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FRESHWATER HABITAT
RELATIONSHIPS
ARCTIC GRAYLING-THYMALLUS ARTICUS
ALASKA DEPARTMENT Of ASH & GAME
HABITAT PROTECTION SECTION
RESOURCE ASSESSMENtT BRANCH
.APRIL, 198 1
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FRESHWATER HASITAT RELATIONSHIPS
ARCTIC GRAYLING (THYMALLUS ARCTICUS )
By
Steven W. Krueger
Alaska Department of Fish and Game
Habitat Division
Resource Assessment Branch
570 West 53rd Avenue
Anchorage, Alaska 99502
May 1981
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ACKNOWLEDGEMENTS
Many people from the Alaska Department of Fish and Game and from the Auke
Bay F·lsheries Laboratory of the Natior.al Marine Fisheries Service freely
gave their time and assistance when contacted about this project and it is
a pleasure 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especially those who provided unpublished data and observations from their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, Hab 4 tat Evaluation Procedure Group, Fort Collins,
Colorado. Contract No. 14-16-0009-79-119.
TABLE OF CONTENTS
Arctic Grayling
Page
I. Introduction 1
A. Pur;Jose 1
B. Distribution 2
D. Life History Summary 4
E. Economic Importance 12
II. Specific Habita t Requirements 13
A. Lake In l ets/Outlets 13
1. Upstream Migration 13
2. Spawning 15
3. Post-Spawning Movements 19
4. Development of Eggs and Alevins 19
5. Sunmer Rearing 20
6. Migration to Overwinter i ng Areas 22
7. Overwintering 22
B. Bog Streams 22
1. Upstream Migration 22
2. Spawning 25
3. Post-Spawning Movements 28
4. OevelorAnent of Eggs and Alevins 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 Streams 34
1. Upstream Migration 34
2. Spawning 35
3. Egg and Alev in Development 36
4. Suntner Rearing 36
5. Migration to Ovel""ttf i ntering Areas 40
6. Winter Rearing 40
D. Spring Streams 42
1. Upstream Migrat i on 42
2. Spawning 43
3. Sunmer Rearing 43
4. Migrati on to Ovel""ttfintering Areas 43
5 . Winter Rearing 43
Habitat-Arctic Grayling Relati onships 44
Deficiencies in Data Base 53
Recommendations and Further Studies 55
Literature Cited 58
I . INTRODUCTION
A. Purpose
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 life stages of Arctic
grayling, Thymallus arcticus {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 Ar·ctic
grayling survival and behavior.
2} To develop an Alaska data base composed 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 recornmend
studies to fill gaps fn the data .
The following "Life Histor:-y 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:
upstream spawning mi gration;
spawning;
post-spawning movements;
incubation;
sunmer rearing;
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migration to overwintering areas; and
winter rearing
B. Distribution
The Arctic grayling is a holarctic species of the genus
Thymallus. Within North America it occurs from Hudson Bay west
through northern Manitoba, Saskatchewan, Alberta and British
Columbia, the Northwest Territories (excluding most islands of
the Arctic archipelago), l.he Yukon and most of Alaska. In
Eurasia it fs found as far west as the Kara and Ob Rivers and
south to Northern Mongolia and North Korea.
Several i solated, relict populations exist in North America. One
is located in a fraction of its original range in the extreme
headwaters of the Missouri River drainage in 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 Arctic grayling population ir the Great Lakes region was
eliminated in 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 Vermont (Scott and Crossman, 1973).
Arcti c grayling are distributed over much of Alaska (Figure 1)
(McClean and Delaney, 1978). Distribution of Arctic grayling
within southeast Alaska is primarily limited to stocked lakes.
Occasionally fish drift downstream in large stream systems, such
as the Stikine River. These rivers often support substantial
populations of grayling in their headwater reaches (McClean and
Delaney, 1978).
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Insert Figure 1
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MA IN STUDY SITES
CD T.AN.AN.A RIV ER
(2] IC.AVIK RIVER
[j] GULIC.AN.A RIVER·
I1J TYEE L.AICE
[3] COLVILLE RIVER
.ALEUTIAN ISlANDS • •
"' • ,A.J7 .t;~.::. ~tl,.,
FIGURE 1. DISTRIBUTION OF ARCTIC GRAYLING IN ALASKA (FROM
ALASKA DEPARTMENT OF FISH AND GAME, 1978)
AND MAIN STUDY SITES .
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( .
Numerous clearwater tributaries and lakes within the upper Copper
and Susitna River drainages contain Arctic grayling. Grayling are
found in the Gulkana and Oshetna Rivers. Arctic grayling are
extremely limited within the Prince William Sound area and lower
Copper River drainage and somewhat limited within the lower
Susitna River drainage . Both of these l arge rivers are glacial
and support relatively small populations. However, a few
clearwater tributaries of. the lower Copper River and many
clearwater tributar1es of the lower Susitna River, such as the
Talachulftna 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).
Numerous 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 streams of the Alaska
Peninsula and Bristol Bay drainages. Especially large
individuals are found fn the Ugashik and Becharof Lake and Togiak
River drainages (McClean and Delaney, 1978).
Arctic grayling are widely distributed in the remaining Arctic
and sub-Arct i c areas of Alaska, with the exception of the
Yukon-Kuskokwim River delta area (McClean and Delaney,· 1978).
Life Hfstorx SummarY
Arctic grayling usually migrate to spawning sites just prior to
or during spring breakup.
migration of this f i sh
Several factors influence the upstream
including distance separating
overwintering and spawning sites, streamflow and water
temperatures. Tack (1 971) reported upstream movement of fish
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within the ice covered Chena River about 2. 5 wee ks prior to
breakup conditions. Tripp and McCart {1974) reported upstream
mov ement of f i sh in the ice cov ered Mackenzie River to ice-free
tri butaries .
Fish passage can be prevented by ice jams, beaver dams or
waterfalls. The swinming perfonnances of adult and juvenne
Arctic grayling are influenced by f i sh 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 j uvenile fish is not
clear. Tack (1980) related this phenomenon to homing. Juvenile
fish may become imprinted on vi sua 1 , o 1 factory or other
conditions and recognize the spawning area upon maturation.
The establishment of male spawning territories may i nitiate
spawning activity {Kruse, 1959; Bishop, 1971 ; Tack, 1971}. Males
select terri toria l sites for various physical conditions where
spawning wi ll eventually take pl ace .
Grayling territories vary in size with respect to stream width,
wa t er depth, current velocity, channel configuration, spawner
density 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, 1 n a wider ( 2. 4-3.0 m, 8-10 ft) reach of the same
stream, territory dimensions were approx i mately 1.2 by 1.2 m (4
by 4 ft). Tack {1971 ) noted that male f ish territori es in the
out l et of Mineral Lake, Al aska 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 r epe ll ed by lateral displays.
sometimes followed by direct attacks (Tac k, 1971).
Grayl i ng spawn in areas with surface current velocities l ess than
1.4 m/sec (4.5 ft/sec ), varying water depths and relatively
small, unimbedded 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 successful 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 i nto t he substrate where eggs and sperm are
released.
The fertilized eggs sink to the bottom of the stream and adhere
to the substrat e. No actua l redd is constructed but the eggs may
be covered by as much as 5 em (2 i n.) of dislodged substrate.
There i s no parenta l care of the eggs. After spawning, the
female resumes her former rest ing 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 known 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 maturity is variable and genera lly is greater i n the
northern rea ch es of this fish 's range . Grayling have been fo und
to reach sexual maturity at age 2 or 3 in Mich i gan (Creaser and
Creaser, 1935 ) and in Montana (K ruse, 1959) streams. Most fish
in t he 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.
Maximum 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 River, 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 Poul i n, 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 12,900
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 in individuals less than 280 mm (11 in)
fork length (fl) to 2,800 in fish exceeding 305 mm (12 in) fl.
Female fish ranging in fork length (fl) from 331 to 373 mm
(x • 353 mm) from Weir Creek, a small tributary of the Kavik
River, Alaska contained from 4,580 to 14 , 730 eggs (i • 8,482
eggs) (Craig and Poulin, 1974). Ten females (295 to 395 mm fl)
from the upper Chandalar River, Alaska drainage contained 2,330
to 9,150 eggs (i • 4,937 eggs) (Craig and Wells, 1974).
Development of Arctic grayling eggs to hatching occurs very
rapidly (13-32 days) and ·fs 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 Crossman,
1973). The yolk sac is completely absorbed in one to two weeks.
Kruse (1959) examined survival of emergent grayling in Grebe
Lake, Wyoming. He estimated that fish surv ival through the fry
stage was about six percent in one of several inlet streams.
Probable causes of this high mortality were egg dislodgment ,
predation and low fertilization . Water vell)ci ty could also
influence egg and a l evin survival. No redds were constructed
during spawning and ferti 1 ized eggs may not have been covered
with gravel .
Alevins hatching within the gravels probably have higher survival
than those hatching on 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 over its range, but
fish from northern reg i ons generally grow more slowly than f i sh
frt~m southern areas (Craig 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 i n) ( Northwest Territories, 1967 ).
Growth rates of young of the year (yoy) Arctic grayli ng can be
extremely variable among drainages due to differences 1n length
of open water (growing) seasons, temperatures and food supplies.
For e.xample, 49 fish within t he outlet of Chick Lake (along the
Donnelly River, a tributary of the Mackenzie River, Northwest
Territories, Canada) atta i ned a mean fork length (fl) of 49 nm ±
4 nm by 8 July, 1973. In contrast 38 individuals from a small
inlet to Chick Lake were only 20 mm ± 4 nm by July (Tripp and
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McCart, 1974). The Chick Lake inlet had lower water temperatures
and lower benth ic invertebrate standing crops than t hose of the
Chick Lake outlet . Elliott (1980) noted substantial differences
in growth rates among yoy Arctic grayling among small bog and
mountain streams within Alaska. He ascribed those differences to
f ood availability, water temperatures and durations of the
growing seasons.
Arctic grayl i ng are opportun i stic 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
Daphnia and chironomid larvae and pupae. Elliott (1980)
inv esti gated the summer food habits of fish in selected spring ,
rapid-runoff and bog streams cros sed by the Trans-Alaska Pipeline
System (TAPS}. Early yoy fish (less than 3.5 mm fl) consumed
about three different aquatic and terrestrial invertebrate taxa
whereas 1 a rge r yoy fi sh { equa 1 to or greater than 3. 5 mm fl )
consumed up to eight taxa. lnmature chironomids were the most
frequently eaten taxon.
La rger fish consume drifting inma ture and mature aquat ic
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 apparent ly 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 wi thin poo 1 s of the Sagavan i rktok and
Co lville Ri vers, Alaska, contained ephemeropteran and plecopteran
nymphs (Alt and Furniss, 1976 ; Bendock, 1980).
Predation on Arctic grayling eggs and alevins by other fishes
coul d s ig nifi cantly reduce fish producti on. Tack (1 971 ) reported
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whitefish preying upon Arct i c grayling eggs at the outlet of
Mineral Lake, Alaska. Rainbow trout (Salmo gairdneri
Richardson), Arctic char (Salvelinus alpinus (Linneaus)), round
whitefish (Prosopium cylindraceum (Pal lu s)), northern pike (Esox
lucius Linnaeus), longnose suckers (Catostomus catostomus
(Forster)), and other fishes may a 1 so consume Arctic gray li ng
eggs and alevins (Bishop, 1971; MacPhee and Watts, 1976; Alt,
1977).
Spawned-out adult fish may remain within spawning areas or
migrate considerable distances to su11111er feeding areas within
lakes or streams. A spawned-out fish tagged in late June 1972 in
a lake outlet entering the Mackenzie River near Norman Wells,
Northwest Territori es 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
small bog stream, within several weeks after spawning and move to
other areas for feeding (MacPhee and Watts, 1976; Williams and
Morgan, 1974).
Movement of juvenile fish out of spawn i ng streams can occ: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 streams ind i cate that
larger individuals progressively move to faster and deeper stream
reaches for increased cover and food availability (Everest and
Chapman, 1972; Lister and Genoe, 1970). Most of the la;'ger
juveniles were found in relatively fast water with some cover and
areas of low current velocity. Everest a.1d Chapman (1972)
speculated that fish hold in areas of low current velocities and
feed in areas of faster velocity with higher pr ey densities .
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Fry (y oy ) may remain within their natal streams or migrate to
other system.s where they feed and grow during the relathe ly
short Arctic and sub-Arctic open water season (McCart and Bain,
1972; MacPhee and Watts, 1976). The movement of larger, older
f ish out of spawning streams may lessen competition among age
classes. Rearing fish are segregated by size (ag e ) with yoy fish
generally occupying areas of lower current velocities, and more
shallow water. Yearling and older fish generally occupy deeper,
slightly faster areas (Chfslett and Stuart, 1979). Larger fish
have been observed in pools upstream of smaller fish; areas which
probably contain higher densities of prey (Vascotto, 1971).
Fish appear to return to the same sunmer rearing areas. Many
tagged individuals have been recovered the following ,year in the
same areas (Tack, 1980).
Limited stud i es monitoring fish movements in small bog and
mountain st~ams have detected a late summer 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 (C raig,
McCart and Bain, 1972). Arctic grayling must migrate to their
overwintering grounds before the streams become impassable from
low flows or ice buildup. Decreasing water temperatures and
flows associated with the onset of winter probably infl•Jence the
timing of migration to overwintering areas.
The winter distribution of Arctic grayling is more restricted
than the summer distribution . Most bog and many small mountain
and lake inlet and outlet streams become dewatered or freeze
so 1 i d during the fa 11 and winter months. Fish overwinter in
Takes, open pools, spring and glacial streams and in spring fed
mountain streams. Fish have been found in pools of large
i nterior Alaska mountain streams, such as the Chena River .
Spring streams in the Tanana River drainage in interior Alaska
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with 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).
D. 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~chniques, 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.
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II. SPECIFIC HABITAT REQUIREMENTS.
A. Lake Inlet/Outlets
1. Upstream Migration
a. Water Temperature
Water temperatures associated with the upstream
migration of Arctic grayling to s pawning areas withir
inlets and outlets of lakes 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 Fieldi ng Lake as
soon as there was flowing water. Water temperatures of
the inlet during the initial phase of the migration
were not given but water temperatures duri ng the las:
two days of the migration were 0.6°C and 1.1°C (33°F
and 35°F).
Many arctic streams may be impassable to Arctic
grayling prior to spring breakup because of ice
conditions or dewater i ng. 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). Ri pe Arctic grayling have been detected
moving upriver within ice-covered spawning streams such
as Trai 1 Creek. a tributary of the Mackenzie River,
Northwest Territories, Canada (Jessop , Chang-Kue,
Lilley and Percy , 1974 }. Arct i c grayling may ascend
rapid runoff and bog streams , and inlets and outlets of
lakes as soon as flow conditions permit passage to
spawning sites.
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Tack (1 972 ) reported Arctic grayl i ng at the roouth of
the Mineral Lake outlet approx i mately nine days after
water temperature o f 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 funr.tion
of the 1 ength of time requ 1 red to move from their
overwintering habitat .
Kruse (1959) noted that Arctic grayling fn Grebe Lake,
Wyoming, begin spawning migrations into four inlet
streams and the outlet stream (the Gibbon River) when
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 the conclusion of the
spawning migrations in the inlet and outlet streams
were 7.2° to 8.3°C (45°-47°F) and 2 .2° to 8.9°C
(36°-48°F). These temperatures are noticeably higher
than those associated wi th spawning migrations in
Fi elding and Mineral Lakes.
Tack (1980) found no correlati on between roomentary
temperature reductions below 1.0°C and upstream
movement of fish in the Chena River. Diel water
temperature patterns may influence upstream fish
movement in other strea~s (Wojcik, 1954, Warner, 1955;
MacPhee and Watts, 1976).
b. Current Velocity
Arctic grayling usually beg i n spawning migrations to
i nlets and outlets of lakes dur i ng breakup conditions .
Wojcik (1954) first observed fish in one of several
inlet streams of Fielding Lake, Alaska as it began
flowing in mid-May, 1954. Warner (1955) observed the
first fish 1 n the mouth of the same stream the
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following year shortly after open water appeared in
early June. Tack (1972) reported Arctic grayling
entering the outlet of Mineral Lake from the Tok River,
via the little Tok River, during high flow conditions.
Mature and innature individuals were captured by
gillnet as they congregated near the lake's ou t let.
2 . Spawning
a. Water Temperature
Water temperature appears to significantly influence
Arctic grayling spawning. Wojcik (1954) observed fish
spawning in an i nlet tributary of Fielding Lake in
water temperatures of 3.3°C (38°F). Three to four days
later, at the termination of spawning activity, water
temperatures redched 7.8°C (46°F). Warner (1955)
stated that fish spawning began in the same stream the
following spring when maximum water temperatures
approached 4.4°C (40°F).
Tack (1972) reported male fish establishing spawning
territories in the outlet of Mineral Lake when water
temperatures approached 4°C (39 .2°F). Spawning ceased
when water temperatures decreased and remained below
4°C (at 2°-3°C) for two days. Fish resumed spawning
when water temperatures reached 4°C (39.2°F) and
continued for four additional days as water
temperatures ranged from 4° to l0°C (39.2°-50°F).
Grayling were reported spawning in southern tributaries
of the Mackenzie River in water temperatures ranging
from 8° to 16°C (46.4°-60.8°F) by Jessop, Chang-Kue,
Lilley and Percy (1974).
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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
(4 2.8°-51.8°F) (Cuccarease, Floyd, Kelly and LaBelle,
1980). Water temperatures of the streams 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 oc to 10°C
(36°F-50cF} (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:, 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} (Warn er, 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) observed fish spawning over
fine {about 1 em) gravel. Much of the stream was
covered by ice and snow, therefore observations were
made a 1 ong an 0. 18 km ( 200 yd) open reach near the
mouth of the stream and in smaller open areas upstream.
Tack (1971) described the spawning substrate in the
outlet of Mineral lake as being "pea size."
Fish spawning has been observed within rif1:les and runs
of four inlet tributaries to Tyee lake near Ketchikan,
Alaska. Spawning substrate ranged from sand to small
cobble. Coarse sand and gra vel to about 2.5 em (1 in)
in diameter was commonly used by most fish (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 imi ted
variety of substrate and other habitat conditions
within the stream and the need to better determine the
characteri sties 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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most material not exceeding 1.25 em (0.5 in } in
d i ameter, within the outlet of Bench Lake, Alaska
(personal communications , Stephen Hammarstrom, 1981).
Similar size substrate is used for spawning in the
outlet of Crescent Lake, Alaska (personal
communication, Ted McHenry, 1981).
d. Water Depth
Arctic grayling spawn in a range of water depths.
Selection of spawning sites is more strongly influenced
by current velocities and substrate conditions. Fish
in a n inlet to Fielding Lake spawned in "shallow back
waters" (Wojcik, 1954) and in depths of 16 em ( 6 in)
(Warner, 1955}.
Water depths measured in 22 fish terri tories in 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 i nlet 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 most intensively
utilized stream and from 0.05 to 0.46 m (0.17-1.5 ft}
in several smalier, shallower streams with
substantially fewer spawners.
-18-
e. Li ght
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~s e, 1959 ).
Netting at the Bench Lake outlet found few Arct i c
grayling spawning during evening and early morning
hours (persona 1 COIIIIlJni cation, Stephen Hanma rstrom,
1981).
3. Post-Spawning Movements
Spawned-out Arct i c grayl i ng (f i sh hav i ng completed spawning )
co1m10nly vacate spawning s i tes 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 Tra il Creek . Food availab ili ty probably infl uences
post-spawning fi sh movement and distri bution (Tac k, 1980 ).
Small lake inlets may become dewatered by mid to late
sunwner. Adu 1 t spawned-out fish typ i ca 11 y 1 eave these
intermittent streams after spawning and enter lak~s (Kruse,
1959).
4 . Development of Eggs and Alevins
a. Water Temperature
Stream water temperatures influence Arctic grayling
deve 1 opment rates within 1 ake in 1 ets and outlets.
Kruse (1959 ) observed that eggs hatched 19 day s after
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fertilizat · n i n 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 Fi elding 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 r·equired 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 1 iterature.
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 becom ing
exposed, desiccated or frozen.
Downstream migration of yoy fish within inlets of lakes
is probably a response to more sui tab 1 e current
ve 1 ociti es and an abundance of food items in 1 akes.
Newly emerged yoy fish held positions in "quiet water
coves and eddies" during the day along an inlet to
-20-
Grebe Lake, Wyoming (Kruse, 1959). At night th e yay
fish vacated areas of low current velocities and
act iv e ly migrated downstream t o Grebe Lake.
Fry have also been found in shallow margins of Tyee
Lake, Alaska and in small , shallow, pools in the delta
area of i nlet streams of Tyee Lake. High aquat i c
invertebrate production in littoral areas provided
ample food (Cuccarese, Floyd, Kelly and LaBelle, 1980).
Fry have not been observed in mainstem reaches of the
inlet streams.
Arctic grayling fry within l ake outlets typica lly
occupy areas of low current velocities. Yoy fish have
been observed in stream margins with shallow depths and
low current velocities {personal commun i cation, Stephen
Hammarston ~nd 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 assoc i ated current velocities and food
ava i labil i ty. Fry within several inlats to Grebe Lake,
Wyoming occupied shallow, slow hab i tats prior to
migrating downstream to Grebe Lake (Kruse, 1959}. Fry
within Tyee Lake, Alaska occupy shallow littoral
reaches ranging in depth from 2 to 46 em (1-18 in) .
They also occupy shallow, q•.;1et pools in delta reg i ons
of the inlet streams rather than the ma in stem reaches
(Cuccarease, Floyd, Kelly and LaBelle, 1980). Yoy fish
have been observed in shallow margins of the outlets of
Bench Lake and Crescent Lake, Alaska {personal
co111T1Unication, Stephen Hanmarston and Ted McHenry,
1981 ).
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6. Migration to Overwintering Areas
No information was found in the 1 iterature concerning
movements of Arctic grayling within lake inlets and out l ets
to overwintering areas. Fish 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 habitat of fish within lakes. The 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 this time.
Upstream migration of yearling, older juvenile and
adult grayl i ng within Poplar Grove Creek usually
commenced when 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 approached 12° to 14°C (54-57°F) during
late Ma_y to early June. Mean water temperatures during
peak upstream migration of yearling fish (6°-12°C) were
consistently higher than 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 grayl i ng entered Weir
Creek , a tri butary of the Kavik Ri ver, Alaska when
water temperatures were about 5°C. The mi gration
ceased when water temperatures approached l2°C (Craig
and Poulin, 1974).
Upstream migration of adults in Weir Creek was similar:
mature fish were first observed in water of about 4°C
(39°F). Migration ceased when temperatures reached
15°C (60°F) (Craig and Poulin, 1974). Maximum water
temperatures at the termination of the juvenile
upstream fish migration reached 20°C (67°F).
b. Current Ve l ocity and Discharge
The upstream migration of juvenile and adult Arctic
grayling in bog streams usually coincides with high
flows result i ng from snow melt and surface run-off
during spring breakup . The first mature •green• fish
were taken nine days after breakup i n Weir Creek,
Alaska duri ng early June (Craig and Poulin, 1974 ). The
t i me between i nitiati on of flow i n Weir Creek and
arri val of fish was probably due to the di stance from
overw i ntering 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 1n the Tanana River, Alaska in
early April 1953 and 1954 wh i le the streams were frozen
and i mpassable to fish. However , as melt water scoured
furrows i n the ice, the fish began migrati ng upstream.
MacPhee and Watts (1976} trapped adult an d juvenile
f i sh i n 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 juvenile
fish generally began moving upstream in mid-May durin~
the initial stages of the open-water season. The
relatively high discharge at this time ranged from
about 1.3 to 4.0 m3tsec (46-141 cfs). However, the
peak of the juvenile fish migration consistently
occurred at lower flows, about five to ten days after
the peak of the adult fish migration. Juvenile
migration continued for several days after the adult
migration. Discharge at the end of the adult and
j uvenile migrat i ons were generally less than 1.1 m3tsec
(38 cfs}. The yearling fish lagged several days behind
the older juven i le fish .
Upstream migration of juvenile and adult Arctic
grayling in Weir Creek, a tributary of the Kavik River,
Alaska, resembled migrations in Poplar Grove Creek.
(McCart, Craig and Bain, 1974}. Adult fish migrated
upstream in Weir Creek. in 1971 between early 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 and adult Arctic grayling migrated upstream in
Nota Creek, a tributary of the Mackenzie River,
Northwest Territories, Canada, during spring breakup in
May. Dur i ng this time dishcarges decreased from
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1.67 m3/sec to 0.38 m3/sec (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 ability of juvenile
fish to pass upstream and could be responsible for t i.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 correlated to water temperature and flow
conditions are available. Selected studies using weirs and
seines noted the spawning condition of fish (•green•, •ripe•
or •spa~med 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 lea Lake, A 1 as k a i n wate 1·
t emperatures of JO C (44 °F) (McCart, Cra ig and Ba i n ,
1972). The Tea Lake system drains a flat, marsh y arec
and is a bog stream.
Bishop (1971) reported that water temperatures of about
go to l0°C (47°-50°F) appeared to s t imulate f ish
spawning in Providence Creek, Northwest Terri tories.
Maximum water temparatures in Weir Creek during the
Arctic grayling spawning period ranged from 4° to 16cC
(39°-61°F ) (Craig and Poulin , 1974). Max i mum water
temperatures in Nota Creek whtm Arctic grayling wen
1'ripe11 ranged from 3.5° to ll°C (38°-51°F). MaximurT.
water temperatures at the peak of spawning ranged fr>m
4.5 ~ to ll °C (40°-52°F) (personal coJTITiunication,
Derrick Tripp, 1981).
Water temperatures in Happy Valley Creek during Arct ·c
grayling spawning ranged from 4° to 12 °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 coiTITiun i cation,
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 grayiing in Poplar Grove
Creek, Alaska spawned in late May to early June as
streamflows decreased from peaks of about 1.1 m3t sec
(23.2 cfs) in 1973 and 4 .0 m3t sec (8 4.8 cfs) in 1971
(MacPhee and Watts, 1976). Bishop (1971) r e ported fish
spawning in Providence Creek , tributary of the
Mackenzie Riv er, during breakup conditions.
c. Substrate
Arctic grayling appear to use a wide range of substrate
s i zes in bog streams f or spawn i ng. They spawned over
gravel ranging from 2.5 to 3 .75 em (1 -li in ) diameter
in Mainl in e 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
(C raig, McCart and Ba i n, 1972).
Substrate used for spawning in Providence Creek,
Northwest Territories, Canada was gravel mixed with
sand. Fish did not spawn over pure mud , sand or clay
(Bishop, 1971 ).
Fish spawned i n orga nic detritus in Mill ion Dollar
Creek, Alaska, along the Trans-Alaska Pipeline
(p ersonal communication, George Elliott, 1980). The
substrate i n Mill i on Dollar Creek is silt and fine sand
overlain by organic muck (Elliot, 1980).
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3.
d. Water Depth
Observations of water depths used for spawning in bog
streams are limited.
Sev era 1 pairs o·f fish spawned in riffles 5 em ( 2 in)
deep in Mainline Spring Creek, Alaska (personal
communication, George Ell iott, 1980).
e. Light
Fish appa rently spawn only during the day , as observed
by Bishop (1971) in Prov i dence Cre ek.
Post-Spawning Movements
Arctic grayling may migrate downstream irrmediately after
spawn ing. In 1973, post-spawners left Weir Creek, Alaska
within two weeks after spawning. Large juvenile Arctic
grayling also emigrated within two weeks. Tagged adu lt fish
from Weir Creek, Alaska were captured later in t he Kavik
River and the Shav iovik River (Craig and Poulin, 1974).
Rapidly decreasing streamflows probably influenced f ish
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 outmi gration of adult and j uvenile Arctic gray l ing
occurred in Poplar Grove Creek during late May and early
J une after s pawning and when streamflow·s were steadily
declining. Emigration of spa wned-out adults extended from
-28-
mid-May through mid-June 1973 as flows steadily declined
from 1.4 m3/sec (49 cfs) to 0.3 m3/sec (11 cfs). Large
juvenile Arctic grayl ing outmigrated after the adults during
mid-J une. Not 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 Grove Creek until the stream
freezes in ft\ 11 .
Adult grayling ususally leave Nota Cr eek, a bog stream
entering the Mackenzie River, Northwest Territories, Cana da,
within t wo weeks after spawn i ng . Some spawned-out adults
may return to food -rich Nota Lake for short periods of time.
Emigrat ion of large five and six year old juveniles was
followed by younger, smaller individuals through early J uly.
By mid-July young of the ye ar Arctic grayling and some
yearli ng and two year olds occupied Nota Creek (p ersonal
communication, Derrick Tripp, 1981). Decreased living space
and food availability associated with low flows are probabl y
important factors influencing f ish move ments .
Tack (198() sugg ested that the outmigration of juvenile and
spawned-out adult fish may allow yay fish to rear and feed
in natal streams without competition. Adult and j uven :le
fish may rear in other stream systems that are rich in food,
such as spring streams.
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4. De velopment of Eggs and Alevins
a. Water Temper~
Extremely limited information is available concerning
egg incubation and water temperature relationships in
bog streams . Bishop (1971) subjected fertilized eggs
to a range of water temperatures. He determined that
eggs hatched in approximately 14 days at a mean water
temperature of 8.8°C (48 °F).
Arctic grayl ing eggs required 18 to 21 days to hatch in
Nota Creek in water temperatures ranging from 5.5° to
l3 °C (42°-55.5°F} and a mean water temperature of 9.6°
to 10 .3°C (49°-50 .5°F) (personal communication, Derrick
Tr i pp, 1981).
b. Current Velocity and Di sc harge
Spates could dislodge and destroy eggs and severely
reduced flows could lead to desiccation.
Aquatic habitat selected by rearing yoy fish in bog
streams may have current velocities of from 0 to 0.15
m/sec . Larger f i sh generally select faster water.
Elliott (1980} measured mean column velocities at
holding positions of 'ea rly' yoy (~ 35 mm fl) and
'late' tOY (> 35 mm fl) fish i n sel ected bog streams
during Ju ne and Augu ~t 1980. The mean column current
velocities assoc i ated with 'early ' yoy fish were 0 .02,
0.07 and 0.03 m/sec i n Million Dol lar Creek (n = 198},
Pamplin 's Potholes (n = 175), a~d the Tea Lak e i nlet,
Ala ska (n =57), respect ivel y.
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Larger 'late' yoy fish were found in slightly faster
mean current velocities: 0.08, 0.09, 0.14 and 0.1
m/sec. in Pampl i n'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 yoy fry select protected stream areas
where current velocities are extremely low (personal
corrmunication, George Elliott, 1980; de Bruyn and
McCart, 1974; McCart, Craig and Bain, 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 rrm fl)
have been observed in bog streams with slightly greate~
current velocities than yoy fish. The average mean
current velocities occupied by juvenile fish in the Tea
Lake inlet (n = 9) were 0.175 m/sec and in Main l ine
Springs Creek, Alaska (n = 16), 0.196 m/sec. Limited
observations of adult Arctic grayling (250 rrm fl) from
bog streams showed adult fish holding in mean c~rrent
ve l ocities of 0.262 m/sec (Elliot, 1980).
b. Substrate
Rearing fish of all ages were associated with a variety
of substrates inclusing c1etritus, silt, sand, and
gravels. Arctic grayling showed little movement in
small bog streams during July and August following fish
spawning and movement to surrmer 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. Water Depth
Water depths occupied by rearing fish vary
consi derably. Newly emerged yoy 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
Ba in, 1972).
Early yoy fish occup ied small bog streams with depths
from about 0.09 to 0.85 m (0.3-2.8 ft). Late yoy
occupied depths (within the same streams) rang ing 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 fo und 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 with the
onset of winter. Emigra tion of yoy and juvenile fish may
also occur duri ng the summer.
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a. Water Temperatur~
Decreasing water temperatures may influence the
downstream movement of Arctic grayling. Significant
numbers of yoy and juvenile fish move d downs tream in
Weir Creek, Alaska during Se ptember 1973. Min imum
Wdter temperatures during earl y , mid and late September
were about 1°, 4° and 0°C, respectively. Downstream
movement of juvenile fish occurred about one week
before yay fish in Weir Creek. No apparent
relationship co uld be demonstrated between downstream
movement of juvenile or yay fish and water temperatures
(Craig and Poulin, 1974).
Sim ilar downstream movements of yay fish occurred in
Poplar Grove Cr,eek, Alaska. Of the 65,536 yoy fish
observed between July 17 and October 18, 1973, 96 %
( 62 ,680 ) were observed i n the 1 ower reaches during
October (MacPh ee and Watts, 1976). In Poplar Grove
Creek, downstream migration of yoy fish may be related
to stream temperatures.
b. Cu rrent Velocity and Discharge
No relationship could be found between the downstream
movement of j uvenile or yay f i sh 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
qui t e distant from summer rearing areas. Fish overwintering
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areas in the Shaviovik River and suiiiTier 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 migration of adult and juvenile Arctic
grayling. Upst r eam migrants were taken in Vermillion
Creek, Northwest Territory, Canada about one week after
breakup when water temperatures ranged from 0° to 3°C
(32°-37°F) (personal coiiiTiunication, Derrick Tripp,
1981). Tack (1980) also found that water temperatures
of at least 1.09 C (34°F) stimulate upstream movement of
Arctic grayling in large mountain streams like the
Chena River near Fairbanks, Alaska.
b. Current Velocity ana Discharge
Upstream migration of adult and juvenile Arctic
grayling usually occurs during high flows at spring
breakup. A weir placed in Vermillion Creek captured
upstrea ,n migrating adult and juvenile Arctic grayling
for t e n days after peak flows in May of 1973 and 1975.
The M~ck enzie River was covered with ice for up to ten
days af ter breakup occurred in Vermillion Creek during
1973 and 1975 (personal communication, Derrick Tripp,
1981), and observations of fish migration were not
made.
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Observations made und e r the ice along the Chen a River
indicate that Arctic grayling initia te upstream
movement pr ior to breakup (Tack, 1980 ). The fish were
probably movi ng to ups tream reaches of the Ch ena Rive r
or its tributari es. The relative i mpor tance of
streamflow and water temperature i n relati on t o
upstream f ish migration is poorly understood .
2. Spa wni ng
a. Water Temperature
Limited data is available concerning t he relationship
between Arctic grayling spawni ng and stream water
tempe ratures . Spaw ning in the Chena River drainage ha s
been obs erved in water temperatures of soc (Ree d, 1964 ;
personal commu nication , J erome Ha l lbe rg). Tack (1980 )
discussed the possi bility of the 4°C isothe rm
influencing the distribution of spawnin g fish in large
streams like the Ch ena River .
b . Current Velocity and Discharge
Fish s pawn in mountain streams i n a wide range of
current velocities . They have been o bse ~ved spa wn ing
in a n ov erflow slough in the Che na River, Alaska a t
I
re'1atively low current velocities (Reed, 1964 ) and i n
riffles of the Ea s t For k Chena River, Alaska where
surface
ft/sec )
1'191).
current veloc ities approach 1.4 m/sec (4 .5
(p ersonal communication, J erome Hallberg,
Bendock (1 979 ) reported spawning in pools of
the Colville River , Alas ka with negligibl e current.
Nelson (19 54 ) obs erved fis h spawning activity along Red
Rock Creek, Montana in the ends of riffles . Fish
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3.
c.
spawning occurred in similar low flow areas along the
East Fork Chena River, Alaska (personal communication,
Jerome Ha 11 berg, 1981).
Substrate
Arctic grayling use a variety of substrates for
spawning includi~g mud, silt and gravel up to 4 em (1 .5
in) in diameter. Bend ock (1979) observed fish spawning
on silt overlaying gravel in the mainstem Colville
River, Alaska and its tributaries . Spa wn ing substrate
used by Arctic grayling i n the E3st Fork Chena River,
Alaska consists of fine gravels from 0 .75 to 4 em
(0 .4-1.5 in) in diameter (p ersonal 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 sub strate of unknown size but not in pure
silt or sand substrates .
Egg and Alevin Development
No informatio n 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 test wa t 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
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-------------~~-----
Fairbanks, Alaska were used for this study. Results
were expressed as median tolerance 1 imit (T LM ), the
temperatu re at which 50 perc en t of the individuals in a
test die (LaPerri er and Car lso n , 1973).
Results indicated t hat young of the year f ish are
apparently mo re tol e ra nt of re latively high water
ter.pe ratu res t han o lder fish . Th e TLM of yay fish
exceeds 24.5°C, the highest test water temperature.
lndividuals of 10 em fl had TLM val ues of 20 .0 to
24.0°C and fish of 20 em fl has TLM val ues of 22.5 to
24 .5°C . The small fish were acclimated at 4°C and t he
larger fish at 8°C.
Bioassay results indicate that water tempe ra ture
diffe rences of 2°C at relati vely high water
~emperatures can cause very different survival rates of
Arctic grayl i ng . For example, survival of 20 em fl
f ish 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 beca us e fish co uld avoid warm wa ter
temp era t ures by moving to coo ler areas.
b. Cu rrent Velocity and Discharge
Recently emerged yoy fry generally occupy areas with
low cu rrent velociti es. The sma 'l new ly emerged fry
(about 20 ITin total length at 14 cays ) have limi~ed
swimming abilities. Chisl ett and Stuart (1979) noted
that newl y emerged f ry clustered i n shallow, protected
reache s of flood cha nne 1 s , backwat er s 1 oughs and
sidechannel pools of the Sekunka River , Br itish
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Columbia. These fish are found in similar habitats in
the East Fork Chena River, Alaska (personal
communication, Jerome Hallberg, 1981).
Nelson (1954) noted that recently emerged yay fish were
distributed in "backwaters and protected areas ... ,
away from strong currents" within Red Rock Creek,
Montana .
Older yay fi5h crcupy progressively faster waters.
'Early' yay fish (S 35 mm fl) occupied a mean current
ve locity of about 0.07 m/sec (0.22 ft/sec) (n = 183 ) in
5el ected headwater areas of the Gulkana River, Alaska
in early July. Larger yay 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) fou nd yay fish (~ 35 mm fl)
occupying slow current areas of backwater and side
chann els in the Sekunka River and Martin Creek, British
Col umbia during July 1978. Side channels contained
flowing water and were less ephemeral tha n backwater
chann e ls. All yay fish were found in low current
velocities.
Most of the backwater hauitats dewa tered during August
1 ow flows and yoy fish (:> 35 rrm fl) inhabited
sidechannel riffle areas and margins of mainstem
riffles.
By September and October yoy fish occupi ed sidechannel
riffles and margins of mainstem riffles where current
velocities approached 0.8 m/s e:. Yay fish at this time
ranged from 50 to 96 mm fl (Ch 1slett and Stuart, 1979 ).
-38-
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+ {old est aged fish ), occupied larger, deeper pools
than the younger fish.
The distribution of juvenile and adult Arctic grayl iu g
in selected Alaskan mountain streams is similar to the
the distribution in the Sekunka River, British Columbia
where adult and juvenile fish are generally restrict~d
to poo ls and sloughs (Alt, 1978; personal
communication, Jerome Hallberg, Joe Webb, Terence
Bendock, 1981).
Fish will mo\e 1nto shallower, faster riffle areas fo r
more food, such as sa 1 mon an c. whitefish eggs ( Sondoc 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 shdllow areas in sidechannels
and backchannels (Chislett and Stu art, 1979). Yoy fi:;h
have been observed in backwater sloug hs and sha~low
pockets of protected water in the East Fork Chen a
River, Alaska (personal communication, Steven Grabacki,
Jerome Hallberg and Sandra Sonnichsen, 1981 ).
Older fish generally select deeper pools (Chislett et
al.; personal co mmunications, Steven Grabacki, Jerome
Hallberg and Sandra Sonnichsen, 1981).
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d. Cover
Recently emerged yoy fish seek various fonns of
in stream cover when disturbed. Young (17 -45 days old)
fry in sha 11 ow, s i 1 tbottomed backchanne 1 s of the
Sekunka River moved into deeper water with various
ty pes of i nstream cover. Similar aged yoy fish in
sidechannels used substrate interstices and shadows of
boulders for cover. Nelson (1954) 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 colllllonly use logs, boulders and turbulence
for i nstream cover (persona l colllllunication, Jerome
Ha 11 berg, 1981 ).
5 . Migration to Overwintering Areas
Little is known about Arctic grayling migration to
overNi ntering areas. Tack (1980) obse rved a slow downstream
moveme nt of Arctic grayling in the Chena River, Alaska and
compared it to the faster emigration of fish in North Slope
mou ntain streams where winter conditions occ ur very early
(Yoshi hara, 1972 ). Yoshihara (1972) observed many fish
moving downst r eam in the Lupine River i1111led iate ly after
water temperatures reached freezing. Age distribution of
• emigrants in the Lupine River is not known .
6 . Winter Rearing
The distribution of overwintering Arctic grayling is more
limit ed than the summer distribution. Streamflows are lo w,
much or a ll of the stream is ice-covered and s t ream reaches
-40-
ca n be frozen solid during the harsh Arctic an d sub -Arct ic
winters . Ov erwintering areas i n mountain streams include
pools of intermittent or flowing streams (such as Colville
and Chena River, Alaska, respectively) or spring fed streams
which remain ope n during winter months (the lower Shaviovik
River, Alaska).
a. Current Velocity
Curre nt velocities in overwintering sites ar e probably
very low. Conventional current velocity meters do not
functio n at air t emperatures below freezing. Fish
overwinter in ·i ntermittent pools of the Colville River
where current ve l ocity is negligible. In the Hulahula
River overwintering sites had current veloc i ties of
0.15 m/sec (0.5 f t;~:ec ).
b. Water Depth
Fish have been o bserved under the ice in pools of at
l~ast 1.4 m (4.6 ft) depth in the Colville , Chena and
East Fork Chena Rivers. Bendock (1980) 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 Che na River and
the mainstem Chena River, Alaska (Tack, 1980) during
the winter months (Hallberg, personal commu nication,
1981).
Arctic gray l ing have been taken by Kaktovik, Alaska
villagers in the Hu lahula P.iver in late April (Furniss ,
1975) and through the ice in the East Fork Cha ndalar
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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Fork Chandalar River was 1.5 m (5 f t }. These sites are
thought t o be im portant overw i ntering areas for Arctic
grayli ng.
Spring fed mountain streams are often the only sites fn
the North Slope where water remains flowing throughout
the winter. These streams are important overwintering
sites for Arctic grayling. Alt and Furn i ss (1976)
captured adult ~~:tic grayling i n a spring fed pool in
the Frankl i n Bluffs area of t he Sagavanirktok River,
Alaska on May 6, 1975 . The approx i mate 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 4.6 mg /1 in Arctic grayling overwinteri ng
sites in the Colv i lle 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
Furniss, 1976 ).
D. Spring Streams
1. Upstream Migration
Limited i nvestigations indicate that Arctic grayling may
enter springfed streams after spawning (Reed, 1964; Pearse,
1974; Tack, 1980).
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2 . Spawning
Arctic gray li ng apparently do n~t spawn in springfed streams
where low water t emperatures may adversely influence egg and
alevin development (Va nHyning, 19 78).
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 in 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 immature, catchable (by rod and
reel) Arctic grayling moved downstream early i n 1963 in the
Delta Cl ea rwater River. Larger adult fish remained in the
river through most of September. Some tagged adu 1 t fish
were fou nd at the mouth of the Delta Cle al"'\'later River in
late October. I
5. Winter Reari ng I
Arctic grayling apparently do not rear in i nterior Alaska I
springfed streams (VanHyning, 1978), but have been found in
springfed streams in the North Slope (Craig and Poulin,
1974).
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III . HABITAT-ARCTIC GRAYLING QELATION SHIPS
Tables I thr ough VI summarize the reported water temperature l evel s
associated with various life stages and activities of Arctic grayling.
Table VII lists the reported current velocities (or discharges )
associated with different life history stages and Table VIII, the
substrate types used for spawning.
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Table 1: Observed water temperatures associated with upstream migration of
Ar ctic grayling to lake inlets/outlets.
Parameter
Water
Temperature
Observed Values
5.6 -7.8°C
Remarks
Maximum water temperatures
during last 2 days of fish
migration
First mature fish captured
during fish migration
Water temperatures of
several inlets to Grebe Lk.
during ini tia l fish spawning
migrati on ac tivity
19531 1954
Water temperatures of outlet
(Gibbon R.) to Grebe Lake,
Wyoming during initial fish
spawning migratiion activity
Location Reference
Inlet to Fielding Warn er (19 55 )
Lake, Alaska
Outlet to Mineral Tack (1972)
Lake. Al,;ka
Four inlets to Kruse (1959)
Grebe Lake.
Wyomtng
Gibbon River,
outlet to Grebe
Lake, Wyoming
Kruse (1959)
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Table If: Observed water temperatures associated with upstream migration of
Arctic grayling within bog (tundra) streams.
Parameter
Water
Temperature
Observed Values Remarks
Maximum water temperatures
during early to late stage
of adult fish spawning migra-
tion . Incomplet e fish sampling
due to high flows, 1973.
Ma xi mum water temperatures
beginning and end of adult
fish upstream migration, 1971.
Weir placed in stream after
fish migration began.
Ma ximum water temperatures
from start to near term1n ~
t1on of juvenile fish
upstream migration, 1971.
Initial average water tempera-
tures of Poplar Grove Creek
during start of adult, sub-
adult, juvenile fish upstream
migration during May 1973 -1975.
Average water temperatures
at end of adult, sub-adult,
juvenile fish migration in
June 1973-1975.
Location
Weir Creek,
Tributary to
Kavfk R., Alaska
Happy Valley Crk,
tributary t!l
Sagavanerktok
River, Alaska
Happy Valley Crk,
Sagavanerktok
River, Alaska
Pop l ar Grove
Crk, tributary
to Gulkana R.,
Alaska
Poplar Grove
Crk, tributary
to Gulkana R.,
Alaska
Reference
Craig and Poulin
(1974)
McCart, Craig and
Bain (19 72)
McCart, Craig and
Ba1n ( 19 72)
MacPhee and Watts
(1976)
MacPh ee and Watts
( 1976)
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Table IJI : Observed wat er temperat ures associated with
Arctic grayling s pawning in mountain streams.
Parameter Observed Valu es Remarks
Wat er 5.6°C In s tantaneous water tempera-
Temperature ture of slough where fish
were obse rved s pawn i ng
Instantaneou s wate r tempera-
ture during fish s pawning
activ1 ty
ln stanta neou ~ water tempera -
ture during fish s pawning
activity
Instantaneous water tempe ra -
t ure during fish spaw ning
acti vity
In stantaneo us water tempera-
t ure during fish s pawning
ac tivity
l ocation Reference
Slo ugh along Reed (1964)
Chena Riv e r,
Ala sk a
Riffle. E. Fork Hall be r g
Chena R 1 ver. (p ersona l
Alaska conmuni cation)
Seab ee, Rainy, 8E;nd ock
Foss i1 Crks , (1970)
tributaries
Co lville R .• Al aska
Nuk a River, trib-Ben do ck
utary, Colville (1 979)
River, Ala s ka
Aniak R, trib-Alt (19 77)
uta ry, Kuskokwim
River , Ala ska
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Tabl e IV: Observed water temperatures asso c iated with
Ar ctic gray l ing spaw ning in l ake inl ets/out l ets.
Parameter
Water
Temp erature
Observed Valu es Remarks
Initi al f i s h s pawning
activity occ urred
Fi s h spa wn ing act i vity was
completed at this temperature
3-4 days after co llJTlen ceme nt
of spawn ing act iv ity.
Firs t s pawning activ i ty
observed
Fi s h s pawnin g ce~se d a s
water temperat ure dropped
bel ow 4°C
Fi s h spa wni ng lc tivity
res ume d and l asted 4
additional days
Fi s h actively s pawning
fi s h actively spa wni ng
Fi s h active ly s pawni ng
Fish actively spawning
Loc ation Reference
Inl e t to Fielding Wojc ik (1954)
Lake, Ala ska
Inl e t to Fie lding ~oj c ik (195 4 )
Lak e , Al as ka
Outlet to Min e r al Tack {1972)
Lak e , Alaska
Outle t to Min era l Tack {197 2 )
Lake, Alaska
Outl et to Mine r a l
lake, Alaska
Severa 1 in 1 ets
to T_yea Lake ,
Alaska
Four inl ets t o
Grebe lk , Wyoming
Outlet to Gr ebe
Lak e, Wyoming
Gibbon River
Inl e t to Agne s
lake , Montana
Ta ck (1 972 )
Cuccarease, Fl oy d
Kell y , LaBe ll e
(1980 )
Kr use {1 959)
Kruse (1959)
Brow n (1938)
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Table V: Observed water temperatures associated with
spawning of Arctic grayling within bog streams .
Parameter
Water
Temperature
Observed Values Remarks
Direct spawning observation,
one temperature reading
Fish spawning activity seemed
to be related to these wate r
temperatures.
Maximum water temperatures
based on occurrence of rfpe
and spawned-out Arctic gray-
ling captured by weir
Maximum water temperatures
during Arctic gra y ~~ng
spawning activity ba sed on
condftfon of fish in weir
Location
Outlet Tyee Lk,
Alaska
Reference
McCa rt, Craig and
Bafn (1 972)
Providence Crk, Bishop (19 71)
tributary to
Mackenzie River
Northwest Territory
Weir Creek, Craig and Poulin
Alaska (1974)
Nota Creek,
Alaska
Tripp (p erCio na l
conmun i cat ton .
1981)
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Table VI: Observed water temperatures associated with
Arctic grayling egg and alevin development.
Parameter
Water
Temperature
Observed Values
6.1 ° -9.4°C x::: 7.JOC
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, Wyoming
Inlet to Fielding
Lake, Alaska
Somewhere in
Alaska
Ref e r e nce
Kru se (1 959)
Wo .i c i k (1954) ;
Warn e r (1955)
Wojcik (1954)
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Tabl e Vlf: Relationship of current velocity (or di scharge)
to specific li fe history stages.
Activity
Spawning
Early Development
Juvenil e Rearing
Adult Summer Habitat
Adu l t Winter Habita t
Up s tream Migration
Current Rate or Flow
s low, shall ow backwater
1.2 m/s
.34 m/s to 1.46 (x =.79)
shall~w riff les
1.1 m3/sec
4.0 m /sec
low flow and riffles of 1.4 m/s
negligible
.02, .07, .03 m/s
sha ll ow, prote cted areas
shallow pools
s hall ow pools
.08 to .195 m/s
.8 m/s
.26 m/s
open areas
at breaku~
1. 3 -4 m /s
2 m3ts3-, 20 m3 /s
1.67 m /s
at breakup
at breakup
high flow
Reference
Wo jci k • 1954
Wa r ne r. 1955
Tack, 197 1
Elliott, 1980
Mac Phee and Wa tts, 1976
Mac Phee and Watts , 1976
Ha 11 berg, 1981
Bendock, 1979
Elli ott, 1980
Chis lett and Stuart, 19 79
Cuccarese et al., 1980
Ha111na rs ton. 1981
Elliott, 1980
Chis lett and St uart , 1979
E 11 i ott , 1980
Chislett and St uart, 1979
MacPhee and Watt s , 1976
MacPhee and Watt s , 1976
McCart et a 1. , 1974
Tripp, 1981
Wojc ik, 1954
Warne r, 1955
Tack, 1972
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Table VII I : Reported substrate types used for spawning.
Substrate
Fine gravel (1 em )
"pea -size"
sand to small cobble
sand -gravel
sane, 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, gravel(~ 4 em)
gravel, .75-4 em
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Reference
Warner, 1955
Tack , 1971
Cucc arease et al., 1980
6rown, 1938
Kruse, 1959
Hammarstron, 1981
McHenry, 1981
Elliott, 1980
C14 aig et a l., 1972
Bishop, 1971
Elliott, 1980
Elliott, 1980
Bendock, 1979
H a 11 berg , 1981
['/. DEF ICIENCI ES W DATA BASE
Facto r s i nf luenci ng the migration of adu lt and j uven il e Ar ctic
grayling from overwintering areas to spawn i ng streams is apparently
infl uenced by flow and wate r temperature conditions. The timing of
adult and juveni le fish migrations are not understood; the juvenile
f i sh run l ags several days to several wee ks behind the adult fish in
ce rta i n areas.
Information about selection of s ite s in mountain, lake inlet and
outlet, bog and spring streams i n relation to current velocity , water
depth and substrate i s limited. Most of the obse r vations were made i n
l ake inlets and outlets and mountain streams.
Habitat selection by spawning Arctic grayling is influenced by at
least three variables -substrate, water depth and current velocity -
which collec ~ively determine t he habitat quality. For example, Arctic
grayling may be excluded from spawning areas by excessive cu rrent
ve l ocities desp ite acceptable substrates and water depth s. Th ere i s
limited i nformat io n on the i nteraction of vari ous physica l parameters.
Factors infl uencing the survival and development of eggs and alevins
are not well understood. Studies indicate that egg dislodgeme nt by
other spawning fish and spates may be a major cause of morta 1 i ty .
Minimum water temperatures required for successful egg development are
not known; however, water temperatures above 6°C (42°F) have been
recoiTITiended.
The movement of Arctic grayl ing to overwintering areas appears to be
i nfluenced by f low and wate r temperatures associated with the onset of
wi nter. This emigrat ion may be of short duration or extend over
several weeks . There is very little information about Arctic gra yling
overwintering ar eas. Fish oveNi ntering in the North Slope are
li mited to open water areas or to streams whi ch do not completely
freeze. Fish oveNintering in i nterior Alaska and Canada may remain
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in spri ngfed and gl acier streams. It is not known whether young of
the year Arct ic grayling burrow into interstices within cobbl e an d
rub bl e as water temperatures approach freezing.
The age structure of Arct i c grayli ng 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 lar~e juvenile and adult
fish . Fish may be either sedentary or nomadic during the open-water
rearing s eason. Expla nations for fish emigration are specula t i ve but
1 iving space and food availability are probably influential. Studies
of j uven il e and adult Arc tic grayling habitats focused on water depth ,
current veloci t i es and substrate i n small bog and mountain streams.
Methods of describing water depth, cu r rent velocity and substrate
characteristics varied among studie ~. Some invest ·gators measured
current velocity at t he site of s pawning Arctic grayli ng; others
estimated surface curren t velocities. Substrate si ze classi f ication
systems also varied; few researchers evaluated su bs trate i mbeddedness
at spawning sites.
The effects of water t emperatures 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 S~UDIES
In depth investigations are needed to detGnnine the relationships
between specifi c chemical and physical features of aquatic habitats
and grayling growth and behavior. For example, investigations should
be designed and conducted to assess the f actors which influence
development and survival of Arctic grayling eggs and alevins in bog.
lake inlet and outlet. spring and mountain streams. In vest igations
should consider egg dislodgement. predation, desiccation, and diel
drift of emergent fry.
Water t emperatures and ice conditions during grayling egg and alev i n
development 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 t he development and s urvival of eggs and
alevins. Water temperatures at or below threshold level s co uld cause
morpholog i cal deformities, slow deve lopment rates and high mortality
among eggs and alevins. The se investigations could explain the
apparent avoidance of spring streams by spawning Arctic grayling.
Weirs could be used to sample grayli ng from bog. mountain, spring and
lake inlet and outlet st reams to relate r esiden ce time and migration
to stream flow. water temperature and other physical and chemical
stream factors.
Unique tags on upstream migrating adult and juvenile f i sh would
provide specifi c migration data. Data from consecutive years could be
used to determine Arct ic grayling homing to specific spawning and
rearing streams.
Arctic grayling spawning sites should be studied more extensively in
northern latitudes. Characterist i cs 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 li months old.
Comprehensive sampling of fry would determine movement of yay Arctic
grayling . Weirs of small mesh screen could be used to monitor yoy
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 physical and
chemical habitat components. Weirs could remain in spring streams
which remain ice-free to monitor the presence of fish .
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Investigations shoul r.l be conducted to determine open-water, lotic
microhabitat selection by juvenile and adult Arctic grayling using
techniques similar to those described by Everest and Chapman (1972 ).
l"hese i nvestigations should describe water depths, current velocities,
substrate, proximity to nearest fish and instream cover. Snorkel i ng,
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
grayling 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.
Ori f t and substrate samp 1 i ng of invertebrates can be used to assess
food availability and its relationship to the distribution of Arctic
gray li ng.
Studies of juvenile and adult Arctic grayling overwintering habitat
should continue. Gillnets, SCUBA and other techniquP.s could be used
to i nvestigate these overwintering habitats.
Fish passage studies should be conducted to assess the ability of
juvenile and adult Arctic grayling to ascend culverts and other high
current velocity areas . MacPhee and Watts (1976) determined 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 possibil i ty 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
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-58-
Brown, C. 1938. Observations on the life history and breeding habits of
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Henshall , J . 1907. Culture of the Montana grayling. U.S. Fisheries
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-60-
J essop, Ch ang-Ku e, Lill ey and Percy. 1974. A further evaluation of th e
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Mac Phee, C. and F. Watts. 1976. Swirrming perfonna nce of Arct ic grayl i ng
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