwintering Areas
Lakes, ~nclud1ng turbid glacial lakes support overwintering
populations of juvenile and adult Dolly Yarde., char (Amstrong.
1965b; Schmidt. Robards and McHugh, 1973). The char typically
remain in Eva Lake from December through mid-March (Armstrong and
Blackett. 1965}. Their distribution within lakes may be quite
restricted (Amstrong 7 1965b; Schmidt et al., 1973).
E. Egg and Alevin Development
1. Water Temperature
Blackett (1968) determined that anadromous Dolly Varden eggs
hatched in 129 days with 675 thenna 1 units. Absorption of
the yolk sac was completed about 65 days later when water
temperatures were 2.2° to 2.8°C,
F. Summer Juvenile Rearing
1. Water Depth
Recent 1 y emerged Do 11 y Varden char ty pi ca 11 y occupy
extremely shallow rivulets. tributaries or streamside
margins {Blackett, 1968; Armstrong and Elliott. 1972). They
may occupy deeper stream reaches as they grow (Armstrong and
Ell i ot. 1972).
2. Current Velocity
Recently emerged Dolly Varden char may occupy extremely
shallow, low current velocity stream reaches (Blackett,
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1968; Annstrong and Elliot, 1972). Minnow traps captured
juvenile, pre-smolt Dolly Varden char in Hood Bay Creek from
a variety of lotic habitat types. Ti'te largest juvenile
char, about 150 rrm fork 1 ength, were captured in r1 ffl es
(Annstrong and Elliott. 1972).
3. Instream Cover
Juvenile Dolly Varden have been obsei"Ved 1n proximity to
various forms of instream cover including root balls, trees
and undercut banks (Anmstrong and Elliott, 1972; Wilson, et
al., 1981).
4. Substrate
Recently emerged fry have been found along stream margins
with varying sizes of substr!te, shallow depths and very low
current velocities (Blackett, 1968; Annstrong and Elliott,
1972). Heiser (1966) noted Juvenile Dolly Varden char
occupying "gravel or muddy substrata" within tributaries of
Eva Lake, Alaska.
Deposition of significant amounts of fine sediment fn
streams with limited flushing ab1li ties could reduce the
quality of Juvenile anadromaus Dclly Varden rearing habitat.
Laboratory stream channels containing unimbedded rubble
(0.30 m in diameter) consistently supported more Juvenile
steel head trout, Salmo gai rdneri (Richardson), and chinook
salmon, Oncorhynchus tsha!ftscha (Walbaum). than stream
channels containing imbedded rubble and with water
temperatures exceeding 511 C (Bjornn, Brusven, Molnau,
Milligan, Klampt, Chacho and Schaye, 1977). Bjornn et al. ·
(1977) ascribed the reductions in fish abundance in the
channels with imbedded rubble to l~~s of intersticial cover.
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G. Juvenile Migration to Overwintering Areas
1. Water Temperatu~
Juvenile ( pre-smolt) Dolly Varden char have been reported
moving upstream in Starrigavin Creek when water temperatures
decreased from about 7° to 49 C. The f1 sh entered Spring
Pond Creek, a spring-fed tributary characterized by more
stable water temp~~atures and flows (Elliott and Reed, 1974;
Elliott, 1975). No fish movement into Spring Pond Creek was
noted after Starrigavin Creek water temperatures decreased
below 4°C.
2. Stream Flow
Freshets within Starrigavin Creek, Alaska appeared to
stimulate immigration of juvenile Dolly Varden char to
Sprfng Pond Creek (Elliott and Reed, 1974; Elliott, 197.5)
until water temperatures decreased below 4°C.
H. Juvenile Overwintering Areas
1. Water Temperature
Fry overwintering areas fn southeast Alaskan streams usually
llave relatively warm water temperatures. Spring Pond Creek,
a tributary of Starrigavin Creek, supports overwintering
Dolly Varden char. This stream usually has winter water
temperatures at or above l.0°C (Elliott and Reed, 1974;
Elliott, 1975).
The headwater reaches of Hood Bay Creek a 1 so appear to
support overwintering juvenile Dolly Varden char. Armstrong {
and Elliott ( 1972) found the greatest nuntlers of j uveni 1 e
fish during March and April in headwater reaches of Hood Bay
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2.
Creek where water temperatures were 5° to 6.l°C. Downstream
reaches were characterized by water temperatures of 3.9°C.
Elliott and Reed (1974} noted that Dolly Varden fry hid
among substrate interst~ces as water temperatures in Spring
Pond Creek decreased to 4°C to 2°C. When water t~peratures
rose above 2°C fn March. fish began to move dbout the
stream.
Stream Flow
Stable winter flow conditions sur1 as those found in Spring
Pond Creek.. are probably very important tc winter survival
(Elliott and Reed, 1974; E11~ott, 1975).
3. Substrate
Debris and large substrate material may enhance the quality
of fish o~erwintering areas. Elliott and Reed {1974) noted
juvenile Dolly Varden char burrowing into logging debris and
slash when water temperatures declined to 4°C or below in
Sp1•ing Pond Creek., Alaska.
Deposition of fine sediments in streams with limited
sediment flushing capabilities could imbed substrate
material and significantly reduce the available
overwintering habitat for juvenile Dolly Varden char.
Experiments of overwinter habitat selection by juvenile
chi nook. and coho salmon and stee 1 head and cutthroat trout
(Salmo clarti (Richardson)) at water temperatures less than
5°C indicate that substrate (15-30 em in diameter) with
inter·stices devoid of fine sediment consistently supported
more fish than substrate imbedded with fine sediment
(Bustard, 1973; ~jornn et al., 1977).
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I. Juvenile Migration to Summer Rearing Areas
1. Water Temperature
Juvenile (pre-smelt) Dolly Varden char were found to
emigrate from Spring Pond Creek, a spring-fed stream
inhabited by ovel""ff'intering juvenile and adult resident and
anadrormus Dolly Varden, to Starrigavin Creek when water
tem~~ratures rose to 4° to soc 1n April 1g74 (Elliott.
lg75).
2. Stream FlOW
E11 i ott (1975) suggested that fl aods in Spring Pond Creek
depressed the downstream movement of juvenile Dolly Varden.
Fish emigration increased when flows decreased.
J. Inmigrant Migration to Sea
1. Water Temp~rature
Most i11111ature and mature Dolly Varden char (nat including
smolts) emigrated from Eva Lake Creek, Alaska shortly after
ice-breakup. Water temperatures ranged from 4. 4° to 6. 7°C
(Annstrong. 1965b).
2. Stream Flow
Although ice-breakup in Eva Lake appeared to strongly
influence fish migration to the sea, peak numbers of
emigrants moved downstream auri ng flood states (Armstrong,
1965b).
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Most Dolly Varden char emigrated from Eva Lake during
darl<ness. During the height of the migration individuals
were detected moving downstream during both night and day
(Armstrong, 1965b).
K. Smolt Migration to Sea
1. Water Temperatures
Water temperatures coinciding with the initiation, peak and
the near-end of the spring Dolly Varden smolt migration in
Hood Bay Cre~,;. Alaska were 3", 5" and *'CIC, respectively,
during 1967, 1968 and 1969 (Armstrong, 1970).
Water temperatures at the beginning, pea I< and end of the
smolt migration during 1962 and 1963 were 3°, SCI and 8°C in
Eva Lal<e, and 6" and 1D°C in Eva Creek (Armstrong. 1970).
These values are somewhat similar to water temperatures
during the spring smo 1 t outmi gration from Hood Bay Creek,
Alaska.
Dolly Varden smolt stopped migration i" mid-June 1957 in the
Anchor River, Alas .~a when the water temperature reached
13.3°C (Allin, 1957).
The autumn smolt mig ation in Hood Bay Creek, Alaska began
when water temperatures were 8°C and ended when water
temperatures were 6~C during 1967, 1g68 and 1969 (Armstrong,
1970).
2. Stream Flow
Fi ood s apparent 1 y influenced the timing of the spring and
autumn Dolly Varden cnar smolt migration in Hood Bay Creek,
A 1 ask a. Peak nunilers of smo 1 ts migrated downstream during
periods of high water (Armstrong, 1970). Smelts have also
been noted emigrating from Mendenhall Lake (near Juneau)
during the spring and early SUIIIIEr. Peak migration~ often
coincide with freshets (Bethers, 1974).
Most smo 1 t migrate downstream in Hood Bay and Eve Creeks
during darkness, although the peak of smolt emigration in
Eva Creek occurred during both night and day (Armstrong,
1970).
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~!I. CONCEPTUAL SUIIABILITY INDEX CURVES
Habitat suitability index curves were not constructed for anadromous Dolly
Varden char. There were limited data relating the various Dolly Varden
life stages to the physical and chemical characteristics of the habitats.
When data were available, they were often not in a fo~ which could be used
to construct habitat suitability curves.
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IV. DEFICIENC:ES IN DATA BASE
A 1 imited number of 1 .westigations indicate that juvenile anaaromous
Dolly 'lar"den char move to spring-fed reaches of streams with
relatively warm water temperatures during the fall and leave these
a rea s the fa 11 owing spring, as e•Ji denced by baited minnow trap samp 1 es
from Hood Bay Cree~ (BlacKett. 1968; Armstrong and Elliott, 1972} and
by weir sam~ling in the Starrigavin Creek watershed (Elliott and Reed,
1914; Elliott, 1975, 1976 and 1977). The Starrigavin watershed was
affected by timber hanest and deposition and removal of logging
debris.
The relationships between upstream swimming capabilities of juvenile
and adult anadromous Dolly Varden and current velocity, water
temperatul"'e, water depth and str"eam gradient have not been
sufficiently investigated. Excessive stream gradient and high current
velocities associated with natural stream features or culverts could
impede migration of juvenile and adult fish to surrmer and winter
rearing and spawning areas. The upstream swinmi ng perlonnance of
anadromous Dolly Varden is probably influenced by the above factors as
well as fish size, spawning condition and, possibly, sex.
Loti c habitat se 1 ect ion by spawning anadromou s Do 11 y Varden char is
probably col iectively influenced by current velocity, water depth,
substra~e composition and imbeddedness and instream and bankside
vegetation. Few studies have obJectively described the above lotic
habitat conditions. If a specific area of a stream is characterized
by Jne unfavorable spawning habitat feature, such as excessive current
velocity or unsatisfactory substrate composition, that par"ticulal"' area
of stream will not be selected by spawning fish despite other
favorable habitat conditions.
Habitat conditions available for Dolly Varden spawning sites influence
selection of spawning areas, a1though, methods of objectively
cescribing this habitat have varied. ~or example. within a
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hypothe~i ca 1 stream9 stream reach A may suppo.-t one pair of spawning
fi sM and reach B, four pairs . These two reaches may contain equa 1
amounts of ~spawning habitat" as defined by current velocity, water
dept~ and substrate composition and imbeddedness. The difference
between the two reaches may be that "spawning habitat" within stream
reach A was concentrated in one a rea Ind "spawning habitat 11 within
stream reach B was scattered among relatively la.-ge substrate and fast
water. The non-contiguous distribution of spawning habitat in stream
reach B probably allows more fish to spawn because of increased cover
and v 1 sua 1 i so 1 at ion. This ex amp 1 e i 11 ust rates the need to examine
entf.-e stream reaches to better unde.-stand selection of spawning
habitat.
The influence of dissolved oxygen levels, rates of intrag.-avel flow,
sediment compositions and water temperatures on the sur~ival and
development of anadromous Dolly Varden cha.-is not understood.
Numencus research focusing on the eggs and alevins of other salmonids
indicate that the physical and chemical factors exert a substantial
and often interactive influence on the survival and fitness on the
development stages.
Adult and inmature (post-smolt) Dolly Varden typically oventi'lte.-in
lakes, although fish have been found in spring-fed reaches of streams
in southeast Alaska. The impo.-tance of glacial or glacial-influenced
lakes and streams has not been adequately examined. The relationships
between oventintering habitat and water depth, current velocity,
substrate composition, water temperature and other variables has not
been detenni ned. The di stri but ion of overwi nter1 ng Dolly V ~ rden in
se 1 ec ted 1 ake s may be quite rest ri c~:ed, although phy sica 1 and chemica 1
factors which may limit the fish's distribution are not known.
Juvenile (pre-smelt; Dolly Varden have been documented moving to
spring-fee tri but aries and burrowing into 1 oggi ng debris when water
temperatu.-es app.-oac!'led 4°C. There were no .. eferences found in the
literatu.-e of juvenile fish using mainstem reaches of rapid-nJnoff
streams for overwintering habitat. The apparent affinity of yoy ar.d
older juve~ile fish to rapid run-off streams during the summer
indicates that. this type of a rea cou 1 d be used for oven;-i nteri ng
habitat.
There is 1 itt 1 e ava i 1 ab 1 e i nfonna ti on concerning the 1 otic sumner
micro-habitat selection by juvenile (pre-smolt) anadromous Dolly
\/arden char with respect to various physical and chemical lotic
habitat variables, food availability and the presence of other fishes.
Some investigators nave attempted to describe juvenile fish habitat
quantitatively by bankside observation, electro-shocking and baited
minnow traps with varying degrees of success. Apparently snorkeling
has not been used for fish observation in clearwater streams.
Snorkeling has been shown to be a valuable fish observation technique
in clearwater streams of the Pacific Northwest and the midwest
(Everest and Chapman, 1972; Fausch, 1978).
Some wor-k has focused on the feedin9 habits of juvenile (pre-smolt)
fish occupying mainstem versus stream margins but no drift or benthos
sampl 1ng was done to fonnulate "forage ratios." Few observations of
the feeding behavior cf juvenile Dolly Varden char have occurred in
streams except for yoy fhh in very sha 11 ow. 1 ow current ve 1 oci ty
areas along stream margins.
So1111i! observa ~ions of j uveni 1 e coho sa 1 mon and Do 11 y Varden char-have
occurred in aquariums and stream$. The behavior of juvenile Dolly
Varden char and other salmoni~s occurring in the same regions has not
been studied adequately.
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v. RECOMMENDATIONS AND FURTHER STUDIES
Studies shou 1 d be designed and conducted to detenni ne the su rv iva 1 ,
movements and behavior of all life stages of anadromous Dolly Varden
char with respect to physical, chemical and biological habitat
components within selected pristine Alaska drainages. The above
relationships should be examined thoroughly both within the drainage
and by supplemental laboratory and field studies prior to the
occurrence of any land use activities which could modify the habitat.
Severa 1 years of study caul d be required to meet this objective.
Investigations should continue during and after land use activities to
adequately monitor the fish life stage-habitat relationships. ~uch
research and supp 1 ementa 1 1 a bora tory and fie 1 d stud iPS cou 1 d provide
land managers with needed information to protect and enhance
anadromous Dolly Varden habitat.
Field and laboratory studies should be designed and conducted to
determine the upstream swimming capability of immature (pre and
post-smelts) and gravid and spawned-out adult anadromous Dolly Varden
char in relation to current velocity, water depth, water temperature,
stream gradient and length of potential migration barriers. Studies
could be similar to those described by iolacPhee and Watts (1976) for
testing Arctic grayling swimming performance. Results of these tests
could be used f~r determining the best methods for installing culverts
to allow fish migration and to designs of culverts and other fish
passage facilities.
Str.dies should be designed and conducted to evaluate the influence of
~ater temperature, dissolved oxygen, rate of intragravel flow,
~uhstrate composition and possibly other physical and chemical habitat
.,~,.iables on the survival and development of anadromous Dolly Varden
:har eggs .tnd alevins and the fitness and survival of emergent fry.
Controlled environmental laboratory tests should complement field
studies.
-29-
Standardized methods should be developed and evaluated to objectively
describe current velocity, water depth, substrate composition and
irmeddedness, instream and bankside cover !nd water temperatures at
anadromous Dolly Varden spawning sites. The above lotic habitat data
collected at a nurmer of r~dds within a stream or stream reach could
be examined by frequency analysis for each lotic habitat component.
These frequency analyses would help describe lotic habitat selection
by spawning fish, in relation to current velocity, water depth,
substrate composition and imbeddedness and possibly other lotic
habitat .ar1ables. The frequency analyses would not detennine fish
spawning habitat preferences because streams and stream reaches are
characterized by a finite combination of acceptable habitat variables.
Readers are urged to consult Appendix III of the Terror Ri•ter, Alaslta
In stream Flow Report by Wilson et al. (1981) which discusses fish
spa~ning habitat selection and the assumptions associated with habitat
suitability curve construction.
Standardized methods should be developed and refined to evaluate
current velocity, water depth, substrate composition and imbeddedness
and instream and bankside cover at anadromous char spawning sites to
better understand nabitat selection within individual streams or
stream reaches. For example, lotic habitat types could be
characterized by current velocity, water depth, instream !nd banltside
cover conditions within those ranges measured at redds. Measurements
of substr!lte composition and imbeddedness which were net used for
spawning habitat by anadromous C"olly varden char could help us to
J
better understand spawning habitat selection of this fish in various
streams.
Weirs should be used to monitor juvenile (pre-smolt) anadromous Dolly
Varden char movements in relation to streamfl~. water temperature and
other physical and chemical habitat variables in small, intennittent
and larger streams within a drainage. Snork.ling and minnow trapping
could supplement sampling with weirs.
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Studies should be designed and conducted to describe sunmer habitat
se 1 ect ion by j uven i 1 e (pre-smo 1 t) anadromou s Do 1 1 y Varden char with
respect to a va~iety of p~ysical and chemical habitat variables, food
avai 1 abi1 i ty and the presence of other fish. Investigations using
snorkling for fish observation should be conducted in clea~ater
st~ams using techniques simila~ to Eve~st and Chapman (1972) and
Fa us ch {1978) . Fish ho 1 ding positions shou 1 d be cha ~acteri zed by
wate~ depth, distance to streambed, lighting, substrate coiii)Josition
and imbeddedness, instream and bankside cover, current velocity and
proximity to other fish, including cha~ and other species. These
investigations would complement fish movement studies along streams
supporting weirs.
Studies of fish feeding behavio~ coupled with benthos and drift
saiii)Jlirg and fish gut analysis should be conducted to bette~
understand the feeding habits and apoarent affinity of Dolly Varden
char to the substrate.
More i nves ti ga ti ens shou 1 d occur to detect and cha ~acteri ze
ove~intering habitat selection by various ages of .iuvenile
(pre-smolt) anadr·omous Dolly Varden char with ~spect to ~:i~er deeth,
current velocity, water temperature, overhanging vegetation,
banks, substrate material and p~oximity to stream
Identification of overwintering fish habitat is ~equired
undercut
margins.
for the
p~otection of this fish species. Various land use activities could
significantly reduce the quality of this habitat {Busta~d, 1973).
Habitat enhancement efforts should be formulated with an understanding
of what constitutes good ove~intering habitat fo~ various ages of
Dolly Varden char.
Lr Jo ra tory and fie 1 d tests , somewhat simi 1 a~ to those conducted by
Bustard (1973), should be jesigned and conducted to complement studies
of ~:~~~ distribution and behavior of overwintering anadromous Dolly
va~den in selected streams. These tests should provide juvenile fish
with a cent i nuum of overwinter habitat types from no cover to tot a 1
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r i pari an cover. u n imbedded to tot a 1 1 y imbedded substrate of sand to
large cobble substrate and a variety of water temperatures. Juvenile
fish of various ages should be tested because fish size may influence
overwinter habitat selection.
Studies should be designed and conducted to determine the presence of
overwintering immature {post-smolt) and adult anadromous Dolly Varden
char in clearwater and glacial lakes and streams. The distribution of
overwintering fish in lakes can be quite limited. Studies of lotte
overwintering areas with respect to various physical and chemical
habitat conditions should be conducted to explain h'!bitat selection
criteria and to predict fish overwintering areas by the character of
the habitat.
Radio telemetry should be considered as a viable technique to monitor
movements of immature and adult anadromous Dolly Varden char in
overwintering areas which are difficult to sample by gillnet or other
methods.
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V!. LITERATURE CITED
Alde~dice, D., W. Wickett and J. Brett. 1958. Some effects of temporary
low dissolved oxygen levels on Pacific salmon eggs. J. Fish. Res.
Boa~d Can. 15(2):229-250.
A11in, R. 1954. St~eam survey of Ancho~ Rive~. Game Fish Investigations
of Alaska. Quarterly Progress Report, Federal Aid in Fish Restoration
Proje~t F-1-R-4. Alaska Game Commission 4:47-66.
Alt, K. 1978. Invento~y and cataloging of sport fish and sport fish
wate~s of western Alaska. Alaska Dept. of Fish and Game. Fede~al Aid
in Fish Restoration, Annua 1 Progress Report, 1977-1978.
19{G-I -P) :36··60.
Armstrong, R. 1963. Investigations of anadromous Dolly Varden populations
1n the Lake Eva-Hanus Bay drainages, southeastern Alaska. Alaska
Dept. of Fish and Game. Federal Aid in Fish Resto~ation, Annual
Progress Report, 1962-1963. Project F-5-R-4:78-122.
Armstrong, R. 1965a. Some feeding habits of the anadromous Dolly Varden,
Salvelinus rnalma (\llalbaum) in southeastern Alaska. AlasKa Dept. of
Fihs and Game Informational Leaflet 51:1-27.
Armstrong, R. 1965b. Some migratory habits of the anadromous Dolly Varden,
Salvel inus mal rna (Walbaum) in southeastern Alaska. Alaska Dept. of
Fish and Game Research Report 3:1-26.
Armstrong, R. 1967. Investigation of anadromous D~lly Vdrden populations
in the Hood Bay drainages, southeastern Alaska. Alaska Dept. of Fish
and Game. Federal Aid in Fish Restoration, annua 1 Progress Report,
1966. Project F-5-R-8:33-56.
Armstrong, R. 1970. Age, food and mig~ation of Dolly Varden smolts in
southeastern Alaska. J. =ish. Res. Bo~rd Can. 27:991-1004.
-33-
A~trong, R. lg74. Migration of anadromaus Dolly Varden {Salvelinus
~lma) in southeastern Alaska. J. Fish. ~es. Board Can. J1:435-444.
Armstrong, Rand S. Elliott. 1g12. A study of Dolly Varden in Alaska.
Alaska Dept. of Fish and Game. Federal Aid in Fish Restoratio1,
Annual Progress Report, 19~1-1g72. Project F-g-4-13:1-34.
Annstrong, R. and P. K:issner. 1g6g, Investigations of anadromous Dally
Varden populations i~ the Hood Bay. southeastern Alaska. Alaska De~t.
of Fish and Game. Federal Aid in Fish Restoration, Annudl Progn~ss
Report, 1g68-lg6g. Project F-5-R-10:45-g2.
Armstrong, R. and R. Winslow. 1gsa. Investigation of anadromous Dolly
Varden popu 1 at ions in the Hood Bay drainages , southeastern A 1 as <a.
Alaska Dept. of Fish and Game. Federal Aid in Fish RestoratiJn,
Annua1 Progress Report, lg67-lg68. Project F-5-R-9:45-80.
Sethers, M. lg74. Mendenhall Lakes salmon rearing facility. Alas5a Delt.
Fish and Game. Feaeral Aid in Fish Restoration, Annual Report of
Progress lg73-1g74, Project F-9-6. l5(AFS-4J).
Bjornn, T., H. Brusven, M. Molnau, J. Miiligan, R. Klampt, E. Chacho and C.
Schaye. 1g77. Transport of granitic 5~diment in streams and its
effects on insects and fish. For Wild 1 i fe and Range Exp. , S tr .. ,
Completion Rep. Water Resour. Res. Inst. Project B-036-1D.~.
University of Idaho, Moscow. 43 p.
Blackett, R. lg68. Spawning behavior, fecundi~y and early life history cf
anadromous Dolly Varden, Salvelinus malma (Walbaum) in southeastern
· Alaska. Alaska Dept. of Fish and Game, Research Report 6:1-85.
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3~acket-:. ~. and ~. Arms~rong. 1965. Investigations of anadromoiJS Dolly
'/arden ;:-..~ulations in the Lake Eva-Hanus Bay
~iaska. ~lask.a Deot. of Fish and Game.
=tes tarat ion, Annual Progress Reoort,
F-5-R-6:23-56.
drainages, so•Jtheastern
Federal Aid in Fish
1964-1965. Project
Bustard, D. 1973. Some aspec!s of the winter ecology of juvenile
salmonids with reference to possible hatitat alteration by logging in
Carnation Creek, Vancouver Island. M.S. thesis. Univer-5·1ty of
British Columbia, Vancouver, B.C. 85 p.
Chapman, D. ~966. Food and space as regulators of saimonid population~ in
streams. Am. Nat. 100:345-357.
Chapman, 0. and T. Sjornn. 1569. Distribution of salmonids in streams,
with speci a 1 reference to food and feeding. H. R. MacMillan Lectures
in Fisheries. Symposium on salroon and trout in streams. Unher-5ity
of British Columbia. p. 153-176.
Coble, 0. 1961. Influence of water exchange and dissolved oxygen in redds
on surviv~l of steelhead t~ut embryos. Trans. Am. Fisn. Soc.
90( 4): 469-471.
Conte, F. and H. Wagner. 1965. Development of osrootic and ionic
regulation in juvenile steelhead trout, Salroo gairdner1. Comp.
Biochem. Physiol. 14:603-620.
£1liott, S. 1975. Ecology of rearing fish. Alaska Dept. of Fish and
Game. Federal Aid in Fish Restoration, Annual Progress Report,
1974-1975. Project F-9-7(D-I-B):23-46.
Elliott, S. 1976. Ecology of rearing fish. Alaska Dept. of Fish and
Game. Federal Aid in Fish Restoration. ~nnual PrQ9ress Re~ort,
1975-1975. Project F-9-8(0-I-8):21-44.
-35-
E11iott, S. 1977. E~oloqy of reari~g fish. Alaska Dept. of Fish ~nd
Game. Feder~l Aid in Fish Restoration, Annual Progress Report.
1976·1977. Project F·9-9-(0-1-B):25-36.
elliott, S. and~. Dinneford. 1975. Establishment of guidelines for
protection o-f the sport fis~ resources during logglng operations.
t.laska :lept. of Fi~h and Game. Federal Aid in Fish Restoration,
Annual Report of Progr~ss, 1g74-1975, Project F-9-7, 16(D-1-A}. 1-22.
Elliott, S. and R. Reed. 1g74. ~cology of rearing fish. Alaska Dept. of
Fish and Game. Federal Aid in Fish Restoration, Annual Prorress
Report, 1973-1974. Project s-g-6(0-I-B):g-43.
Everest, F. H. and 0. w. Chapman. 1972. Habitat selection and spatial ~Y
juvenile chi nook salmon and s tee 1 head trout in two 'daho s t ree...ns. J.
Fi$h. Res. Board Can. 29:g1-100.
Fausch, K. 0. 1978. Competition between brook and rainbow trout for
resting positions in a stream. M.S. thesis. ~ichigan State
University, East Lansing, ~1chigan. 100 p.
Hartmann, G. 1968. G~n rate and distribution of some fishes in the
Chill tvack, South Alouette and Salmon Rivers. B.C. Fish and Wildl.
Branch, Mgmt. Publ. No. 11, 33 p.
Heiser, D. 1g66. Age and growth of anadromous Doliy Varden char,
Salvelinus .alma (Walba~). in Eva Creek, Baranof Island. southeastern
Alaska. Alaska Dept. of Fish and Game, Research Report 5:1-2g.
Koski, K. 1966. The survival of coho salmon (Oncorhynchus k1sutr.h1 from
egg deposition to emer-ge11ce fn th~t! Oregon coa~tal streams. M.S.
thesis. Oregon State Unfver~ity, Corvalis, Oregon. 34 p.
-36 ..
Legget, J. 1959. 7he reprod~.o~·ive biology of t he Dolly Varden char
(Salvelinus :nalma (\olalbaum)}. "1.S . thesis . Unh1ersity Clf Vic~oria,
Victoria, B.C. lll p.
•l..:'le'il. "· :966. Affect of the spa~o~nir.g bee environment on reproduction
of pink and chu m salrtWJn. U.S. Fish and 'Jildl. Serv . Fish. Bull.
65:!95-5l3 .
~cNeil, ~. and W. Ahnell . 1964 . Success of pink salmon spawning relative
to size of spa~o~ning bed materials. U.S. Fish and Wildl. Serv. Scec .
Sr:i. Re. Fish. No. 469. 15 p.
"1acPhee, C. and F. Watts. 1976 .
in highway culverts. Final
Anchorage. Alaska. Contract
Swimming perfonr~nce of Arctic grayling
Report to U.S. Fish and Wildlife Service,
14-16-001-5207. 41 p.
Morrow, J. 1980. The fresh~o~ater fishes of Alaska. Alaska Northwest Pub.
Co ., Anchorage, Alaska. 248 p.
~e~d. R. and R. Armstrong.
area . Alaska Dept.
1971. Dolly Varden soort fishery -Juneau
of Fish and Game. Federal Aid in Fish
~estoration, Annual Progress Report, 1970-1971.
12:1-105.
Project F -9-3.
RP.imers, N. 1957. Some aspects of the relation between stream foods and
trout survival. Calif . Fish and Game. 43(1):4J-6g.
Reiser, D. and T. Bjornn. lg79. Habitat requirements of anadromous
salmonids -lnfluencP. of forest and rangeland management on anadrornous
fish habitat in t~e western United States and Canada . U.S.O.A. Forest
Ser"vice. Gen. Tech. Rep. PNW-96 . Pac1 fie Northwest FQrest and Range
Exp. Stat. Por"tland. Oregon. 54 p.
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Sc!"'midt, A., S. Qob:H·ds and I'll. McHugh. 1973. Inventory and cataioging cf
sport fish and sport fish waters in southeast Alaska. Alaska Deot. of
Fish ilnd Gilme. Jederal Aid in Fish Restoration, Annual Progress
Reoort, 1972-1973. Project F-9-5 1 14:1-62.
Scott, '.J. and E. Crossma.,. 1973. Fresh_,ater fishes C'f Canada. Bull. 184.
Ffsh. =<e~. 3oar1 of Can. 1 Ottawa. 9Ei6 p.
Sheridan, W. 1962. Waterflow through a salmon spawning riffle in
southeastern Alaska. u.~. Fish and Wil dl. Serv. Spec. Sci. Rep. Fish
No. 407. 22 p.
Silver, S. • C. ~a r-r-en and P. Doudoroff. 1963. Dissolved oxygen
requirements of developing steelhead trout and chinook salmon embryos
at different water velocities. irans. Am. Fish. Soc. 92{4):327-343.
Vaux 1 W. A. 1962. Interchange of str~am and intragravel water in a salmon
spawning riffle. U.S. Fish and Wildl. Serv. Spec. Sci. Rep. Fish No.
405. 11 p.
Wickett, W. 1958. Review of certain environ~~ntal factors affecting the
production of pink and chum salmon. J. Fish. ~es. Board Can.
15(5): 1103-1126.
Wic~ham, G. 1967. Physical microhabitat of trout. M.S. thesis. Colorado
State Univer-sity, For-t Collins. 42 p.
Wilson. w., E. Trihey, J. Baldrige, C. Evans. J. Thiele and D. Trudgen.
1981. A:1 assessment of environmental effects of con~truction and
operation of th~ proposed Terror Lake hydroelectric facility, Kodiak,
Alaska. Instream flo-, studies final report. Arctic Environmental
Information and Da:~ Center. University of Alaska, Anchor-age, Alaska.
419 p.
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STLAWRENCE
~AND
0
MAIN STUDY SITES
IIJ HOOD BAY CREEK
[i) RODMAN CREEK
00 EVA LAKE
~ STARRIGAVIN CREEK
ALEUTIAN ISLANDS
0 •
..,::?~
" 0 ~Jt:/)'
DISTRIBUTION OF DOLLY VARDEN -SAL VEL/NUS MALMA (WALBAUM)
FROM SCOTT AND CROSSMAN 1973, AND MAIN STUDY SITES.
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FRESHWATER HABITAT
RELATIONSHIPS
THREESPINE STICKLEBACK -GASTEROSTEUS ACULEATUS
AlASKA DEPARTMENT OF FISH & GAME
HABITAT PROTECTION SECTION
RESOURCE ASSESSMENT BRANCH
APRIL, 1981
FRESHWATER HABI~'AT RELATIONSHIPS
THREESPINE STICKLEBACK (GASTEROSTEUS ACULEATUS)
By
Stephen S. Hale
Alaska Department of Fish and Game
Habitat Division
Resource Assessment Branch
570 West 53rd Street
Anchorage, Alaska 99502
May 1981
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AC~NOWLEDGEMENTS
Many people from the Alaska Department of Fish and Game and from the Auke
Bay Fisheries Laboratory of the National Marine Fisheries Service freely
gave their time and assistance when contacted about this project and it is
a p~easure to thank them and fishery biologists from other agencies,
especially those who provided unpublished data and observations from their
own work. The librarians of the Alaska Resources Library and the U.S. Fish
and Wildlife Service were of great help.
This project was funded by the U.S. Fish and Wildlife Service, Western
Energy and Land Use Team, Habitat Evaluation Proce~ure Group, Fort Collins,
Colorado. Contract ~o. 14-16-0009-79-119.
TA8LE OF CONTENTS
THREESPINE STICKLEBACK
Page
I. INTRODUCTION 1
A, Purpose 1
B. Distribution 2
c. Life History Sunmary 2
1. Size and Age 4
2. Maturity and ~eproduction 6
3. Feeding and Competition 9
4. Oistrihution and Behavior 13
[\, Ecological and Economic Importance 15
II. SPECIFIC HABITAT RELATIONSHIPS 16
A. Spawning 16
1. Temperature 16
2. Water Depth 16
3. Substrate 17
4. Aquatic Vegetation 17
5. Light 18
6. Size of Territory 19
B. Incubation 19
1. Temperature 19
2. Dissolved Oxygen 21
c. Juvenile Rearing 21
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D. .~Jults
1. Temperature
2. Water-Depth
3. Current Velocity
4, Dissolved Oxygen
5. Chemical Parameters
6. Feeding
III. CONCEPTUAL SUITABILITY INDEX CURVES
IV. DEFICIENCIES IN DATA BASE AND RECOMMENDATIONS
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27
40
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LIST OF FIGURES
1. Distribution of threespine stickleback in
Alaska (Morrow, 1980) and main study sites.
2. Conceptual model of relationship between
threespine stickleback embryos and
temperature.
3. Conceptual model of relationship between
threespi~e stickleback adults and water
temperature. current velocity, and water
depth.
4. Conceptual model of relationship between
thre~spine stickleback adults and dissolved
oxygen concentration and pH.
LIST OF TABLES
I. Data table -Spawning
II. Data table -Incubation of embryos
III. Data table-Adults
3
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I. INTRODUCTION
A. Puroose
This report comp i 1 es existing i nfonna t ion on the freshwater
habitat requirements, tolerances, and preferences of the
threespine stickleback, Gasterosteus aculeatus (L.) and provides
a data b~se for habitat evaluation procedures. The threespine
stickleback is generally regarded as a hardy species, tolerant of
a wide range of habitat conditions.
Infonnation on physiological tolerances and requirements have
been gathered from throughout the range of the species. However,
observations on actual conditions observed in oodies of water
where sticklebacks occur are generally restricted to Alaska. The
threes pine has been extensively used as a laboratory fish for
behavioral and pollution studies. Much work has been done on the
physioiogy of the species in Europe, but relathely little in
North Arneri ca. It is recognized that habitat requirements may
differ for stocks from different geographic areas, but present
data available from Alaska are insufficient to demonstrate this
within the State.
Within the Gasterosteus acul eatus comp 1 ex, three fonns are
genera 1ly recognized (McPhail and Lindsey, 1970}. There is a
partially plated freshwater fonn (called leiurus}. a heavily
plated marine fonn (called trachurus), and an intermediate fonn
(semiannatus). This report is restricted to the totally
freshwater fonn (leiurus), even though the anadromous fonns may
use freshwater habitat during the br~eding season.
This report emphasizes habitat requi re:nents, prima ri 1 y those of a
physical and chemical nature. Certain biological factors
affecting the well being of the population, such as feeding,
-1-
predation~ c~moetition, parasites. and disease, are not
comprehensively treated.
B. Distribution
The threespine 5tick1eback is widely dis":ributed in the no:-th·~rn
hemisphere in North Arne rica, Europe, and Asia. Except for
Europe, it is not found more than a few hundred kilometers f1~
the coast. On the At~ antic coast of North Arne rica, it rangi!S
from Baffin Island and the Hudson Bay area down to Chesapeate
Bay. On the Pacific ~oast of North America, it occurs fron
Alaska to Baja California. On the e~stern coast of Asia, • t
ranges from the Bering Strait south to Japan and Korea (Scott and
Crossman, 1973; Wootton, 1976).
In Alaska, the threespine stickleback occurs in all coastal ~reas
from Dixon Entrance to the Alaska Peninsula, the Aleutian Island~
and Bristol Bay (Morrow, 1980). It also occurs in the west•!rn
tip of the Seward Peninsula and on St. Lawrence Island; howe\·er,
the freshwater leiurus form is not thought to be present in the
1 a tter areas (McPha i 1 and lindsey, 1970; Wootton, 1976). ! St!e
Figure I.)
The leiurus form occurs in both lakes and streams.
C. Life History Su~ry
An excellent synthesis of threespine stickleback life history has
been presented by Wootton { 1976). Behavi ora 1 aspects of
threespine stickleback life history regarding reproduction
(spawning and incubation of eggs) have been extensively
documented and wi 11 not be dealt with in this report. Sec ti or•
II, Specific Habitat Requirements, will examine data on
tolerances and preferences for physical and chemical paramete~;
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nu, t.
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,
MAIN STUDY SITES
[i] Wood ._.,., Lallea
til ll&acll Lake. CltfQftlll Lalle
(I] a... Lale. Kartull Lalla
[!i L••• ..._ LU.e
I • '
Dlltrllautlon of Ttlreeeplne atlcklebacll itt Alaaka (Morrow, 1980)
a11d •aln atudr altea.
.. J ...
history aspects other t~an reproductive behavior for juveniles
and adults, emphasizing Alaskan studies. Much of this section
has been provided by Cannon (1981), with permission of the
author.
1. Size and Age
Cannon (1981) identified three discrete size classes of
threespine sticklebacks in lower Jean Lake on the kenai
Peninsula. Standard lengths ranged from 22~71 mm. One size
class (SL • 32-33 mm) was captured in surface tows in June
and early July; weekly abundance for this size class in the
catch decreased during this time. A smaller size class (~
• 28·29 mm) entered the catch in mid-June and rapidly
dominated the catch by mid-July. During July and August,
two size c1asses were distinguished in baited minnow traps
fished on the lake bottom. Stickleback (SL • JR-40 mm) were
taken in littoral areas; a larger size class (Sl a 51-52 mm)
was captured at depths of 19-20 m.
In Lake Nerka (Wood River lakes) Burgner (1958) observed a
trfmodality of size distributions which suggested the
presence of three age groups in early summer. Stickleback
fry did not ap~ear in catches until August and were abundant
in September. The maximum age was reached at three years or
more and in adult fish, females were larger than males. The
life history of populations inhabiting Karluk and Bare Lake
on kodiak Island was studied by Greenbank and Nelson (1959).
Lifespan was determined to be 2+ years, some individua1s
probably lived past a third winter. Young of the year were
f1rst observed in collections made in early July; this s1ze
class was reported to be very abundant and soon dominated
the catch. Mature females attained a somewhat larger size
than did males. Narver (1968) described the general life
history of threespine sticklebacks in the Chignik Lakes on
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the Alaska Peninsula. Determination of age was accomplished
by examination of length-frequency. An interpretation of
size distribution for a September sample placpd Age 0 fishes
within the 20-30 mm fork length (F.L.) size range, Age 1+
within 40-45 mm range and Age 2+ within 55-65 mm range. Fry
first appeared in tow net catches in ea r1y August; thef r
density increased rapidly. Enge 1 ( 1 971) a 1 so 1 dent iff ed
three size classes in beach seine and minnow trap catches in
Johnson. Scout. ind Bear Lakes on the Kenai Pen1 nsul a.
Length of life was estimated to be 2+ years. mean lengths
for ripe males and females in Bear Lake were 50.5 mm (F.L.}
and 49.5 mm respectively. These lengths correspond to those
found by Greenbank and Nelson for Age 2 fish. The abundance
and size of threespine found in Lake Aleknagik (Wood River
1 a kes) were ex ami ned by Rogers ( 1972). Age groups were
assigned on the basis of length frequency. Age 0 fish were
approximately 10-20 mm (SL) by the first week of August and
initially appeared in beach seine hauls in mid-July. Age 1
were 30-35 mm, Age 2 were 40-50 mm, and Age 3 were 55-65 mm.
The variations in age and size structure found in these
studie5 undoubtedly reflects differences in genetic and
environmental influences between the respective populations.
Direct comparisons are confounded by d~fferences in sampling
gear and measurement methods used. Rogers (1972)
demonstrated that in years of unfavorable growth, major
stickleback age groups showed distinctive length
frequencieSi but in years of good growth, the older year
classes overlapped.
-5-
2. Maturity ar~ Reproduction
Initial sexual maturity in the Lower Jean Lake stickleback
population probably occu~s during the second summer of life
(Cannon, 1981). Fishes thought to be Age 1 which were
captured in tows and minnow pots were sexually mature; a
sma11er size ~lass, Age 0, was not mature. Only one age
class at sexual maturity was reported by Greenbank and
Nelson (1959) and McPhail and Lindsey (1970). Rogers
(lg68), Burgner (1958), and Narver (1968) suggested that
threespfnes mature at Age 2. Carl (1g53) and Jones and
Hynes (lgso) stated that breeding occurred during the first
year of life .(Age 0). Variation in these estimates cannot
be explained with available info~tfon; however,
differences in length of season, food availability and
genetic character we~e p~obably influencing factors.
A decrease in Age 1 stickleback catch per unit effort for
surface tows in lower Jean lake during the course of the
summer and the presence of a similar ~ize class in shallow
shoreline areas suggested that there was a migration from
pelagic to littoral habitats (Cannon, 1981). Because Age 1
fish were sexua11y mature and were strongly suspected of
spawning fn littoral areas during this time, the movement
appeared to be associated with reproductive activity.
Tfnbergen ( 1952) and Narver ( 1968) described a spawning
migration of schools of threespine stickleback into
shallows. Males then left schools to establish territories.
Baerends (1957) stated that when reproductive instinct fn
threesp1ne 1s activated, adult fish begin to migrate. These
migrations appeared to correspo"d with a reproductive
stimulus that is only satisfied when the fish reach shallow,
warm wate~s with abundant vegetation.
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Although characteristic behavior a .. sociated with breeding
was not observed by Cannon (1981) in Lower Jean Lake, the
majority of the lake population apparently spawned in the
lake. Spawning emigrations from the lake were not observed
during the summer, however, they may have occurred prior to
mid-May. Sticklebacks were not found in high densities
milling near the tnlet and outlet creek weirs nor attempting
to pass through them. Greenbank and Nelson (1959) reported
that in karluk Lake, threespine in breeding cond;tion were
observed in the lake, but nesting was not witnessed.
Various authors have noted a variety of materi~ls used in
construction of ne~ts. Ttnbergen (1952) described nests
constructed of small twigs, grass and other debrh;
Greenbank and Nels;nn (1959) and Vrat (1949} discovered nests
formerl from cemented sand grftins.
Male sticklebacks apparently reQuire th~ sight of aquatic
vegetation for the initfatiOn-of riest building ( Pellcwijk and
T f nbe rgen , 1937 , c i ted by A ron~on , 1 gs7) . f~a rver ( 1968)
noted that the occurrence of stickleback tn spawning
condition was greatest in areas where submergent flora was
plentiful. Hagen (1967) observed sticklebacks nesting in
still and standing backwaters near dense stands of aquat1c
vegetation. Hale sticklebacks exhibiting breeding colors
an~ females with d1stenderl abdomens were observed in ~r near
thick patches of Chara .!!· fn Lower Jean Lake (Cannon,
1981).
Egg production in European populations has been estimat~d to
b~ 100-200 eggs per spawning (Assen, 1967; ~ootton, 1973 b).
McPhail and Lindsey, 1970) reported that in populations from
the Pacific Northwest, gravid females will often spawn
sever!l times per season and l"Y 50 to 200 eggs ~t a time.
The number of eggs per spawning and the ~umber of spawnings
-7-
per season have been shown to be positively correlated to
the size of th~ f~male (Wootton, 1973 ~). ~e suggested that
pouu1~t1an estimates which fgnorP.d the high potential of egg
production a!'u:! ~urv1va1 provided by t~e J~R~lti-seasonal
spawn i 'lg capac 1 ty of 1 a rge fe!l'la 1 es B nd the protective
nestin~ behavior of males could grossly underestiMte
stickleback produc:t1on and would incorrectly assess the
importance of this species to the energetics of the
freshwater com.unity. Potapova et aT. (1966) found that the
quantity and vhbiHty of egg production fs related to
growth rate and lipid storage; large females exhibit higher
fecundity.
The rf gors of breed i ng ev i dent 1 y 1 ead to i nc reased
mortali~v. This view is supported bv detailed descriptions
of the complex nature of stickleback reproductive behavior
provided by Tinbergen (1952) and van Ierse1 {1953, cited bv
Wootton, 1976). High mortalities reported by Rogers fn lake
Aleknagik may have resulted from a decreased resistance to
high temperature due to the stress of spawning. Narver
(1g68) observed high death rates associated with spawning
stress in the Chignik Lakes, Alaska. These high mortalities
and the territorial spawning of sticklebacks were considered
possible density-dependent population regulators. Hagen
(1967} described post-reproductive mortality in the
threespine population inhabiting the Little Campbell River,
B.C. Many adults were found in extremely poor condition;
fungus and other parasitic eruptions were noted.
Sticklebacks in Karluk and Bare lakes on Kodiak Island
apparently incurred a high mortality after spawning and
appeared nutritionally deprived (Greenbank and Nelson,
1959). Engel (1971) reported post-spawning mortality fn
study lakes on the Kenai Peninsula, Alaska.
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3. Feeding and Competition
Feeding investigations of stickle~acks in North America have
determined that their diet is composed mainly of zooplankton
and insects. The fish are opportunistic and food sources
:tre related to seasonal and regional availability. Carl
(lg53) found cladocera and copepods in the maj~rity of
stickleback stomachs collected in Cowichan Lake, B.C.;
insects, ostracods, anphipods and algae were also observed •
Greenbank and Nelson (19Sg) discussed the importance of
chironomid larvae and pupae, copepods and cladocera in the
feeding of sticklebacks in Karluk and Bare Lakes. Other
items encountered occas 1 ona 11 y were pea c 1 ams, ostracods,
rotifers, snails, leacht , planarians, fish eggs and water
mites. The importance of chironomids decreased over the
summer. Most larger sticklebacks fed on ostracods and fish
eggs; fewer of them fed on cladocera. Chironomid larvae and
entomostraca were found to be primary food types in
threespine sticklebacks in littoral areas of Lake Aleknagik
with Entomostraca being their major prey in limnetic areas
(Rogers, lg68). Sticklebacks inhabiting Black Lake, a
shallow lake, fed predominately on insect larvae;
threespines in Chignik Lake. which has only a small littoral
zone, fed totally on zooplankton (Parr, 1972).
In Lower Jean Lake, feeding habits of threespines reflected
differences in their distribution (Cannon, 1981) and
seasonal avail~bility. ~ish utilizing bottom waters preyed
on food species which were not caught in p 1 ank ton hat~l s
because most of these species live on or near the bottom.
Threespines foraging in the limnetic surface zone preyed
largely on planktonic invertebrates. Age 1 sticklebacks
{~. 30-32 mm) fed predominately on copepods in mid-June;
but for Age 0 sticklebacks ('~L 28-30 mn) rotifers were
dominate prey during J~ly. The availability of prey species
-9------~ ------
in surface waters may have fluctuated during the summer, due
to variations in their abundance or distribution. Surface
and bottom feeding threespine sticklebacks in Lower Jean
Lake were eco 1 ogi ca 11 y separated (Cannon, 198 1).
Distributional differences were considered a mechanism by
which stickleback age classes minimized competitive stress
and mu:1m1zed the uti 1fzat1on of Take resources. The
ability of threespfnes to utilize 11mnetfc waters as well as
bottom environments demonstrated their adaptability to use a
wide rangl! of conditions thus allowing exoloftation of
diverse aquatic niches. Separation of age classes between
limnetfc and bottom habitats were believed to be associated
with spawning migrations. Exposure to a wide variety of
habitat types was facilitated by these movements.
Anatomical and physiological differences between age groups
may have enforced separation.
Because spawning and early rearing occur in shallows,
stickleback life history is closely associated with littoral
habitats. The quantity of suitable littoral area
potentially serves as an important stickleback population
regulating factor (Narver, 1968). Ecological expansion
would depend on the physical condition and the magnitude of
co-actions {competition and predation) found in adjacent
environs. For these reasons, the variety of potentia 1
niches accessible to sticklebacks and consequently their
distributional patterns can be expP.cted to vary seasonally
and between systems.
Spatial differences exhibited between Age 0 and Age 2
sticklebacks and rearing sockeye salmon in L~~er Jean Lake
provided a mechanism to reduce i nterspeci fi c competition
(Cannon, 1981}. Rearing sockeye were not caught in surface
tows after mid-July; this decline occurred in conjunction
with a large recruitment of Age 0 sticklebacks.
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Feeding similarities between sticklebacks and rearing
sockeye have been reported . by Burgner ( 1958) in the Wood
River Lakes9 Alaska. Narver (1968) in the Chignik Lakes,
Alaska, and Krokhin (1957) in the Kamchatka Lakes, USSR.
Common prey preferences have suggested potential competition
for food. In nature, many similar species appear to coexist
while seemingly in competition; however9 detailed
observations have revealed differences in habitat and
behavior that permit coexistence.
Rogers (1968) compared the food of sockeye salmon fry and
threespine sticklebacks in the Wood River Lakes. In
littoral areas, t~e benthos was used more by sticklebacks
than sockeye fry; surface insects were uti1 ized more by
sockeye. In 1 imnetic areas, winged insects were an
important prey for sockeye, but were rarely ingested by
sticklebacks. Parr (1972), who studied the feeding of
juvenile sockeye and resident fish species including
threespine in the Chignik Lakes, indicated that a dis-
similarity existed between stickleback and sockeye feeding.
Sockeye more frequently fed on winged and pupal insects. In
Lower Jean Lake, a comparison of feeding between these
species (although limited to only a five week period) found
similar differences (Cannon, 1981). Sticklebacks and
sockeye i nhabi ti ng surface waters during this period
utilized similar prey, but sockeye appeared to prefer winged
insect species more than St!Cklebacks.
Partitioning rations among sticklebacks and rearing sockeye
in Lower Jean Lake appeared to be accomplished through
differences in spatial distribution (Cannon, 1981). Age 2
threespine utilized bottom habitats. Age 1 sticklebacks and
juvenile sockeye exhibited similar distribution patterns and
feeding; however, spatial overlap and prey similarity are
only prerequisites to potential competition. A significant
-11-
decrease in Age 1 threespines was observed during the summer
in Lower Jean Lake. This reduction was believed to be
associated with reproductive migrations which brought adult
fish bottom envil·onments. The rapid recruitment of Age 0
sticklebacks into the near surface waters during July may
have influenced the apparent shift in vertical distribution
of juvenile sockeye into deeper water stratums and
restricted the return of Age 1 sticklebacks into limnetic
areas. The high preference of Age 0 sticklebacks for
rotifet·s was not shared with sockeye or adult threespine, a
small mouth size possibly was responsible for the apparent
differences in feeding preference, although Age 1 threespine
of approximately the same size fed predominately on copepods
in June when rotifers were abundant in the net plankton.
Temporal variations in the vertical distributions of prey
species may have been involved. Rogers (1968) suggested
that temporal as well as spatial differences in
stickleback-sockeye feeding relationships could exist.
Narver (1968) developed a conceptual 1110del of sockeye-
stickleback co-actions for populations in the Chignik Lakes.
He concluded that threespine sticklebacks which wer£
ecologically a littoral species would be displaced from
n mnet i c a rea s by rea r1 ng sockeye if the young sa 1 mon
remained abundant. Parr (1972) determined that the
abundance of Age 0 sockeye i, the Chignik Lakes had an
adverse effect on resident fish populations including
threespine sticklebacks. Burgner (1958), Kerns (1965), and
Rogers (1972) have suggested a similar population regulation
of stickleback abundance by large Age 0 sockeye salmon
recruitments in the Wood River Lakes. Reduced spawning
escapement due to overharvest or environmental factors have
resulted in periods of reduced sockeye fry abundance in the
Wood River and Chignik Lakes. During these years,
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stickleback:; were able to rapidly occupy vacant niches
(Burgner 9 1958; Narver, 1968).
Ttle threespine stickleback is tne dominant fish speciu
inhabiting Lower Jean Lake {Cannon, 1981). Its dominance is
favored by a prolonged spawning time, a substantial littoral
spawning habitat with abundant submergent vegetation. an
ability to utilize deep and shallow bottom habi~ats as well
as pelagic areas for feeding, reduced intraspecific
competition between age classes via differences in their
spatial ben~vior and diurnal migrations, and an apparently
low level of interspecific competition frc,r. rearing sockeye
salmon in limnetic areas.
4. Distribution and Behavior
An apparent diel migration of threespines in surface waters
of Lower Jean Lake was observed by Cannon (1981) during the
summer. The abundance of sticklebacks in surface tow net
samples collected in the Wood River Lakes by Burgner {1958)
remained high throughout the day and night. Diel
zooplankton migrations prob!bly influenced stickl~back
distribution in Lower Jean Lake. Stomach analysis of
threespine from different periods showed no significant
decrease in numbers of whole undigested zooplankton. If
feeding continued throughout the day and night, sticklebacks
conceivably would concentrate at depths where prey density
was high. Vertical migrations of l1metic zooplanktens
comnon 1 y occu rri ng in A 1 as ka 1a kes have been observed
{ Rogers • 197 4 ) .
A reduction in interspecific competition between
sticklebacks and sockeyes could result from seasonal
va~iations in vertical distribution. Because threespine are
more resistant to high temperature and illumination and
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because they are equipped with a se 1 f-conta i nen defense
mechanism, near-surface water residence would be less
ecologically intolerable for sticklfback than for sockeye.
Outside of breeding season, stiddt:~ack feed in schools
(Tinbergen 1952). Generally, schooling fish are the same
size because fish of similar size swim at the same speed.
Size in sticklebacks varys between sexes and schools
comprised of all males or all females have been observed
{Narver, 1968). Possible benefits attributed to schooling
behavior have included increased feeding efficiency,
predator detection and defense, increased ability to locate
a mate, enhanced learning ability, and efficiency of
movement (Eggers, 1975). Schooling was observed in
sticklebacks in surfac ~ waters (juveniles) and in shallow
littoral areas (adults) of lower Jean Lake (Cannon, 1981).
MacMahon (1946) described the threespine stickleback as the
fiercest "reshwater fish in Britain for its size. Its
highly flexible fin motions are well suited to feeding in
the dense vegetation and submerged debris of the littoral
environment. Hagen (1967), who conducted dispersi~n studies
of threespine, reported that sticklebacks (leiurus) were a
sedentary fish. Recaptures of marked fish were never made
beyond 200 m from the point of release. Sticklebacks in
Lower Jean Lake generally exhibited -3 lethargic swimning
activity (Cannon, 1!:181). Adult sticklebacks observed in
schools feeding in algae beds and solitary f1sh which were
probably spawning moved slowly through the water. Even
pelagically feeding schools of Age 0 threespine swam
sluggishly near the surface. Rapid swimming motions
occurred during terri tori a 1 defense and occasionally in
pursuit of prey, but only for short distances.
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0. Ecological and Economic lmoortance
The threespine stickleback is often abundant whe~e it is found
and plays a significant ro~e as a predator, competitor, and prey
species in many lake ecosystems. There has been cor1cern
regarding the stickleback a~ a potential competitor with sockeye
salmon {Oncorhynchus nerka) fry but recent studies {Cannon, 1981;
Manzer, 1976; Rogers, 1972; Wootton, 1976} have indicated that
competition to the detriment of the sockeye fry does not often
occur. (See also discussion in Section I.C., life History
Summary.) In artificial situations, such as the reclamation and
re-stocking of lakes with rainbow trout(~ gairdneri), the
presence of threespine sticklebacks may be detrimental to the
trout population (Engel, 1971). In many areas, sticklebacks are
an important prey species for predaceous fish such as trout,
salmon, and northern pf ke (McPhai 1 and l1 ndsey, 1 970; Wootton
1976) and for fish-eating birds (Scott and Crossman, 1973}.
The threespine stickleback has been harvested only t'l a minor
extent. They have been used for oil, meal, fertilizer, and
animal food, including sled dog food (Wootton, 1976}.
Because the threespine stickleback is hardy, easy to keep, and
widely distributed, it has proven to be an important laboratory
fish. It has been extensively used for behavfo~al studies and
for studies on the effect of water pollution (Wootton, 1976).
-15-
II. SPECIFIC HABITAT REQUIREMENTS
A. Spawning
1. Temperature
The surface water temperature during the breeding season in
Lower Jean Lake ranged from about 12 to l8°C (Cannon, 1981).
In a st:-eam of southern British Columbia, the average
temperature during the breeding season was 16°C (Hagen,
19€7) and a stream in England during the spawning season had
temperatures ranging from 16 to l9°C (Lindsey, 1962).
Greenbank and Nelson {1959) stated that the water
temperature i~ two lakes on Kodi~~ Island may influence the
time of spawning. Threespine sticklebacks spawned earlier
in Bare Lake (surface water temperatures from mid-May to end
of July were 4.4-22.8°C) than in the deeper Karluk Lake
( 3. 3-13. 9°C). Baggennan ( 1 957, cited by Wootton, 1976)
found that increased temperatures (about 20°C). given a
sufficiently 1 ong dayl ength, accelerate the maturation
process.
2. Water Depth
Male threespine sticklebacks build their ne~ts in streams or
in shallow areas of lake shores. Hagen (1967) found that
the average depth of nests in Little Campbell River, B.C.,
was 24 em; some nests were built in water as shallow as 4
em.
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3. Substrate
The nests are constructed of small twigs, plant material,
and sand. Some nests may be constructed mostly of sand
grains (Greenbank and Nelson, 1959; McPhail and lindsey,
1970).
Nests in the River Wear, England, were built on a •muddy•
bottom (Wootton, 1976). The same substrate was used by the
le1urus form in little Campbell River, B.C. (Hagen, 1967).
Hagen gave male sticklebacks a choice between "s1nd 11 and
11 111Jd" substrates 1 n the 1 aboratory and found that they
demonstrated a strong preference to build their nests on the
mud. The leiurus form in Hayer Lake, B.C. also occurred on
a soft 111Jd bottom (Moodie, 1972). Scott and Crossman (1973)
state that threespines prefer sandy areas for nest building
(form not mentioned).
4. Aquatic Vegetation
Stickleback nests are usually found in or nP.ar aquatic
vegetation.
Hagen ( 1967) obser-ved stick 1 eback ( 1 ei urus} nests in the
little Campbell River, B. C. near dense stands of aquatic
vegetation such as Oenanthe, Potomogeton, Nuphar, Carex,
Myosotfs, Glyceria, Typha. Lemma, and green algae. The fish
always nested among broadleaved vegetation.
In the laboratory, Hagen ( 1967) presented ma 1 es of the
leiurus form with a choice between Oenanthe (a plant found
in the headwaters) and Elodea (a lower river plant) and
found a strong preference for the former. In a sur-vey of
streams on Vancouver Island, during breeding season, Hagen
found leiurus plentiful only in areas with dense aquatic
•
vegetation. The leiurus fonn in Mayer Lake, B. C., occurs
only among the thick vegetation of inlet stream margins and
stream mouths and apparently does not occur in open water
(Moodie, 1972). The littoral zone of Hayer Lake is densely
covered with Sphagnum and emergent grasses.
Na rver ( 1 g66) reported that spawning stick 1 ebacks in the
Chignik Lakes were most abundant where aquatic vegetation
was plentiful. However, Karluk and Bare lakes on Kodiak
Island, which have good populations of sticklebacks, have
only sparse aquatic vegetation (Greenbank and Nelson, 1959).
5. Light
Gonad maturation of three~pine sticklebacks in the spring is
depend~nt on both an adequate light intensity and on an
adequate daylength (Baggerman, 1957, cited by Mcinerney and
Evans, 1970). Baggenman found that high temperature (20°C)
by itself is not effective in inducing sexual maturation,
long photoperiods are also required. Baggerman also showed
that sticklebacks exposed to 269-323 lux (25-30 ft-candles}
matured slightly more rapidly than those exposed to 161 lux
(15 ft-candles). Mcinerney and Evans (1970) reported that
sticklebacks (presumably the trachurus fonn) exposed to an
energy level of 370 ergstcm 2-sec in the laboratory had
maturation rates comparable to those of wild fish. This
energy level is equivalent to illuminance levels ranging
from 230 lux (at the green wavelength) to 5 lux (at the
purple wavelength}. Mcinerney and Evans also tested the
effect of light quality (wavelength) by exposing fish to
four segments of the visible spectrum ranging from 388-653
millimicrons (long ultraviolet to short red). They found no
major differences among the four in affecting the rate of
gonad maturation.
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6. Size of Territory
Wootton (1976) states that the maximum density of threespine
stickleback nests is 4-51m 2 and that the minimum distance
between nests is 30-50 em. The average distance between
nests in the River Wear, England, ranged from 143-237 em
(Wootton, 1976).
B. Incubation of Embryos
1. Temperature
The time to hatching is directly dependent on temperature.
Wooton (1976} plotted the data of several investigators and
found that the time to hatching varies from about 5 days at
25CIC to about 15-43 days at 8°C. At 18-l9°C, hatching
occurs in about 8 days and the yolk sac is absorbed in
another 4 days. Studies in Alaska have reported hatching
times of 14 days at a water temperature ranging from 9-l6°C
{Kodiak Island; Greenbank and Nelson, 1959) and 5 days at a
water temperature varying from 21.1-22.8°C (Kenai Peninsula;
En~el, 1971). Neither of these studies was in situ.
Heuts (1947), in laboratory studies in Belgium, found that
the best survival of eggs incubated in freshwater occurred
at water temperatures of 1~-26°C, Heuts showed that the
eggs are adapted to a narrow range of temperatures. Lindsey
( 1962) reared the freshwater form in the laboratory at
temperatures ranging from 10-28°C. No eggs were
successfully reared in freshwater at 10°, 12°,14°, or 28°C,
and survival at 16° and 18°C was less than optimum.
However, there may have been factors present other than
temperature which lowered the survival rate. Lindsey also
found that fish reared at higher temperatures (22°C and
greater) had a higher proportion of females than fish reared
-19-
at 20°C and below. Lindsey further showed that eggs from
females with few lateral plates have a higher optimum
development temperature than eggs from females with more
plates. The approximate optima were: 26°C (2 maternal
plates), 20-27°C (4 or 5 maternal plates}, 20-22°C (6
maternal plates). and l6°C (7 maternal plates).
Wootton (1976). citing the work of Swarup (1958, 1959),
stated that abnonm~l development occurs when newly
fertilized eggs are exposed to very low {0°C) or very high
(33°C) temperatures for a duration as short as 1.5-3.0
hours. One of the abnonm~lities was the production of fish
with a triploid number of chromosones. These fish developed
and grew at the same rate as nonm~l (diploid) fish, but were
misshapen.
Hagen {1967} measured water temperatures ranging from 16 to
23°C during the nesting of sticklebacks in Little Campbell
River. B.C. Some approximate surface water temperatures for
various lakes in Alaska measured at the approximate time
incubation occurs are: Bare Lake, 6? -23°C, and Karluk
Lake, 4? -l5°C (Greenbank and Nelson. 1959); Lower Jean
Lake, 13 -18°C (Cannon, 1981); Johnson and Scout Lakes on
the western Kenai Peninsula, 12 -18°C, and Bear Lake in the
Kenai Mountains, 7 -l5°C (Engel, 1971); Wood River Lakes,
10 -14°C (Rogers, 1968); Lake Nerka. 9 -l8°C (Burgner,
1958); Black Lake, 12 -l5°C, and Chignik Lake, 9 -l3°C
(Parr, 1972).
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2. Dissolved Oxygen
Given the fact that sticklebacks nest in shallow areas with
adequate light penetration and aquatic vegetation. dissolved
oxygen l eve 1 t 1 n the water near the nests are probab 1 y
rarely a limiting factor. However. the eggs are placed
inside covered nests, often on muddy bottoms containing much
dead organic matter, and often in areas of 1 ittle or no
current, so circulation of water through the nest to replace
the oxygen used by the eneryos is necessary and is
accomplished by the fanning of the mele. In the absence of
fanning, van lersel (1953, ci.ted by Wooton, 1976) noted that
eggs became moldy and died. The rate of fanning reaches a
peak shortly before hatching; presumably. this is when the
oxygen requirement of the et!Cryos is highest. When van
Iersel ran water low in dissolved oxygen and high in carbon
d i oxide through a nest in the 1 a bora tory, the amount of
fanning by the male increased.
C. Juvenile Rearing
After leaving their nests, young sticklebacks form schools, which
may be an adaptation for cover, and eventually join the adults.
There is little information to suggest that young of the year
have habitat tolerances, preferences, or requirements different
from those of adults.
D. Adult L1 fe
1. Temperature
The adult threespine stickleback is usually regarded as a
eurythermal fish (Wootton, 1976). Jordan and Garside (1972)
studied the upper lethal temperature of threespine
sticklebacks (probably the trachurus form) fram the harbor
-21-
of Halifax, Nova Scotia (where the salinity ranges from
20-30 ppt) which had been acclinaated to various combinations
of temperature and salinity. The highest upper lethal
temperature {28.8°C) was noted for fish acclimated to 20°C
and 30 ppt salinity, tested at 12 ppt, and the lowest upper
lethal temperature (21.6°C) was shown by fish acclimated to
1o•c and a salinity of 0 ppt, tested at 30 ppt. Fish which
had been acclimated to freshwater at 20°C, and tested in
freshwater, had an upper lethal temperature of 27 .2•c.
There were no significant differences in survival among the
d1fferent sized fish (total lengths ranged from 30-80 m)
tested. Coad and Power (1973), citing Bertin (1955) state
thctt the threespine can tolerate temperatures around 25°C
an~ is also tolerant of temperature changes. Heuts (1947}
found with the leiurus form collected at 0°C and placed into
water at 25 -2a•c that fis~ with fewer lateral plates
survived longer than fish with more latera.l plates. Mean
survival time was around 40 hours. Heuts also stated that
the freshwater form will not tolerate '10111' temperatures.
The threespine stickleback is found in Alas.kan lakes with
late spring to early fall temperature ranges of anywhere
from o• to 23°C (Burgner, 1958; Cannon, 1981; Engel, 1971;
Greenbank and Nelson, 1959; Parr, 1972; Rogers, 1968 and
1972). The stickleback may not be found at the extremes of
this range if they have a choice of more moderate
temperatures. Little information is available on
teft1)erature preferences or on tMPerature distribution
durfng late fall, winter, and early spring. In the su~m~er,
sticklebacks move into warmer, shallower, water but this is
probably more a function of reproductive and fe-eding
requirements than a demon~tration of temperature
preferences.
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Temperature influences the rate of growth. Wootton (1976),
citing the work of Cole, reported that the mean growth
efficiency increases from 5.9~ at 7,0°C to 11.3~ at 20.0°C.
Beukema ( 1968, cited by Wootton, 1976) found that the
feeding rate also depends on temperature, the ri!'te of
stomach evacuation ranging from about 16 hours at l'i-12°C to
"one night" at 18-20°C.
2. Water Depth
3.
The depth distribution of threespine sticklebacks in lakes
resu 1 ts from temperature preferences, feeding mi gra t 1 ons,
reproduction requirements, predation and competition. They
are generally a ~nallow water fish, particularly in the
su11111er when they move into the shoa 1s along the shore.
During the summer they range from the surface to the bottom
in lower Jean Lake (21.3m) (Cannon, 1981) and Johnson (4.0
m), Scout (6.1 m), and Bear (18.3 m) lakes (Engel, 1971) on
the Kenai Peninsula. In Karluk Lake on Kodiak Island, they
have been caught from the surface down to 24.4 m, but none
were caught in attempts at 38.4 m or at 61.0 m (Greenbank
and Nelson, 1959). Very few fish were caught below surface
waters in Lake Nerka during the sunner; the deepest
stickleback was caught at 7.3 m (Burgner, 1958).
Current Velocity
Although threespine sticklebacks commonly occur in streams,
they prefer areas with little or no current (Hagen, 1967).
Hagen conducted a thorough study in Little Campbell River,
B.C., and determined that swift waters are an unfavorable
habitat. The average current velocity in a section of the
stream where the leiurus form was plentiful was 3 em/sec
(gradient about l . 5 m/km). A section with an average
velocity of 23 em/sec had only a few sticklebacks of the
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leiurus form and a riffle section with an average velocity
of 74 em/sec had none. When Hagen transplanted the fish
from the low current area into an area of fast current, they
migrated into areas of slower current. When Hagen gave
males a choice between standing water and moving water in
an aquarium, they demonstrated a strong preference for
nesting in the standing water. Hagen also reported that
some fish successfully passed through a 70 m long culvert
with a current velocity of g2 em/sec, although this was not
a common occurrence.
4. Dissolved Oxygen
KroKhin (1957) calculated that the oxygen consumption of
threespine sticklebacks in KamchatKa lakes where the
temperature ranges from 2.0 to 14.3°C would range from 0.127
-0.365 mg 02/hr-g live weight. He found that fish in the
laboratory at a temperature range of 0.5 -19.5°C used
oxygen at a rate ranging from 0.12-0.55 mg 02/hr-g live
weight. Threespine sticklebacKs from some English streams
had an oxygen consumption rate ranging from 1.0 microliters
o2thr-mg dry weight (1 microliter= 1.43 micrograms) for 500
mg dry weight fish to about 2.5 microliters/hr-mg dry weight
for 70 mg dry weight fish (temperature not given; Lewis et
a 1. , 1972).
Jones {1948 and 1952, cited by Wootton, 1976} stated that
the minimum dissolved oxygen (DO) level at which threespine
sticklebacks can exist is about 0.25 -0.50 mg/1. Jones
found that the avoidance response of sticKlebacKs is
triggered when the fish are exposed to water with a DO level
of 0.3 mg/1 at low temperatures. At 20°C, the response
occurs at 2.0 mg/1, indicating the fish have a lower
tolerance for low DO conditions at higher temperatures. In
a survey of some English strea~s. Lewis et al. (1972) found
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that threespine sticklebacks were most abundant in waters
with 8 -12 mg 0211, less abundant in waters with 6-8 mg
0211, and absent from waters with 2-5 mg 0211.
In Lower Jean Lake, sticklebacks were caught near the lake
bottom during the s~~r where DO levels ranged from 3.0 -
6.5 mg/1 (Cannon. 1981). These levels apparently had no
effect on feeding activity. Dissolved oxygen levels above
the thermocline r!~ged from 10.1-14.0 mg/1. Greenbank and
Nelson (1959) reported that the ~tfckleback population fn
Karluk and Bare Lakes are in waters where there 1s an
11 abundance" of dissolved oxygen at all depths during the
summer. They also stated that the threespine stickleback
"is known to survive over winter in shallow lakes in
northern temperate and subarctic zones, where •••• dissolved
oxygen sinks to a trace •••• ".
5. Chemical Parameters
Threes pine stick 1 ebacks have been reported to occur in
waters with a pH range of: 7.0 • 8.7 in Karluk and Bare
Lakes (Greenbank and Nelson, lg59); 6.3 -7.0 in three Kenai
Peninsula lakes (Engel, 1971); and 6.8 in Little Campbell
River, B.C. (Hagen, lg67). Jones (1948, cited by Wootton.
1976) tested the tolerance of the fish to a wide pH range
and reported that they avoided a pH of less than 5.6 or
greater than 11.4.
A survey of some English streams conducted by Lewis et al.
(lg72) showed that the threespine did not occur in waters
which smelled of hydrogen sulfide and rotting vegetation.
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6. Feeding
Krokhin (1957) calculated that threespine sticklebacks in
some Kamchatka lakes, where the annual temperature range is
2.0 -14.3°C, consume daily between 0.08 g/fish in winter and
0.23 g/fish in August. This represents 1.8-5.1~ of body
weight. The monthly food consumption ranged from 2. 70 -
6.77 g/fish, for an annual total of 42.49 g. which is
equivalent to 8 or 9 times the average weight of an adult
(4.5 g). In Lake Dal'neye, Krokhin (1g7Q) calculated that . 2
the monthly food consumption ranged from 0.80 9/g 1 he
weight in January to 2.185 g/g live weight in August.
Manzer (1976) reported that the daily ration of threespine
stickleback in Great Central Lake on Vancouver Island, B.C.,
was 6.6~ of the body weight in July and 7.8~ in October.
Wootton (1976) states that, to support a population of
threespine sticklebacks, an area must produce suitable food
on the order of 10 -100 gtm 2-yr.
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III. CONCEPTUAL SUITABILITY INDEX CURVES
Conceptual suitability index curves are presented for water
temperature, current velocity, water depth, dissolved oxygen
concentration, and pH in Figures 2 through 4. Data to support the
curves is included 1 n Tab 1 es I , ll, and II I. These curves hou 1 d .1ot
be construed as a graphical presentation of actual data. Rather, chey
are intended to be hypothetical models of the relationship between
threespine sticklebacks and certain environmental paramete1·s. As with
any hypothesis, they must be tested and verified before being applied
to any particular situation.
The curve5 are based on pub 1 i shed and unpub 1 i shed data and on
conversations ~ith fishery biologists who ha~~ ~orked with threespine
sticklebacks. Both experimental laboratory data and field
measurements and observations were used. The suitability index for
each environmental parameter ranges from zero to one. An index of one
indicates an optimum or preferred level of that particular parameter
and an index of zero indicates a completely unsuitable level.
Data from laboratory physiological studies in European populations of
threespine sticklebacks indicates that the optimum temperature during
incubation of embryos is 8 or 9°C higher than the average surface
water temperature during the presumed incubation period in several
Alaskan lakes (Figure 2). Until the temperature tolerance of Alaskan
sticklebacks has been studied in physiological laboratory studiest it
must be assumed that the optimum temperature for European stickleoacks
{about 22ac) does not apply in Alaskan waters which rarely reach 22ac.
A problem encountered in constructing the curves is that much of the
data in the literature concerning environmental parameters of
threespine stickleback habitat does not relate various levels of the
parameters to some measure of habitat suitability. Often, ranges of
the parameter are given based on measurements taken throughout the
area occupied by threespines but there is no indication that one point
-27-
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on th~ range is any better or worse than any other point in terms of
habitat suitability.
The curves are drawn using data from throughout the natural rar.ge of
the threespine stickleback. Although there probably are differences
in habitat preferences and tolerances for different geographical arecs
or even in different streams of the same geographical area, there i!.
not enough data to support drawing separate curves at this time.
However, one must be aware that any point on the curve, especially
toward either extreme, may be unsuitable for a particular stock. Hew
far the stock deviates from tne curve l'lkiSt be determined by field
measurements and experimentation with tnat particular stock.
A second precaution regarding the curves concerns tne interaction 1f
various parameters. A given level of one parameter can have a
different effect on the fish as the level of another parameter varins.
For example, a dissolved oxygen concentration of 3 mg/1 may be
suitable at a water temperature of 5°C, but unsuitable at a
temperat~re of 20°C. Ideally, given enougn data, a separata dissol\ed
oxygen curve snould be drawn for each of several different
temperatures. Tne overall suitability of any particular habitat is a
summation of tne interacting effects of many parameters.
A third precaution to consider is tnat the effect of less than opti~Jm
levels of any parameter on the fisn depends strongly on the duration
of exposure. Stickleback embryos exposed to a water temperature of
33°C for 1.5-3.0 hours e'perience an abnormal development (Swarup,
1958 and 1959, cited by Wotton, 1976)1 but survive. Embryos exposec
to 33°~ for a longer period of time would die.
A 1 so, different ages wi tni n a 1 i fe stage probab 1 y have di ffe ren 1•
habitat requirements. For example, tne oxygen requirement of emtryo~
is highest just prior to natching. However, because of insufficient
data, it is not possible to draw separate curves for different ages.
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Overall, the suitability index curves presented in this report provide
an indication of conditions which make a desirable threespine habitat
and conditions which make a less desirable habitat. Also, although
the curves are general, they can show differences in habitat needs
among threespi nes and other species. Further, the process of
constructing these curves is beneficial in defining areas where more
data is needed. Lastly, these curves can aid in the design of
experiments and sampling programs. As more data becomes available,
these hypothetical curves can be further refined.
~29~
Table I.
Parameter
Temperature, oc
Water depth,
em
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THREESPINE STICKLEBACK
Adult -Spawning
Observed Values
ca. 12 -18
5.6 -22.8
3.3 -13.9
16
16 -19
4
21J
Remarks location
surface water temperature during Lower Jean Lake
breeding season
surface water temperatures Bare Lake
during June and July Karlu~ lake
average during breeding seaso~ Little Campbell
River, B.C.
during spawning season Hobson's Brook,
England
shallowest nest little Campbell
River, B.C.
average depth for nests
Reterence
Cannon (1981)
Greenbank and
Nelson (1959)
Hagen ( 1967)
lindsey ( 1962)
Hagen ( 1967)
-------------------Table ll. THREESPINE STICKLEBACK
tncub~Lion of Embryos
Parameter Observed Values Remarks Location Reference
Temperature, 16 -26 best survival rate Belgium (laboratory) Heu t s ( 194 7) oc
10 -14 no eggs successfully reared England (laboratory) Lindsey (1962)
16 -18 survival less than optimum
20 -26 best survival
28 no eggs successfully reared
> 22 lower male:female ratio
than fish reared at ~ooc
16 -26 optimum development temperatures; the
I fish with fewer maternal plates t.J ..... l•dV1n9 htght:r optimum temperatures I
0, 33 exposure to these temperatures England 1 Swarup (1958 and
for 1.5-3.0 hours causes (laboratory) 1959. cited b~
abnonmal development Wootton. 1976
8 time to hatching 1s lS -43 days various Wootton ( 1976) ..
5 time to hatching ts S days
25 time to hatching 1s S days
13 -18 approximate surfac~ water temp. lower Jean lake Cannon ( 1g81)
during t ncuba t! 'ln ~~eri od
12 -18 approximate surface water temp. Johnson lake Engel (1971)
during tncubatton period
Table II. Cont'd THREESPINE STICKLEBACK
Incubation of Embryos
Parameter Observed Va1uP.s Remarks location Reference
Temperature 12 -18 approximate surface water temp. Scout lake oc during incubation period
7 ~ 15 approximate surface water temp. Bear lake
during tn~ubation period
6? -23 approximate surface water temp. Bare lake Greenbank and
during incubation period Ne 1 son (lg5g )
4? -15 approximate surface water temp. Karluk Lake
during incubation period
10 -14 approKimate surface water temp. ~Jood River lakes Rogers ( 1g68)
during incubation period
.)
9 -18 approximate surface water ;) temp. lake Nerka Burgner ( 1958)
during incubation period
12 -15 approximate surface water teinp. Black lake Parr (1g72)
during incubation period
g -13 approximate surface water temp. Chignik Lake
during incubation period
--
-------------------Table III. THREESPINE STICKLEBACKS
Adults
Parameter Observed Values Remarks Location Reference
Temperature, 27.2 Upper lethal temperature for the Halifax. Nova .Jordan and Garside ac trachurus (1) form acclimated to Scotia ( 1972) 2ooc in freshwater (laboratory)
25 tolerable general Bertin {1925, cited
by Coad and Power,
1973)
25 -28 The leiurus form collected at 0° Belgium Heuts ( 1947)
and placed into water with these ( 1 a bora tory)
temperatures died after an average
of 40 hours; fish with fewer
lateral plates survived longer
than fish with more plates
I w w 7.0 mean growth efficiency of 5.9~ 1 Cole (unpublished, I
cited by Wootton,
1976)
20.0 mean growth efficiency of 11.3t
11 -12 rate of stomach evacuation was 1 Beukema (1g6R,
16 hours cited by Wootton,
1g76)
18 -20 rate of stomach evacuation was
••one night 11
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Table Ill. Cont'd THREESPINE STICKlERACK~
Adults
Parameter Observed Values Remarks location Reference
Temperature, 6.9-17.7 early June-mid-July; obsP.rved water temp. lower Jean lake Cannon (1981) oc fish present
3.9 -15.0 Jun 1 -Sep.15; observed water temp.
fish present
Johnson and Scout Engel (1971)
lakes
8.9 -18.3
4.4 -22.8
3.3-15.0
6.7-14.2
0.0 -20.0
11.5-15.0
9.0 -13.0
Jun 1-Sep.15~ observed water temp.
fish present
mid Hay-mid Sept.; observed water temp.
fish present
mid Hay-mid Sep.; observed water temp.
fish present
Aug .• observed water temp. fish present
Jun -Sep; 6 yrs; observed water temp.
fish present
Bear lake
Bare lake
Karluk lake
Wood River lakes
lake Nerka
early Jul -early Sep.; observed water temp Black lake
fish present
late Jun-early Sep.; observed water temp Chignik lake
fish present
Greenbank and
Nelson (1959)
Rogers (1968)
Rurgner ( l'l58)
Parr ( 1972)
------.. ------------Tahle I I I. Cont 'd THREE('" INE STICKLEBACKS
Aduits
Parameter Observed Valu~s Remarks location Referen :e
Water Depth, 0-21.3 (bottom) SUIIJI"'er lower Jean Lake Cannon (1981)
m 0 -4.0 (bottom) SUill!ler Johnson Lake Enge 1 ( 1971)
0 -6.1 (bottom) SUill!ler Scout lake
0 -18.3 (bottom) SUill!ler Bear lake
0 -24.4 SUITiller Karluk Lake Greenbank and Nelson
(1959)
38.4, 61.0 nnne caught; sunmer
7.3 greatest depth at which fish lake Nerka Burgner ( 1958)
I were caught; w SUill!ler
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0 -10 Jun -SeJ,tt. Kamchatka ~rokhin ( 1957)
lakes, USSR
50 -60 Dec., Jan., Feb.
Current 3 average in leiurus habitat Little Campbe 11 Hagen ( 1 96 7)
Velocity, R1ver, B.C.
em/sec
(stream 23 rna rgina 1 habitat
population)
74 poor hab tta t
92 fish were able to negotiate
a 70 m culvert with this velocity
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TablP. Ill. Cont'd THREESPiN£ STIC~lEOACKS
Paran1eter
Dissolved
Oxygen
mg/1
plf
Adults
Observed Values
0.25 -0.50
0.3
2.0
8 .. 12
6 - 8
2 -5
3.0 -6.5
10.1 .. 14.0
< 5.6 or>ll .4
6.A
6.3-7.0
7.0-8.7
Remarks
minimum level tolerable
level at which avoidance response
is triggered at low temps.
level at which avoidance response
is triggered at 20°C
fish most abundant in reaches with
this 1 evel
fish less abundant in reaches w1th
this level
fish absent from reaches with this
level
healthy fish caught near lake bottom
at this level; summer
level above thermocline, summer
avoided by fish
measured value
r..easured value
measured value
Location
Great Britain
(laboratory)
England
lower Jean Lake
Great Britain
( 1 a bora tory)
little Campbell
River. B.C.
Kena 1 Pen i nsu 1a
lakes
Karluk and Bar~
lakes
Reference
Jones (19t18 and
1952, cited by
Wootton. 1976)
lewis et a 1 .•
( 1972)
Cannon (1981)
Jones (1948, cited
by Wootton, 1976)
Hagen ( 1967)
Engel (1971)
Greenbank
and Nelson (1959)
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THREESP1NE
5 T; CK LE SACK
I NCUSATlON
F1gure 2 Conceptual model of relct•onsh•p between
Threesp1ne stJck.leback embryos and toctmperoture .
See text for Qualifications for use or this curve
(NOT RECOMMENDED II'OR APPUCATION TO I"ECIFIC WATERSHEDS
WITHOUT FIELD YER1f'ICAT10N)
-37·
THREESP!"lE
STI(t<LEBACK
1 o-,..
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os,
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10
ADULTS
20
See text for Qualific a tiona
for use of these curves
(NOT IIIECOMMENDED I'OR APfiiLICATION
TO lnCIPtC WAT!Ra.t!.DS W'.THOUT
I'IELD Vlfi.IC.,TION)
\
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\
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30
Water temperature. •c
0~--------+-------~---------+
20 60
Water deptn, m
F1gure 3 . Conceptual model of relot1on~l'\lp betwee-n Tnreesp1ne
St1c:klebac:k adults and water temperature, ~'.lrrent
1eloc1ty (stream populot1ons only}. end wcrer depth
-38-
I THREESPINE
STICKL EB.ACK
I See text for qualifications
for use of these curves
(MOT ._I COMMENDED ,OR A~ICA nON
I WITHOUT
ADULTS
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I 1.0
I
I O.S
I
I 0
0 A 8 12
I )( Dissolved lftg/1 w nxygen,
0 z
I > .... 1.0
~
I CD < .... -
I ~
V'l 0.5
I
I 0
0 6 8 10 12
I pH
I
Figure A. Conceptual model of relot1onship between
I Threespine st1ckleback adults ond dessolved
o~eygen concentration and pH.
I
-39-
IV. DEFICIENCIES IN DATA BASE AND RECOMMENDATIONS
Although extensively used in behavioral and pollution studies
elsewhere, the interest in the threespine stickleback in Alaska has
centered mainly on its supposed role as a competitor with rearing
sockeye salmon and rainbow trout. These Alaskan studies, therefore,
have naturally concentrated on the distribution and feeding of
sticklebacks.
Physiological studies on Alaskan populations are needed to define the
to 1 era nces to en vi ronmenta 1 pa ramete ~ such as temperature and
dissolved oxygen concentration. This need is dramatically shown by
the difference between European physiological studies of temperature
tolerance and the known temperature regime existing in Alaskan lakes
where the sticklebacks are present (Fig. 2).
Infonmation is needed on stream populations of threespine sticklebacks
within the State. To date, Statewide investigations have concentrated
on lake populations.
More data from Southeast Alaska is needed to describe regional
differences within the State. The Southeast populations may be more
similar to the populations described -in British Columbia than they are
to the Bristol Bay populations. Additionally, there is some
indication that sticklebacks in the Kenai Peninsula area are exposed
to and adapted to higher temperature regimes than sticklebacks in the
Br1stol Bay area.
A study in Alaska of differences in habitat requirements among the
three fonns of Gasterosteus aculeatus (leiuru~, trachurus, and
semianmatus) would be beneficial as strong differences have been
demonstrated elsewhere; for example, Hagen's (1967) work in British
Columbia. Virtually nothing is known of the populations on the Seward
Peninsula and Saint Lawrence Island (presumably trachurus}. Because
these populations are on the edge of the species range, studies of
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these fish may provide interesting information about the environmen1.al
limits of the species.
Very little infonna.tion is available on the habitat needs of the
threespine sticKleback during the stage immediately after hatching lUt
prior to 1 eavi rg the nests. During this period, they are sti 11 ~mt1~r
the parent a 1 care of the rna l e. The r~ are a 1 so many quest i ens i bout
their life history between leaving the nest until they first begin to
show ur in min :lOW traps and fyke nets as young of the year. In
Alaska, few observations have ever been made of actual nests
-41-
LITERATURE CITED
Aronson, l. R. 1957. ~eproductive and parental behavior Chapt. III,
Part 3, 271 -3J4 in Margaret E. Brown (ed. ). The physiology of
fishes. Vol. II. Behavior. Academic Press, New York. 526 pp.
Assen, J. van den. 1967. Territory in the threespined stickleback
Gasterosteus aculeatus L. An experimental study in intra-specific
competition. Behavior Suppl. 16: 1-164.
Baerends, G. P. 1957. Ethological analysis of fish behavior. Pages
229-269 in Margaret E. Brown. (ed.). The physiology of fishes.
Vol. I. Academic Press, New York. pp.
Burgner, Robert L. 1958. A study of fluctuations in abundance, growth and
survival in early life stages of the red salmon (Oncorhynchyus nerka
Walbaum) of the Wood River Lakes, Bristol Bay, A1aska. Ph.D. Thesis.
University of Washington, Seattle. 200 pp.
Cannon, Richard. 1981. Summer feeding and distributional behavior of
threespined stickleback, Gasterosteus aculeatus, in Lower Jean Lake,
Alaska. 1974. M.S. Thesis, University of Alaska, Fairbanks. pp.
Carl, Clifford G. 1953. Limnobiology of Cowichan Lake, British Columbia.
J. Fish. Res. Bd. Can., 9(9): 417-449.
Coad, R. W. and G. Power. 1973. Observations on the ecology and
phenotypic variation of the threespine stickleback, Gasterosteus
aculeatus L., 1758, and the blackspotted stickleback,§.. wheatlandi,
Putnam 1867, (Osteichthyes: Gasterosteidae) in Armory Cove, Quebec.
Canadian Field Naturalist Si: 113-122.
Eggers, Douglas M. 1975. A synthesis of the feeding behavior and growth
of juvenile sockeye salmon in the limnetic environment. Phd Thesis,
Univers i ty of Washington, Seattle. 217 pp.
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II
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Engel, Larry J. 1971. Evaluation of sport fish stocking on the Kenai
Peninsula-Cook Inlet Areas. Alaska Department of Fish and Game.
Federal Aid in Fish Restoration, Annual Report of Progress, 1970-
1971. Project F-9-3. Job No. G-11-F. Vol. 12: 1-34.
Greenbank, John, and Philip R. Nelson. 1g59, Life history of the
threespine stickleback Gasterosteus aculeatus Linnaeus in Karluk Lake
and Bare Lake, Kodiak Island, Alaska. u.s. Fish and Wildlife Ser.
Fish. Bull. 153. Vol. 59: 537-S5g.
Hagen, M. J. 1967. Isolating mechanisns in threespine sticklebacks
(Gasterosteus). J. Fish. Res. Bd. Canada, 24 (8): 1637-1692.
Heuts, M. J. 1947. Experimental studies on adaptive evolution in
Gasterosteus aculeatus L. Evolution. 1:89-102.
Jones, J. W., and H.B.N. Hynes. 1950. The age and growth of Gasterosteus
aculeatus, Pygosteus pungitius, and Spinachia vulgaris, as shown by
their otoliths. Jour. An. Ecol. 19: 59-73.
Jordan, C. M., and E. T. Garside. 1972. Upper lethal temperatures of
threespine stickleback, Gasterosteus aculeatus (l. ), in relation to
thermal and ostomotic accli~ation, ambient salinity, and size. Can.
J. Zool. 50: 1405-1411.
Kerns, Orra E. 1965. Abundance and size of juvenile red salmon and
major competitor species in Iliamna Lake and lake Clark, 1g62 and
1963. Univ. Washington Fish. Res. lnst. Circ. 231. 35 pp.
Krokhin, E. M. 1957. Determination of the daily food rat1on of young
sockeye and three-spined stickleback by the respiration method.
(Transl. from Russian). Fish. Res. Bd. Can. Transl. Ser. No. 209.
14 pp.
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Krokhin~ Ye . M. 1970. Estimation of the biomass and abundance of the
threespine stickleback (Gasterosteus aculeatus L.) in Lake Del 'neye
based on the food consumption of plankton-feeding fishes. Problems of
Ichthyology. 4:471-481.
Lewis, David B., M. Walkey, and H.J.G. Oartnall. 1972. Some effects of
low oxygen tensions on the distribution of the three-spined
stickleback Gasteroseus aculeatus l. and the nine-spined stickleback
Pung1t1us pungitius (l.). J. Fish. Biol. 4:103-108.
Lindsey, C. C. 1962. Experimental study of meristic variation in a
population of threespine sticklebacks, Gasterosteus aculeatus. Can.
J. Zool. 40: 271-312.
Mcinerney, J. E., ~nd 0. 0. Evans. 1g1o. Action spectrum of the
photoperiod mechanism controlling sexual maturation in the threespine
stickleback., Gasterosteus acul ,eatus. J. Fish. Res. Bd. Canada 27:
749-763.
MacMahon, A.F.M. 1946. Fishlore. London, Penguin Books . 208 pp .
McPhail, J . 0., and C. C. Lindsey. 1970. Freshwater fishes of
northwestern Canada and Alaska. Fish. Res. Bd. Can. Bul. 173. 381 pp.
Manzer, J. I. lg76. Distribution, food and feeding of the threespine
stickleback, Gasterosteus aculeatus. in Great Central lake, Vancouver
Island, with comnents on competition for food with juvenile socke.ve
salmon, Oncorhynchus nerka. Fish. Bull. 74 (3): 647-668.
Moodie, G.E.E. 1972. Morphology, life history, and ecology of an unusual
stickleback (Gasterosteus aculeatus) in the Queen Charlotte Islands.
Canada. Can. J. Zoo1. 50:721-732.
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tl
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II
II
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11
Morrow, James E. 1980. The freshwater fishes of Alaska. Alaska
Northwest Publishing Co .• Anchorage. 248 pp .
Narver. David W. 1968. Pelagial ecology and carrying capacity of sockeye
salmon in the Chignik Lakes, Alaska. Ph.D. Thesis, University of
Washington, Seattle. 309 pp.
Parr, WilHam H., Jr. 1972. Interactions between sockeye salmon and
lake resident fish in the Chignik Lakes, Alaska. M.S. Thesis, Univ.
Washington. Seattle. 102 pp.
Potapova, T. L., T.V. Legedeva. and M.:. Shantunovkiy. 1966. Dif-
ferences in the condition of females and eggs of the threespined
stickleback Gasterosteus aculeatus L. Prob. of Ichth. 8: 143-146.
Rogers, Donald F. 1968. A comparison of the food of sockeye salmon fry
and threespine sticklebacks 1n the Wood River Lakes. Pages 1-43 in R.
L. Burgner (ed). Further studies of Alaska sockeye salmon. University
of Washington. Publications in Fisheries, Vol. III, Seattle.
Rogers. Donald E. 1972. Abundance and size of juvenile sockeye sal~~.
Oncorhynchus nerka, and associated species in Lake Aleknagik, Alaska,
in relation to their environment. U.S. Fish and Wildlife Serv.
Fishery Bulletin: 71: {4),
Rpgers, D. E. 1974. Alaska Salmon Studies: The study of red salmon in
the Nushagak District. Periodic Report No. 4. U.S. Dept. of
Conmerce. _ PP.
Scott, W. 8., and E. J. Cross~n. 1973. Freshwater fishes of Canada.
Fhh. Res. Bd. Canada Bull. 184. Ottaw ... 966 pp.
Tinbergen, N. 1952. The curious behavior of the stickleback. Scientific
American. 121: 2-6.
Vrat. Ved. 1949. Reproductive behavior and development of th~ threespined
stickleback (Gasterosteus acu1eatus) of California . Copeia. 4 :
252-260.
Wootton, R. J. 1973a. The effect of size of food ration on egg production
in the female threespined stickleback, Gasterosteus acu1eatus L. J.
Fish Biol. 5: 89-96.
Wootton, R. J. 1973b. Fecundity of the three-spined sticklebacks,
Gasterosteus aculeatus (L.). J. Fish. Biol. 5: 683-688.
• Wootton, R. J. 1976. The biology of the sticklebacks. Academic Press,
london. 387 pp.
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