FRESHWATER FISHES OF ALASKA
Their Biology, Distribution and Value
by
James E. Morrow
University of Alaska
Fairbanks, Alaska 99701
Grant # R803845-01-0
Project Officer
Eldor W. Schallock
Arctic Environmental Research Station
Corvallis Environmental Research Laboratory
College, Alaska 99701
CORVALLIS ENVIRONMENTAL RESEARCH LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U.S. ENVIRONMENTAL PROTECTION AGENCY
CORVALLIS, OREGON 97330
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DISCLAIMER
This report has been reviewed by the Corvallis Environmental Research
Laboratory, U. S. Environmental Protection Agency, and approved for publi-
cation. Approval does not signify that the contents necessarily reflect the
views and policies of the U. S. Environmental Protection Agency, nor does
mention of trade names or commercial products constitute endorsement or rec-
ommendation for use.
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FOREWARD
Effective regulatory and enforcement actions by the Environmental Pro-
tection Agency would be virtually impossible without sound scientific data on
pollutants and their impact on environmental stability and human health. Re-
sponsibility for building this data base has been assigned to EPA's Office of
Research and Development and its 15 major field installations, one of which is
the Corvallis Environmental Research Laboratory (CERL).
The primary mission of the Corvallis Laboratory is research on the effects
of environmental pollutants on terrestrial, freshwater, and marine ecosystems;
the behavior, effects and control of pollutants in lake systems; and the de-
velopment of predictive models on the movement of pollutants in the biosohere.
This monograph assembles and organizes under one cover a vast amount of
information about freshwater fishes of Alaska that was previously scattered in
various, and at times obscure, sources. This valuable contribution to the
knowledge of fishery biology will be utilized by many scientific disciplines.
A. F. Bartsch
Director, CERL
m
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ABSTRACT
A summary of knowledge of the freshwater fishes of Alaska is provided.
Covered are 56 species in 34 genera and 15 families, including strictly fresh-
water species, anadromous forms and those which normally are marine but which
occasionally or regularly enter fresh water. For each species, a brief de-
scription is given, followed by discussion of its range and abundance, its
general biology and its importance to man, as far as presently known.
This report was submitted in fulfillment of Grant No. R803845-01-0 by the
University of Alaska under the sponsorship of the Environmental Protection
Agency. This report covers the period 1 June 1975 to 31 December 1976 and was
completed as of 31 December 1976.
IV
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CONTENTS
Foreword i i i
Abstract. iv
Acknowledgments ix
Introduction 1
Lampreys - Fami 1 y Petromyzonti dae 2
Paci fi c 1 amprey, Enjbosphenus trvdentatus 3
Arcti c 1 amprey, Lampetra j apon i ca 5
River lamprey, Lampetra• ayresj_ 8
Western brook 1 amprey, Lampetra richardsoni 9
Sturgeons - Fami ly Acipenseridae 11
Green sturgeon, Acj_P_ense_r_ me^ijro^tn_j_ 12
White sturgeon, Acjjpensejr tjransmonjbajiu^s ., 14
Herrings - Family Clupeidae 17
Paci f i c herri ng» Clupea harengus pallasj 18
American shad, A1 os a sap i d i s s i ma 21
Salmons, Trouts, Charrs, Whitefishes and Grayling - Family
Salmonidae : 25
Incormu or Sheefish, Stenpdus leucichtjiys 26
Least cisco, Coregonus sardine 11 a 30
Bering cisco, CDregpnus laurettae 33
Arctic cisco> Coregonus autumnal i s 35
Pygmy whi tefi sh, Pros opium coulteri , 37
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Round whitefish, Pros opium cylindraceum. 39
Broad whitefish, Coregonus nasus. 41
Humpback whitefishes , "Coregonus clupeaformis complex" 44
Alaska whitefish, Coregonus nelsoni .. 44
Humpback whitefish, Coregonus pjdschian 47
Lake whitefish, Coregonus clupeaformis 48
Cutthroat trout, Sajmo clarkl , 51
Rainbow trout, Salmo gairdneri 55
Brook trout, Salvelinus fontinalis 60
Lake trout, Sal veljnjjs^ namaycush 63
Arctic charr, Salvelinus alpinus ,. 68
Do!ly Varden, Salvelinus malma.. 71
Angayukaksurak charr, Salvelinus anaktuyu_k_ensis 75
Pink salmon, Oncorhynchus gorbuscna. 77
Chinook salmon, Oncorhynchus tshawytscha. 81
Chum salmon, Qncorhynchus keta 85
Coho salmon, Oncorhynchus ki s u ten 89
Sockeye salmon and Kokanee, Oncorhynchus nerka. 93
Arcti c grayl i ng, Thy mall us a re tic us. 98
Smelts - Family Osmeridae 101
Longfin smelt, Sp i ri nc hus thai ei cji t hys 102
Pond smelt, Hypomesu_s olid us .104
Surf smel t, Hy pomes us pretiosus , 106
Eulachon, Thaleichthys paci fi cus. 109
Rainbow smelt, Osmerus mordax 112
Mudminnows and Blackfish - Family Umbridae ...117
VI
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Alaska blackfish, Dal 11 a pec to rails 118
Pikes - Family Esocidae 121
Northern pi ke, Esox lucius 122
Minnows - Family Cyprinidae 128
Lake chub, Couesius pi umbeus 129
Suckers - Fami ly Catostomidae 132
Longnose sucker, Catosjoinijs catostomus. 133
Trout-perches - Fami ly Percopsidae 137
Trout-perch, Pejxopsj^ omisconiaycus. 138
Codfishes - Family Gadidae , 141
Burbot, Lota Iota 142
Arcti c cod» Boreogadus saida 145
Paci fi c cod, Gadus macrocephalus. 148
Saffron cod, Elegirius gracilis 150
\
Sticklebacks - Family Gasterosteidae 153
Threespine stickleback, Gasterosteus aculeatus 154
Ninespine stickleback, Pungitius pjngitius. 158
Surfperches - Fami ly Embi otoci dae 162
Shiner perch, Cyrnatpgaster aggregate^. .163
Sculpins - Family Cottidae 166
Pacific staghorn sculpin, Leptocottus ajrmatus 167
Slimy sculpin, Cottus cognatus. .......169
Prickly sculpin, Cpt_tus_ a_s_pe_r 173
Coastrange sculpin, Cottus aleuticus 176
Fourhorn sculpin, Mypxocephalus quadricornis. .....178
Sharpnose sculpin, C1 i n oc o 11 us a cut ice ps ....182
vii
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Flounders - Family Pleuronectidae 184
Arcti c f 1 ounder, Liopsetta glacial is 185
Starry f 1 ounder, Platlchthys stellatus. 187
Bibliography .191
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ACKNOWLEDGEMENTS
The author wishes to acknowledge the financial support of the U. S. En-
vironmental Protection Agency which made possible a major portion of this work.
In particular, the author is appreciative of the enthusiastic encouragement
provided by Mr. Eldor W. Schallock, project officer; Mr, Richard Latimer, farrier
Director of EPA's Arctic Environmental Research Station at College, Alaska; and
Dr. Ronald Gordon, Acting Director of the Arctic Environmental Research Station.
Mr. Kenneth Coyle, Marine Sorting Center, University of Alaska Museum, provided
invaluable assistance by translating some of the Russian literature.
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SECTION 1
INTRODUCTION
Since 1961 the author has studied the taxonomy, biology and distribution
of various Alaskan freshwater fishes. These studies, combined with those of
graduate students at the University of Alaska and related projects of under-
graduates, resulted in the accumulation of extensive notes and manuscripts
based on original investigations and a comprehensive file of references.
With the rapidly increasing industrialization of Alaska, an oil pipeline
nearly ready to go into operation, a qas pipeline proposed and the increasing
need for environmental impact statements, it seeired imperative to make this
accumulation of data on fishes available. Those who plan projects, those who
execute them, and those who determine feasibility, cost/benefit ratios, and
probable environmental effects need as much accurate information as possible.
This monograph includes information on all fishes known to occur requ-
larly or occasionally in the fresh waters of Alaska. Some species are
strictly freshwater forms, never entering the sea. Others are anadromous,
spending part of their lives in salt or brackish water and part in fresh
water. Still others are normally marine forms which regularly or occasion-
ally enter rivers. A few of this last group have not yet been recorded from
fresh water in the State of Alaska but are included because they are known
from this environment in other parts of their ranges. Discussion for each
species includes a technical description, the distribution and abundance, a
summary of the biology and behavior as far as known and a brief discussion of
its importance to man. No attempt has been made to provide keys or other
methods for identification, or a glossary of ichthyological terminology,
since this information is available in McPhail and Lindsey (1970), Scott and
Crossman (1973) and'Morrow (1974).
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SECTION 2
LAMPREYS—FAMILY PETROMYZONTIDAE
The lampreys are a primitive group of fish-like animals. Characteris-
tically, they have no jaws, the mouth beinq a round, suctorial opening on the
ventral side of the head and containing numbers of horny teeth. There is
only a single nostril, located on the dorsal side of the head just before the
eyes. The gill openings appear as a row of seven round holes on each side of
the head, behind 'the eyes. The body is long and eel-like, with no paired
fins. The dorsal fin is in two parts, the anal one, and both are more or
less confluent with the caudal fin. The dorsal fin is larger and better
developed in males than in females, while the anal fin is well developed in
females but is almost non-existent in males. The skeleton is entirely of
cartilage and contains no bone.
Some species of lampreys are parasitic on fishes. They attach themselve^
to the host with the oral sucker, rasp away the skin and scales with their
horny teeth and suck the body juices. Parasitism appears to be found chiefly
in species which are anadromous and is confined almost entirely to the marine
phase of the life history. Non-parasitic species are much smaller and gener-
ally have blunt rather than sharp teeth.
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PACIFIC LAMPREY
Entosphenus tridentatus (Gairdner)
DISTINCTIVE CHARACTERS
The Pacific lamprey is characterized by the presence of three (rarely
only two) large, sharp teeth on the supraoral bar, and three sharp cusps on
each of the central lateral tooth plates.
DESCRIPTION
Body elongate, eel-like, more or less cylindrical anteriorly, compressed
posteriorly. Depth about 1% of total length. Head moderate, its length
about 15% of TL. Snout rounded, its length about 66% of head length. Eye
round, small, its diameter about 191 of head length, about equal to postorbi-
tal distance (from posterior edge of eye to anterior margin of first gill
opening). A single nostril on median dorsal line just in front of eyes, in
a short tube. Oral sucker round to oval. Gill openings 7, their vertical
height about 60% of eye diameter, spaces between openings averaging a little
less than half eye diameter (43-53%). Fifty eight - seventy four myotomes
between last gill opening and anus. Anus at anterior end of anal fin, A
slender, fairly prominent genital papilla present in males.
Teeth
Supraoral bar with three (rarely only two, the center one reduced or
absent) large, sharp points. Lateral tooth plates with three sharp points on
each of the central plates. A row of posterial teeth present. Tongue with
15-25 small, fine teeth, all about the same size. Infraoral bar with 5-8
points.
Fins
Two dorsal fins, placed far back on body, the anterior one lower and
shorter than the posterior, the two fins separated by a notch. Dorsal fins
higher in males than in females. Caudal fin more or less pointed, lower lobe
larger than upper, the lobes joined to dorsal and anal fins. Anal fin rudi-
mentary, virtually absent in males. No paired fins.
Color
Adults fresh from the sea are usually blue-black to greenish above,
silvery to white below. Spawning adults become reddish brown.
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Size
Reaches a length of 760 inn (30 in) and a weight of about 1/2 kg (1 Ib).
RANGE AND ABUNDANCE
The Pacific lamprey has been recorded from Alaskan waters from Nome
(UAFC #1119), St. Matthew Island, Wood River, Unalaska Island and southward.
It is also known from Hokkaido Island, Japan. One was taken from the stomach
of a whale captured off Bering Island, near the Kamschatkan Peninsula. On
the American side of the Pacific, the range extends from the northern Bering
Sea south to southern California. This lamprey is fairly abundant throughout
its range.
HABITS
Adults return from the sea and run upstream in fresh water in the spring
and summer, showing considerable ability to overcome obstacles such as dams
and waterfalls. In this, they hold on with their oral suckers to the smooth
face of the obstruction, then let go and swim vigorously upward and catch a
new hold. This procedure is repeated until the obstacle has been passed. At
the time of upstream migration, the lampreys are not yet sexually mature.
They spend the following fall and winter in the streams, often burrowing into
the stream bottom. Spawning takes place in the spring following the migration
into fresh water. The spawning areas are usually in the upper reaches of the
streams, most often in fine gravel at the upper ends of riffles. The nest is
a shallow pit, about 550 mm in diameter, and is dug by both males and females.
In this activity, each lamprey affixes itself to a stone by means of its oral
sucker, then both members of a pair thrash about with their bodies, disturbing
the fine sand and debris which are carried downstream by the current. They
may also pick up small stones with their suckers and carry them out of the
nest.
In the spawning act, the female attaches herself to a stone and the male
fastens his sucker on the head of the female. The two arch and twist their
bodies towards each other, the male more or less wrapping his body around
that of the female. This brings the male's genital papilla in close proximity
to the genital pore of the female. The two then vibrate and eggs and sperm
are released. A large female may produce up to about 100,000 eggs. The
adults die soon after spawning.
The small, whitish eggs fall to the bottom of the nest, where the
parents leave them. Hatching occurs a week or two after spawning, the time
depending largely on temperature. The newly hatched larvae (ammocoetes) are
blind and have no sucking disc. They remain for several years more or less
buried in the fine sand and mud of the stream bottom, feeding by filtering
plankton and tiny particles of algae and debris from the water. Finally, they
metamorphose into the adult form and go to sea. A number of landlocked popu-
lations are known, in which the adults remain in fresh water throughout their
entire lives (Fletcher, 1963).
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During the marine life, the adults are parasitic, attaching themselves
by means of the oral sucker to the bodies of larger fishes. With their sharp,
horny teeth, they rasp through the scales and skin of the host and feed on the
blood and body fluids. They will probably attack any fish of suitable size,
but are particularly known to parasitize the various species of salmons and
trouts. Adults do not feed in fresh water. In turn, the lampreys are fed
upon by many predacious fishes such as pike, trout, salmon and so on.
IMPORTANCE TO MAN
The Pacific lamprey is of greatest importance as a parasite of more
desirable fishes. It is not known whether this lamprey actually kills a sig-
nificant number of fishes, but certainly a fair percentage are found bearing
scars which may be attributable to IE. tridentatus. The Pacific lamprey is
not marketed, but there is no reason to think that it is any less edible than
the European lampreys, which have long been considered a delicacy.
ARCTIC LAMPREY
Lampjtra japomca (Martens)
DISTINCTIVE CHARACTERS
Two large teeth on the supraoral bar, only two points on the central pair
of lateral tooth plates, and the presence of a row of posterial teeth distin-
guish L_. japonica.
DESCRIPTION
Body elongate and eel-like, depth about equal to width anteriorly, be-
coming compressed posteriorly, depth about 7% (5.8-9.1%) of total length.
Head short, about 12.4% (11.7-13.31) of total length. Snout rounded an-
teriorly, its length about 66% (55-68%) of head. Eye round, its diameter 2035
(15-281) of head, 1351 (114-152%) of postorbital distance. The eye is
relatively larger, over 20% of head, in specimens under about 150 mm total
length, while in individuals over about 200 mm the eye is only 15-17% of
head length. The eye-postorbital relationship, however, appears to remain
constant. Nostril on median dorsal line, just before eyes, in a short,
fairly prominent tube. Oral sucker more or less round. Gill openings 7,
approximately round, their diameter about 36% (22-62%) of eye. Spaces be-
tween gill openings averaging a little more than half eye diameter. Sixty-
five - eighty myotomes between last gill opening and anus. Anus^located at
anterior end of anal fin. A slender, fairly prominent genital papilla
present in males, barely developed in females.
Sensory pores on head region, each in a small tubercle, as follows:
Laterally on each side a row of about six pores extending backward from tip
of snout; a row of about nine from ventral margin of eye dorsally and an-
teriorly; a row of about seven posteriorly from postero-dorsal margin of eye.
On dorsal side, a transverse row, often broken, of four to six pores across
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top of head at level of posterior margin of eye; two rows of widely spaced
pores extending backwards from just anterior to first gill opening, the more
dorsal row reaching somewhat behind last opening, the more lateral row reach-
ing the last opening. On ventral side5 a fairly prominent row more or less
encircling posterior part of oral sucker; two rows from behind oral sucker to
about the level of the fourth gill opening.
Teeth
Supraoral bar normally with two large points (rarely a small third cen-
tral point). Lateral tooth plates with two points on central pair. A row of
posterial teeth present. Tongue with a large median tooth point and six or
seven small points on each side. Infraoral bar with 5-10 usually blunt
points.
F i ns
Two dorsal fins, anterior one lower than posterior, arising far back on
body, the fins separated from each other by at least a definite notch, higher
in males than in females. Caudal fin with lower lobe somewhat larger than
upper, the fin joined to both dorsal and anal fins. Anal fin small, in males
represented only by a low ridge.
Color
Variable, from brown to olive or grayish above, paler beneath.
Sjze
Known to reach a length of over 600 mm (about 2 ft), but generally rather
smaller. The strictly freshwater, non-anadromous form rarely exceeds 180 mm
(7 in).
RANGE AND ABUNDANCE
In Alaska, the Arctic lamprey ranges from the Kenai Peninsula north to
the Arctic coast and east as far as the Anderson River in Canada. It has
also been found on St. Lawrence Island. Inland, it ranges up the Yukon River
into the Yukon Territory, and is also present in the Kuskokwim and Tanana
river drainages. World-wide, this species is almost completely circumpolar,
from Lapland south to the Caspian Sea, eastward to Kamschatka, south to
Korea, and eastward across North America to the Northwest Territories of
Canada.
Abundance varies locally and seasonally. Large numbers are known to
ascend the lower Yukon River, and the population in the Naknek River also
seems to be relatively large. A small population is known to spawn in the
Chaktanika River a few kilometers below the Elliott Highway bridge.
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HABITS
Relatively little is known of the life history of this species. In the
Naknek River of Alaska, and doubtless in other areas as well, the population
appears to be composed of both anadrotnous and strictly freshwater forms. The
Chatanika population is probably strictly freshwater, while those of the lower
Yukon seem to be mostly anadromous. Both the anadromous and the freshwater
forms are reported to be parasitic. Spawning takes place in the spring. Both
males and females engage in nest-building, removing pebbles and small rocks
from the stream bottom by picking them up with their suckers and depositing
them downstream from the nest, or by attaching themselves to a large rock and
thrashing violently, thus raising small stones and silt which are carried
downstream by the current. Nests are located out of the main current, in
water from a few centimeters to nearly a meter deep, flowing at between 0.16
and 0.3 meters per second. In the breeding act, the male attaches himself
by means of his sucker to the head of the female. The two arch their bodies
and the male wraps himself around the female so that his genital papilla is
close to her genital pore. Both members of the pair vibrate rapidly and eggs
and sperm are extruded into the nest. Occasionally, two males may mate
simultaneously with a single female. Females will mate several times before
their egg supply is exhausted, usually with several males. A large female
may produce more than 100,000 eggs.
Time for development of the eggs is probably a few weeks. The larvae
(ammocoetes) apparently spend at least one and possibly two years in this
stage, for in the Naknek River two distinct size groups have been observed.
Upon metamorphosis, the young adults descend the stream either to the sea or
to lakes or larger rivers. Adults are parasitic on many species of fishes,
attaching themselves to their hosts by means of the oral sucker, rasping
through the skin and scales and sucking the blood and body fluids. Host
species which have been recorded include sockeye salmon (Oncorhynchus nerka),
pink salmon (0_. gorbuscha), chum salmon (0_. keta), chinook salmon (0. tsha-
wytscha), starry flounder (P1 at ichthys st e 11 at.us), pygmy whitefish TProsppium
cpujten') and three-spine stick!eback (Gastergsteus aculeatus) (Berg, 1948;
Birman, 1950; Heard, 1966; McPhail and lindsey, 197QTTFTs" probable that
this lamprey will parasitize any species of fish, providing only that the in-
dividual is of suitable size.
Length of life is unknown.
IMPORTANCE TO
The Arctic lamprey is probably of no direct importance to man. It is
possible that lamprey parasitation may have a bad effect on the health of the
host, but, at least in the Naknek River, there is no evidence that this species
has had an adverse effect on the fisheries (Heard, 1966). In the Yukon River,
lampreys were formerly taken in large numbers with dip nets and used for food
(Evermann and Goldsborough, 1907). However, at the present time this species
is but little utilized for any purpose, although a market for smoked lampreys
might well be developed.
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RIVER LAMPREY
Lampetra ayresi (Gunther)
DISTINCTIVE CHARACTERS
The river lamprey has two large teeth on the supraoral bar, a large
middle tooth on the tongue, three points (rarely two) on each central lateral
tooth plate, and no posterial teeth,
DESCRIPTION
Body elongate, eel-like, nearly cylindrical as far back as the dorsal
and anal fins, where it becomes compressed. Depth about 8-111 of total
length. Head moderate, 12.5% (11-141) of TL. Snout rounded, its length
about 601 of head. Eye large, round, its diameter about 1451 (100-200%) of
postorbital. Nostril on median dorsal line, barely in front of eyes, in a
short tube. Gill openings 7, their vertical height about 24% of eye diameter,
spaces between openings equal to about 501 of eye. Trunk tnyotomes usually
65-71, average 68, rarely as low as 60, between last gill opening and anus.
Anus at anterior end of anal fin. A slender anal papilla present in males.
Teeth
Supraoral bar with two sharp points. Three lateral tooth plates on each
side, the central ones with three points, the others with two. Tongue with a
large, triangular central tooth and about seven small ones on each side. In-
fraoral bar with 7-10 (average 9) teeth. No posterial teeth.
Fins
Two dorsal fins, the anterior the lower, the fins separate in non-breeding
individuals but coming in contact at spawning. Caudal fin pointed, lobes
about equal, lower lobe joined to anal. Anal fin virtually absent in males.
Color
Dark brown or brownish grey on sides and back, belly whitish, silvery
around head, gill openings and lower sides. Caudal fin with a band of dark
pigment inside its margin, symmetrical on both lobes.
S i ze
This is a rather small lamprey, maximum length about 311 mm (1 ft).
Average length seems to be about 170 mm (6 1/2 in) (Vladykov and Follett,
1958).
RANGE AND ABUNDANCE
The river lamprey is confined to the west coast of North America, from
as far south as San Francisco Bay (Vladykov and Follett, 1958) north to Tee
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Harbor, 24 km northwest of Juneau, Alaska (Scott and Grossman, 1973). It
does not appear to be particularly abundant anywhere within its range.
HABITS
Like most parasitic lampreys, L_. ay_res_i_ is anadromous. Spawning occurs
in the spring (in late April through May in California). Spawning behavior
has not been described, nor is the length of larval or adult life known. Egg
numbers per female have been estimated at 37,288 for a specimen 175 mm long,
and 11,398 for one of 230 mm. These numbers resulted in estimates of 5,108
and 5,428 eggs per gram of ovary, respectively. The river lamprey is known
to parasitize small coho and kokanee salmon, and is in turn eaten by other
fishes such as ling cod (Vladikov and Follett, 1958).
IMPORTANCE TO MAN
As with other lampreys, the importance of this species is indirect, in
that it parasitizes more desirable fishes. However, its limited abundance
makes it of little concern.
WESTERN BROOK LAMPREY
L ampetra ri chard son1 Vladykov and Follett
DISTINCTIVE CHARACTERS
The very blunt teeth, the lack of a distinct median tooth on the tongue
and the lack of posterial teeth distinguish this species from others found in
Alaska.
DESCRIPTION
Body elongate and eel-like, more or less cylindrical anteriorly but
becoming compressed posteriorly. Depth of body about 7% of total length.
Head short, about 12.5% of total length. Snout rounded anteriorly, its
length about 60% of head. Eye small, round, diameter about 2.3% of total
length, about equal to postorbital distance. A single nostril on dorsal
median line of head, a short distance before eyes, in a short, fairly promi-
nent tube. Oral sucker round, its diameter about 59-60% of snout length.
Gill openings 7 on each side. Trunk myotomes 60-67 between last gill opening
and anus. Anus just before anal fin. A small but obvious genital papilla in
males. The number and arrangement of the sensory pores on the head seem to
be as in L_. ja^onjca.
Teeth
Two broad, blunt teeth on supraoral bar. Two blunt teeth on each lateral
tooth plate. About five blunt teeth on tongue. Infraoral bar with about
seven blunt teeth. No posterial teeth.
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Fins
Two dorsal fins, the anterior lower thin the posterior, arising well back
on body. Fins separate from each other in young, but come together and at
least touch (may fuse) with onset of sexual maturity. Caudal fin joined to
both dorsal and anal fins. Anal fin small, represented in males only by a
low ridge.
Color
Sides and back dark grayish to brownish, ventral side of body whitish.
Caudal fin and posterior tip of body conspicuously dark.
Size
This is quite a small species, the largest specimen recorded being only
154 mm (about 6 in) long. Ammocoetes range up to 175 nm (Vladykov and Follett,
1965).
RANGE AND ABUNDANCE
In Alaska the western brook lamprey has been recorded only from McDonald
Lake, on the Cleveland Peninsula in southeastern Alaska just north of Yes Bay.
However, it ranges southward in coastal streams in British Columbia, Washing-
ton and Oregon as far south as the Umpqua River, near Reedsport, Oregon. It
is not particularly abundant anywhere, as far as is known.
HABITS
Nothing is known of the biology of this species. Presumably, its life
history is like that of other non-parasitic lampreys, in which the larvae
burrow into the bottoms of brooks or hide under stones, hence are not easily
discovered. The adults do not feed and probably do not live more than a few
months. They die after spawning.
IMPORTANCE TO MAN
This species is of no importance to man, except for its scientific and
aesthetic interest. It probably contributes a minor portion of the food of
predacious fishes, insects, etc., wherever it is found.
10
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SECTION 3
STURGEONS—FAMILY ACIPENSERIDAE
This group includes some of the largest fishes in the world, some ex-
ceeding 1,000 kg (a ton) in weight and 6 meters (20 ft) in length. They are
heavy fishes, with more or less cylindrical bodies, a flat, extended snout
and a toothless, protrusible mouth on the under side of the snout, with four
barbels in front of it. The tail is upturned, like that of sharks, with the
upper lobe notably longer than the lower. The body shows five rows of large,
bony plates^ a dorsal row along the back, a lateral row on each side, and two
ventro-lateral rows. The plates are sharp-pointed in the young, but become
blunt in adults. The skin between the plates appears naked but actually bears
patches of fine denticles. The head is covered by bony plates. The first ray
of the pectoral fin is heavy and ossified. The two species of sturgeon found
in Alaska are anadrorrous, spending most of their lives in the sea but breeding
in fresh water. Other species are confined to fresh water.
n
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GREEN STURGEON
Acipenser mediostris Ayres
DISTINCTIVE CHARACTERS
A single row of 1 to 4 bony plates along the mid-ventral line between
the anus and the anal fin, and about 33-35 dorsal rays characterize the green
sturgeon.
DESCRIPTION
Body rather robust, more or less five-sided in cross-section, depth
about 10% of total length. Head flattened, its length about 25% of total
length. Snout produced, flattened, somewhate concave in profile. Mouth
snail, transverse, protrusible, on ventral side, toothless. Four sensory
barbels in front of mouth, their bases nearer to mouth than to tip of snout.
Eyes small, round. Gill rakers 18-20. Body with five longitudinal rows of
bony, keeled plates, the keels sharp-pointed in young but becoming more blunt
and rounded in older individuals. Plates in one dorsal row of 8-11 plates,
two lateral rows of 23-30 plates, and two ventro-lateral rows of 7-10 plates
each. Skin between plates with numerous small, rough plates or denticles.
A single row of 1-4 small bony plates on ventral mid-line between'anus and
origin of anal fin.
Fins
Dorsal arising at posterior third of total length, usually with 33-35
rays. Anal originating under posterior part of dorsal, 22-28 rays. Pectoral
origins low on body, just behind gill openings, fins large and rounded. Pel-
vic fins arising near anus. Caudal heterocercal, lower lobe about 3/4 as
long as upper.
Color
Generally olive to dark green, lower parts more or less whitish green.
A longitudinal olive green stripe on side between lateral and ventro-lateral
plates, another on mid-ventral surface. Fins grayish to pale green.
Size
Reaches a length of over 2 meters (7 ft) and a weight of about 160 kg
(350 Ib) (Dees, 1961).
12
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RANGE AND ABUNDANCE
In Alaska, the green sturgeon has been found as far north as the north-
west side of Unalaska Island in the Aleutians, but it is rather uncommon in
the state (Wilimovsky, 1964). The range extends southward along the Pacific
coast of North America to southern California, with two specimens recorded
from just south of Los Angeles (Roedel, 1941; Norris, 1957). It has also
been reported from Ensenada, Baha California (Miller and Lea, 1972). On the
western side of the Pacific, this species ranges from the Amur River south to
Hokkaido, Peter the Great Bay and Kunsan, Korea. The area of greatest abun-
dance seems to be from the Columbia River in Washington and Oregon to the
Fraser River in British Columbia.
HABITS •
Very little is known of the biology of the green sturgeon. The little
information available suggests that its life history is like that of other
anadromous sturgeons. However, it is not known to ascend streams to any
great distance. It is most commonly found in estuaries, in the lower reaches
of large rivers and in salt or brackish water off river mouths.
The green sturgeon probably spawns in fresh water. It is captured in
nets and by anglers in the lower Columbia and Fraser rivers in late summer
to winter. Presumably, these fish move into the fresh water in the late
summer and fall to spawn the following spring.
Green sturgeon may undertake considerable movements in the ocean. Three
fish tagged in 1954 in San Pablo Bay, California, near the mouth of the Sacra-
mento River, were recovered one to four years later in Oregon. One was re-
captured in Winchester Bay, the other two in the lower Columbia River (Chad-
wick, 1959).
Nothing is known of growth rates, age, etc.
Food of the green sturgeon probably consists largely of bottom-dwelling
invertebrates and small fishes. There is one report of a green sturgeon
having fed on sand lance (Anon., 1954).
IMPORTANCE TO MAN
The green sturgeon is virtually unutilized in North America, although it
is taken commercially in the Bering Sea by Russian fishermen (Magnin, 1959).
As a food fish, the green sturgeon is considered inferior. The flesh is said
to be dark, with a strong, disagreeable taste and an unpleasant odor. It was
once thought to be poisonous. At least in the past, the roe was not utilized
(Jordan and Evermann, 1908). The green sturgeon does not seem to be particu-
larly common, although occasional concentrations (feeding aggregations?) may
be encountered. One such was reported off Kyuquot Sound on the northwest
coast of Vancouver Island, where 75 fish were taken in a single day (Anon.,
1954).
13
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WHITE STURGEON
Acipenser transition tanus Richardson
DISTINCTIVE CHARACTERS
Two rows, each of 4-8 bony plates along mid-ventral line between anus
and anal fin, and about 45 rays in the dorsal fin serve to distinguish the
white sturgeon.
DESCRIPTION
Body elongate, sub-cylindrical, depth about 10$ of total length. Head
more or less conical, somewhat flattened above, its length 19-231 of total
length. Snout moderately produced, blunt, relatively shorter in adults than
in young. Eyes small. Nostrils prominent. Mouth wide, transverse., pro-
tractile and toothless, located on ventral side of head below eyes. Four
tactile barbels on ventral side of snout in front of mouth, a little closer
to tip of snout than to mouth. Gill rakers 34-36. Body with five longitudi-
nal rows of keeled, bony plates. Dorsal row 11-14, lateral rows 38-48, ventro-
lateral rows 9-12. Rarely there may be a secondary row on each side between
the dorsal and lateral rows (Bajkov, 1955). Skin between plates with numer-
ous, rather inconspicuous, stellate tubercles. Two rows of 4-8 plates each
on ventral side between anus and anal fin.
Fins
Dorsal fin originating near posterior third of total length, 44-48 rays.
Anal origin under posterior part of dorsal, the fin with 28-31 rays. Pectoral
fins arising low on body, just behind gill openings, upper rays longer than
lower, first ray notably enlarged and bony. Pelvic origins slightly before
anus. Caudal heterocercal, upper lobe about twice as long as lower.
Col or
Generally gray or brownish above, paler below. Fins gray.
Size
The largest freshwater fish in North America, the white sturgeon is re-
puted to attain a length of 6 meters (20 feet) and weight to over 860 kg
(1,900 Ib). However, the heaviest documented specimen weighed 630 kg
(1,387 Ib) (Hart, 1973).
RANGE AND ABUNDANCE
Pacific coast of North America, from Ensenada, Baha California in the
south (Miller and Lea, 1972) north to the Gulf of Alaska. Found in rivers
and estuaries, and in the sea to as deep as 122 meters (400 ft). Most abun-
dant in and around the mouth of the Columbia River. It has been suggested
that the Lake Iliamna "sea monster" may be a large white sturgeon. Not com-
mon in Alaska.
14
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HABITS
The white sturgeon is anadromous, although some individuals apparently
spend much of their lives in fresh water. Mature fish enter the estuaries
and lower reaches of large rivers any time from fall to early spring, although
the majority seem to leave the sea in the spring. Upstream migration begins
in April or May, with spawning taking place in May and June in most areas
(Bajkov, 1951).
Some fish may move great distances, as far up the Columbia River as
Flathead Lake, Montana (Brunson and Block, 1957). Such fish may well start
their upstream migration earlier (perhaps even the previous year) and spawn
later than fish moving shorter distances.
Spawning areas are said to be in swift water over rocky bottoms. Water
temperature during the spawning period has been reported as S.O-16.7°C
(Scott and Grossman, 1973).
First spawning occurs at about 10 years of age for males, 11 to 12 years
for females of southern stocks (Carlander, 1969; Pycha, 1956). In the Fraser
River, males mature at 11 to 22 years, females at 26 to 34 (Semakula and
Larkin, 1968). Adults return to the sea after spawning, but do not breed
annually. Intervals of 2-9 years may occur between breeding periods in the
Fraser River stock (Semakula and Larkin, 1968).
Fecundity varies with the size of the individual, from a few hundred
thousand eggs in small females to about 2,000,000 or more in the largest
adults. Ovary weights may reach more than 110 kg.
The eggs are brown in color. They are sticky and adhere to the stream
bottom. Time required for hatching is unknown. Growth of the young is quite
rapid for the first few years. One-year-old fish from San Pablo Bay, Califor-
nia, averaged about 260 mm total length, five-year-olds about 800 mm. After
the 8th year, the fish grew at a fairly constant rate of 50-60 mm per year.
Similar growth rates were found in white sturgeon of the Fraser River (Pycha,
1956; Semakula and Larkin, 1968).
Adult white sturgeon may reach great size, but very large fish are rare
nowadays. The average weight of net-caught fish seems to be on the order of
16-25 kg. However, around the turn of the century, very large fish appear to
have been more common. One of 862 kg (1,900 Ib) was supposed to have been
taken at Astoria, Oregon. Another (possibly the same fish) of about 908 kg
(2,000 Ib) was supposed to have been exhibited at the Chicago World's Fair
in 1893, but neither the specimen nor records of it now exist (Jordan et al,
1930; Gudger, 1942). Other large white sturgeon include one of 817 kg (1800
Ib) from the Fraser River at Mission, B.C., sometime before 1897, one of 630
kg (1,387 Ib) from the Fraser at New Westminster in 1897, and one of 583 kg
(1,285 Ib) and 3.81 m long (12 ft, 6 in) from the Columbia River at Vancouver,
Washington in 1912 (Clemens and Wilby, 1946, 1961; Gudger, 1942).
15
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The maximum age recorded for the white sturgeon is 82 years (Carlander,
1969). Extrapolation of data given by Pycha (1956) suggests that the very
large fish mentioned above must have been on the order of 100 years old,
possibly older.
The white sturgeon spends much of its life in the sea, where it is
thought to remain fairly close to shore. It is known from depths of 30 m or
less and has been taken in salt, brackish and fresh waters with temperatures
between 0° and 23.3°C. Migrations and movements in the sea are not well
known, but they certainly exist, for a specimen tagged in San Pablo Bay,
California, was recaptured about 10 months later at the mouth of the Columbia
River, a minimum distance traveled of 1,056 km (Chadwick, 1959).
The white sturgeon is primarily an omnivorous scavenger, but seems also
to be actively piscivorous, much more so than other sturgeons. Eulachon and
lampreys appear to be favored foods, along with sculpins, sticklebacks, sal-
monids, crayfish, frogs, molluscs, insect larvae and various other organisms
(Gudger, 1942; Merrell, 1961; Radtke, 1966; Carlander, 1969; Scott and
Grossman, 1973). The young feed primarily on plankton, amphipods, shrimp and
tendepedid larvae. Other food organisms include mysids, Daphnia, Chaoborus
larvae, copepods and various immature insects (Carlander, 1969; Schreiber,
1961; Scott and Grossman, 1973).
IMPORTANCE TO MAN
Before about 1880, white sturgeon were utilized almost entirely in sub-
sistence fisheries. As the Pacific salmon fisheries developed, the sturgeon
came to be considered a nuisance. However, about 1886 or 1887, "an eastern
firm" began to ship frozen sturgeon to eastern markets. The enterprise was
successful, and by 1892 the Oregon landings from the Columbia River were in
excess of 1,498,000 kg (3,300,000 IDS). The greater part of the flesh was
smoked after arrival in the east, the air bladder made into isinglass, the
eggs into caviar. Indeed, every part of the fish was used, even the backbone
(McGuire, 1896). This state of affairs did not last long. By 1897 the total
catch for Washington and Oregon had dropped to just under 629,142 kg (1,386,000
Ib), and by 1900 had declined further to a mere 55,152 kg (121,500 lb). The
fishery continued at a low ebb until the mid-1940's, when catches began to
rise. Current landings (1970) are on the order of 227,000 kg (500,000-Ib).
The catch from the Fraser River has averaged about 13,600 kg (30,000 Ib) an-
nually since 1941 (Hart, 1973).
Flesh of the white sturgeon is highly esteemed, either fresh or smoked,
and the eggs make excellent caviar. However, there is no demand for isinglass
nowadays.
16
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SECTION 4
HERRINGS—FAMILY CLUPEIDAE
This is a large family, containing about 70 genera and more than 150
species, but only two members, the Pacific herring, Clupea harengus pallasi,
and the American shad, Alosa sapidissima, are found in Alaska,
Characteristically, the members of this group have slender, compressed
bodies, although some are short and deep and a few are sub-cylindrical. The
scales are cycloid and in most are extremely deciduous. The ventral side of
the body is compressed to a sharp edge, usually serrate. There is no lateral
line. The mouth is large, teeth small or absent in almost all. The dorsal
fin is located about at the mid-length of the body and there is no adipose
fin. The caudal fin is forked.
This is one of the most important of the families of fishes from the
human economic standpoint. Vast numbers are taken for food, fresh, canned,
smoked, pickled, etc. Other species are utilized for fish meal and oil. The
smaller species and the young of the larger ones serve as food for a tremen-
dous variety of large fishes, aquatic mammals} birds and turtles.
17
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PACIFIC HERRING
Clupea harengus pal 1 as1 Valenciennes
DISTINCTIVE CHARACTERS
The Pacific herring may be distinguished from the American shad by the
origin of the dorsal fin about mid-way between the tip of the snout and the
base of the tail (further forward in the shad); the body depth 25% or less
of the length (more in the shad); and the presence of teeth on the vomer bone
(absent in the shad).
DESCRIPTION
Body elongate, compressed, its depth less than 25% of total length. Head
compressed, its length about equal to body depth. Snout moderate, about equal
to eye diameter. Eyes round, adipose lids present. Mouth fairly large, no
teeth on jaws but a patch of fine teeth on vomer in roof of mouth. Gill
rakers long, those at angle of gill arch about as long as eye diameter, fine,
63-73, the number apparently increasing with size of fish. Lateral line ab-
sent.
Fins
D. 15-21, origin of fin about equidistant between tip of snout and base
of tail. A. 14-20. P-|. 17-19. ?2- 8~9.' abdominal in position, each with
a fleshy axillary process above its origin. Caudal fin forked.
Scales
Large, cycloid, deciduous, 38-54 rows along side of body. On mid-ventral
line, scales modified, with moderate keels on those before pelvic fins,
strong keels between pel vies and anal.
Color
Dark blue to olivaceous above, shading to silver below.
Size
Said to reach 38 cm (15 in) in Alaskan waters, but the average is
probably around 25 cm (10 in).
18
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RANGE AND ABUNDANCE
On the North American coast, the Pacific herring is found from northern
Baha California (Miner and Lea, 1972) to Bathurst Inlet in the eastern
Beaufort Sea (Richardson, 1836). Commercial fisheries exist as far south
as San Francisco, but the region of greatest abundance is along the coasts
of British Columbia and southeastern and central Alaska.
t
On the western side of the Pacific, this herring ranges from the Japanese
islands and Korea north and west to the Laptev Sea on the arctic coast of
Siberia. Very closely related forms are found still further west in the
Kara Sea, Barents Sea and White Sea (Andriyashev, 1954; Nikolskii, 1961).
Although generally considered a marine species, the Pacific herring
occasionally enters estuaries. Freshwater, landlocked races have been re-
ported from Japan (Fujita and Kokubo, 1927). On this basis, the Pacific
herring is included here,
HABITS
The Pacific herring spawn from December to July. The winter spawning
occurs at the southern end of its range, around San Diego, California, and
occurs progressively later in the year in more northern populations
(Schaefer, 1937; Scattergood et al, 1959; Barraclough, 1967). In Alaska,
most spawning activity goes on between mid-March and mid-April in south-
eastern; April-May in Prince William Sound; and May-June around Kodiak
Island (Scattergood et al, 1959). Spawning has been recorded in the Bering
Sea near St. Michael's in June (Nelson, 1887). Spawning activity is roughly
related to temperature and occurs at water temperatures between 3.0 and 12.3°C.
It is uncertain whether the herring of the Chukchi and Beaufort seas are
self-sustaining populations or represent migrants from farther south.
The eggs are 1.2-1.75 mm in diameter after being spawned. Fecundity is
related to size and age, and northern fish appear to produce more eggs than
do southern ones of the same size (Nagasaki, 1958), although there is con-
siderable variation among populations (Katz, 1948). Average egg numbers along
the coast of North America range from a little over 8,000 in two-year-old
fish from southern British Columbia to more than 59,000 in seven-year-olds
(Nagasaki, 1958). A fish from Peter the Great Bay, near Vladivostok, was re-
ported to contain more than 134,000 eggs (Katz, 1948, citing Russian data).
Spawning of the Pacific herring appears to be a mass proposition, with
no definite pairing of males and females, although Rounsefell (1930) described
the male following "a few inches behind the famale covering the attached eggs
with a stream of milt."
Spawning takes place in shallow water, from the very edge of the high
tide line to a depth of about 7 meters (23 ft), most commonly over vegetation
such as eel grass or Fucus, or on brush, pilings, rocks, etc. In the spawning
act, the female turns on her side, fins extended and body nearly rigid. Vi-
brating her tail, she moves slowly forward, her vent brushing the eel grass
19
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or other substrate, depositing her eggs as she goes. The posterior edge of
the vent is equipped with a fleshy lobe which blocks extrusion of the eggs.
Brushing the vent against the substrate pulls this lobe out of the way,
allovnng the eggs to escape. The female remains on her side for 2-5 seconds,
then resumes upright posture. This procedure is repeated again and again
until all the eggs have been laid, which may take several days (Schaefer,
1937). While the females are depositing their eggs, the males in the area
are releasing* milt. The milt mixes and spreads in the water, fertilizing
the eggs, and the whitish, milt-filled water may extend for several miles
along the shore.
The eggs are sticky and adhere to whatever they touch. They hatch in
about 10 days at the usual temperatures, yielding yolk-sac larvae about
7.5 mm (0.3 in) long. Feeding begins two weeks or less after hatching, as
soon as the yolk has been absorbed. The first foods are mainly copepods,
invertebrate eggs and diatoms. The young fish are in turn eaten by various
filter-feeders such as pilchards, and entanglers like jelly fishes, as well
as by young salmons.
During this early growth period, the young herring may be dispersed by
waves and currents. Those which are carried out to sea apparently perish and
this off-shore transport seems to be the major cause of mortality at this
time (Stevenson, 1962). Those not carried out to sea subsequently congregate
in suitable shallow bays, inlets and channels (Hourston, 1958, 1959). By
the end of the summer, the young may be as long as 10 cm (4 in). They move
into deeper water in the fall and virtually disappear from the fishing
grounds for the next couple of years.
After the first year or two, northern herring grow faster than do those
from California, They also live longer and get bigger. The oldest Califor-
nia herring was a 9+, but a 19+ has been recorded from Alaska. Most herring
in the Alaskan catch are between 4 and 9 years old (Rounsefell, 1930;
Miller and Schmidtke, 1956).
As the fish grow, larger food items become more important in the diet.
Copepods remain important to the young, but the diet also includes euphau-
siids, amphipods, cladocerans, decapod and mollusc larvae, etc. Adults ex-
hibit a similar diet, but also eat larger crustaceans and a wide variety of
small fishes. In turn, adult herring are preyed upon by virtually every
animal large enough to eat them - dogfish and other sharks, salmon, cods,
mackerel, squid, seals and sea lions and man.
Pacific herring usually move offshore in the winter and back onshore for
spawning, but apparently do not undertake extensive coastwise migrations.
Mixing of local populations is relatively rare, so particular groups can be
recognized by certain physical characteristics all the way from Alaska to
southern California.
IMPORTANCE TO MAN
The Pacific herring has never rivaled its Atlantic relative in terms of
tonnage landed, but it is nevertheless a significant part of the fisheries
20
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of the North Pacific. Along the Siberian coast, "the annual catch exceeds
6,000,000 centners" (300,000,000 kg) (Andriyashev, 1954). High production
values for the North American coast are generally slightly lower. Almost all
the North American herring catch comes from Alaska and British Columbia.
The Alaskan herring fishery began in the late 19th century. Statistics
for 1882 show landings of 1,363,636 kg (3,000,000 Ib). Over the next 34
years, the catch fluctuated between 2.5 and 14.6 million kg (5.6-32.1 million
Ib), averaging about 6.6 million kg (14.5 million Ib). The majority of the
catch went into salted herring in various forms. World War I increased the
demand for Alaska salt herring and catches increased accordingly. By the
mid-1920's, however, the Alaskan salteries could no longer compete, either in
price or in quality of product, with the resurgent European industry. Never-
theless, a great expansion of the reduction industry led to further increased
landings. From 1924 to 1941, Alaskan landings were more than 45.5 million kg
(100 million Ib) per year, peaking at more than 118.6 million kg (261 million
Ib) in 1937. For the next 10 years or so, catches again varied widely, then,
beginning in the early 1950's, dropped rapidly to their present low level.
This has been ascribed to a sudden paucity of fish, increased competition
from foreign sources and other factors such as greater profit in other fish-
eries, which attracted operators away from herring. Probably all three were
important. In recent years, the Alaskan herring fishery has produced be-
tween 4.5 and 9.1 million kg (10-20 million Ib) annually, rrost of it being
used for bait (Anderson and Power, 1946, 1957; Anderson and Peterson, 1953;
Power, 1962; Lyles, 1969; Anonymous, 1971, 1972, 1973; Browning, 1974).
AMERICAN SHAD
A]psa sapidissima Wilson
DISTINCTIVE CHARACTERS
The American shad is easily distinguished from similar-appearing Alaskan
fishes by the rather"pointed lower jaw which fits into a deep notch in the
upper jaw; the coarse, curved, radiating striae on the gill cover; and the
row (sometimes two or even three rows) of dark spots on the upper sides be-
hind the head.
DESCRIPTION
Body compressed, fairly deep, depth 30-37% of standard length. Head
moderate, compressed, its length 23-28% of SL. Snout moderate, 27-32% of
head length. Eye about equal to snout, adipose lids well developed. Mouth
terminal, lower jaw fitting into a notch in upper jaw in adults, but this
notch lacking in young less than about 150 mm long. Upper jaw (maxilla)
reaches middle of eye in young, to below posterior margin of eye in adults.
Teeth absent, or at best few and small in adults, present but minute on jaws
and mid-line of tongue in young specimens. Gill rakers long and slender,
those at angle of gill arch shorter than snout in young but longer than snout
in adults, number increasing with age and size, 26-43 on lower limb in young,
59-73 in adults. Lateral line absent.
' 21
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Fins
Dorsal elevated anteriorly, margin slightly concave, its origin slightly
forward of pelvic bases, rays 15-19, usually 17 or 18. No adipose dorsal
Anal long and low, 20-23 rays. Pectorals small, low on sides, 14-18 rays.
Pel vies with 9 rays. A pelvic axillary process present. Caudal deeply forked,
upper and lower lobes about equal.
Scales
Moderately adherent, cycloid, with crenulate borders, 52-62 rows along
middle of side, 15-16 between pelvic base and dorsal origin. Ventral scutes
well developed, 20-22 before pelvics, 14-17 behind.
Cojor
Metallic greenish or bluish above shading to silvery on sides and belly.
A row of dark spots on upper part of sides behind head, sometimes a second,
rarely a third row below the first. Fins pale to greenish, dorsal and caudal
dusky in large specimens, tips of caudal sometimes dark.
Size
The largest North American member of its family, the shad reaches a
length of 760 mm (30 in) and a weight of 6.4 kg (14 Ib). However, the aver-
age weight of adult shad is probably around 2-2.5 kg (4.5-5.5 Ib). Females
are usually larger and heavier than males.
RANGE AND ABUNDANCE
The normal range of the shad on the east coast is from the St. Lawrence
River and Nova Scotia south to the Indian River, Florida. It has been report-
ed from as far north as Bull's Bay, Newfoundland. On the Pacific side, the
shad ranges from Todos Santos Bay, Baha California, to southeastern Alaska,
with strays north and west to Cook Inlet, Kodiak Island and Kamschatka.
In the Major portions of its range, the shad is quite abundant, but it
is quite scarce in Alaska. Fishermen in southeastern pick up a few in the
salmon nets every year, but, as far as is known, there are no breeding popu-
lations of shad in the state.
HABITS
The shad is anadromous, spending most of its life in the sea but return-
ing to freshwater streams to breed. Some spawn almost immediately on enter-
ing fresh water, while others may undertake fairly long journeys, as much as
630 km (394 mi) upstream to their favored spawning grounds (Hildebrand, 1963).
Timing of the run is governed by water temperature. The shad enter the rivers
when the temperature is between 10 and 13°C. Hence, in the southern part of
the range, spawning may occur as early as March, but not until late June at
the northernmost end.
22
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Spawning usually takes place in the evening, between dusk and 10 p.m.,
on sandy or pebbly shallows. No nest or redd is made. The fish pair and
swin1 close together, side by side, releasing eggs and milt. A single female
may require several evenings to complete spawning, as all eggs apparently are
not shed at once. The eggs are slightly denser than water, non-adhesive, and
drop loosely and singly to the bottom, where they lodge in crevices between
the pebbles. Fecundity estimates range from 116,000 to 616,000 eggs per fe-
male, with egg number directly related to the size and age of the individual
fish. Early estimates of average egg counts of about 30,000 and maximum
156,000 apparently referred to hatchery egg-takes in the late 19th century.
On the Atlantic coast, fish in the southern part of the range produce more
eggs than do northern fish, but it is not known if this also holds true along
the Pacific coast (Lehman, 1953; Davis, 1957).
The spent fish return to the sea immediately after spawning. They begin
to feed while still in fresh water and may recover a good deal of fat before
reaching the sea. Adult shad return to fresh water year after year to spawn
in their natal streams (Hollis, 1948; Talbot and Sykes, 1958; Cheek, 1968).
The fertile eggs, about 3.5 mm {0.14 in) in diameter and pale pink to
amber in color, hatch in 12 to 15 days at 12°C, 11 days at 13.3°C, 6 to 8 days
at 14 to 17°C, and only 3 days at 23.3°C (Rice, 1884; Ryder, 1884b; Leim, 1924).
The young are about 9-10 mm long at hatching. The yolk sac is absorbed in
4-5 days and the young begin to feed 10-12 days after hatching. The first
food consists primarily of copepods and chironomid larvae (Ryder, 1884a; Leim,
1924), but as the young fish grow they take larger and larger particles. They
have doubled their length, to about 20 mm, in three to four weeks, by which
time the fins are fully developed and metamorphosis is virtually complete. A
young shad about 50 mm (2 in) long closely resembles the adult, except that
the body is more slender, the shoulder spots are not developed and the ventral
scutes are more prominent. Adult fish feed on copepods, mysids, shrimps, bar-
nacle larvae, ostracods, amphipods, insects and, rarely, small fishes (Bigelow
and Schroeder, 1953; Hildebrand, 1963).
Growth proceeds rapidly during the summer in fresh water. By the time
the young fish move downstream in the fall, they are between 37 and 112 mm
(1.5-4.5 in) long. Growth rate of the young in salt water is unknown. How-
ever, on the basis of back-calculations made in scale studies, the lengths at
various ages are approximately as follows: 1+, 12.7-19.3 cm; 2+, 22.9-29.5 cm;
3+, 25.4-37.6 cm; 4+, 37.6-42.9 cm; 5+, 40.4-48.3 cm; 6+, 51.3 cm (LaPointe,
1958; Cheek, 1968). Growth rates vary from one population to another. Shad
attain at least 11 years, but age determinations on old fish are difficult be-
cause of the extensive regression of the edges of the scales at spawning
(Gating, 1953).
After leaving the natal streams, the young shad spend 3-4 years in the
sea. Sexual maturity is usually attained in the fourth year for males, the
fifth for females, and spawning occurs annually thereafter. An exception
seems to lie in fish of the southern Atlantic states, from North Carolina
south, which spawn but once and then die (Cheek, 1968).
23
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Aside from the spawning migrations, little is known of the movements of
the shad. Atlantic fish, from Chesapeake Bay northward, congregate in the
Gulf of Maine after spawning and presumably winter in deep water off the mid-
Atlantic coast (Talbot and Sykes, 1958). What happens with Pacific popula-
tions is unknown, but, at least in the early years following the first trans-
plants, the Pacific shad must have wandered considerably.
The American shad was first introduced into Pacific waters in 1871, when
about 10,000 eggs were placed in the Sacramento River, California, at Tehama,
on 26 June. Another 35,000 eggs were planted on 2 July 1873, and between
1876 and 1880 almost 600,000 more were added to the Sacramento. Plants were
made 'in the Columbia River basin (910,000) in 1885 and 1886. Additional
plants, totalling more than 9.6 million eggs, were made in the Colorado River
and in Utah and Idaho between 1873 and 1891, but these seem to have been un-
successful (Smith, 1896). Shad appeared at Vancouver Island, B.C., in 1876
(McDonald, 1891; Clemens and Wilby, 1946), in the Stikine River near Wrangell,
Alaska in 1891 (Smith, 1896), and by 1904 had reached Cook Inlet, Alaska
(Evermann and Goldsborough, 1907). In 1926 and again in 1937, single indivi-
duals were taken at Kodiak Island (Welander, 1940). Expansion southward also
took place, with shad found near San Diego in 1916 (Starks, 1918) and in Todos
Santos Bay, Baha California, Mexico, in 1958 (Claussen, 1959). Occasional
specimens are taken in Kamschatkan waters (Nikolskii, 1961), Thus, although
the shad normally returns to the natal stream, it has been something of a
wanderer in the new environment of the Pacific.
Despite the rapid expansion of range, however, shad have never been more
than strays in Alaska. The statement by Evermann and Goldsborough (1907) that
"The cannery at Fairhaven took one about July 1, 1903, and the fishermen at
Birch Point got about 3,000 in one day" has at times been taken to indicate
that shad were once abundant in Alaska. However, there is no Birch Point in
the state, and Fairhaven, near Juneau, was not officially named until 1962
(Orth, 1967). The Fairhaven and Birch Point referred to by Evermann and
Goldsborough are undoubtedly at Bellingham, Washington.
IMPORTANCE TO MAN
The shad is an important sport and commercial fish on the east coast,
with total commercial landings in excess of 4.5 million kg (9.9 million Ib)
in most years. Pacific coast landings are about a third of this figure as a
rule. As a sport fish, the shad is highly regarded in the east, for it
readily takes a fly or small spinner and is a game fighter. Because of its
tender mouth, light tackle is imperative. Oddly enough, there is relatively
little sport fishing for shad on the west coast. Because of its scarcity in
the state, the shad is of no importance in the Alaskan fisheries, either
sport or commercial.
24
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SECTION 5
SALMONS, TROUTS, CHARRS, WHITEFISHES AND GRAYLING -- FAMILY SALHONIDAE
The salmonoids are freshwater or anadromous fishes, mostly medium to
large sized, with more or less round to moderately compressed bodies. The
fins are all soft-rayed and a dorsal adipose fin is present. Scales are
cycloid, with a distinct, elongate axillary scale or process present at the
base of each pelvic fin. The mouth may be large and well-toothed to small
and toothless. The swim-bladder is connected to the esophagus. Pyloric
caeca are numerous. Sexual dimorphism is often apparent at spawning time.
The group is conveniently divided into three subfamilies: Coregoninae,
the whitefishes and ciscoes; Salmoninae, the salmons, trouts and charrs; and
Thymallinae, the graylings.
The Salmonidae is the dominant family of fishes in the northern parts
of Europe, Asia and North America, with a number of species reaching the
high Arctic. Various members of the group, especially the Pacific salmons,
genus Oncorhynchus, are of great commercial importance, while others, as the
troutsTSalmo), charrs (Salvelinus), and grayling (Thymallus} are noted
chiefly as sport fishes.
Characteristically, the Salmoninae have small scales, 115-200 along the
lateral line; well developed teeth on jaws and vomer; a truncate caudal fin
(forked in the lake trout); and prominent parr marks on the young (absent in
pink salmon).
The Coregoninae possess relatively large scales, not more than 100 along
the lateral line; teeth are small and weak, or absent; and parr marks are
absent except in the genus Prosopium.
The Thymallinae, with only the arctic grayling, Thyma11us arcticus,
present in Alaska, agree generally with the Coregoninae, but parr marks are
present in the young and the dorsal fin of adults is greatly enlarged,
especially in mature males.
25
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INCONNU OR SHEEPISH
Stenodus leucichthys (Gilldenstadt)
DISTINCTIVE CHARACTERS
The inconnu is easily distinguished from other coregonines by the large
mouth, the protruding lower jaw, and only 13-17 gill rakers on the lower limb
of the first gill arch.
DESCRIPTION
Body elongate, little compressed. Depth 20-23% of total length, deepest
just behind pectoral fins. Head lonq, 24-28% of total length, depth of head
about half its length. Snout prominent, about 23% of head length. Eye round,
its diameter averaging about 15% of head length. Nostrils with a double flap
between openings on each side. Mouth large, lower jaw protruding, maxillary
reaching at least to vertical through middle of eye. Teeth very small in
velvet-like bands on anterior portion of both jaws and on tongue, vomer and
palatines. Gill rakers 17-24. Branchiostegals 9-12. Lateral line scales
90-115. Pyloric caeca 144-211. Vertebrae 63-69.
Fins
D. 11-19, the fin high and pointed.
axillary process well developed. Caudal
A. 14-19.
fin forked.
14-17. P2. 11-12,
Scales
Large, cycloid.
Color
Overall color silvery, with the back usually greenish, bluish or pale
brown. Silvery white below. Dorsal and caudal fins with dusky margins,
other fins pale.
Size
Up to 27 kg in the Kobuk River, Alaska (Alt, 1969}, to 28.6 kg in the
Mackenzie River (Dymond, 1943} and to 40 kg in Siberia (Wynne-Edwards, 1952}
26
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RANGE AND ABUNDANCE
In Alaska, the inconnu Is known from the Kuskokwim, Yukon, Selawik and
Kobuk drainages. It is found in^the Yukon and its tributaries all the way
from the mouth to Teslin Lake, but apparently does not ascend the Tanana much
beyond Fairbanks. It migrates up the Koyokuk River at .least as far as Alatna.
Except for a few reported from the Meade River, it is not found north and
east of the Kobuk River. In Canada, the inconnu is present in most of the
MacKenzie drainage and east to the Anderson River.
The inconnu ranges westward across Siberia to the White Sea, south to
Kamschatka. Another subspecies is present in Caspian Sea drainages.
Because of its migratory habits, the abundance of the inconnu is both
local and seasonal. However, within these limits it is abundant in most
areas where it is found.
HABITS
Inconnu spawn in the fall, late September and early October, in Alaska
at water temperatures of 1.4-4.6°C. The spawning grounds are located in clear,
fairly swift streams, over bottom composed of differentially-sized gravel and
sand under 1-3 m of water. Spawning takes place in the evening, usually be-
ginning about dusk and continuing well into the night. In the spawning acts
a female, accompanied by a male (rarely by two or more males), rises to the
surface near the upstream end of the spawning grounds. She moves rapidly
across the current, usually in an upright position but sometimes tilted almost
horizontal with the abdomen upstream, extruding eggs as she goes. This activ-
ity lasts 1-3 seconds. The male, meanwhile, stays below the female. The eggs,
averaging about 2.5 mm in diameter, sink to the bottom through the cloud of
sperm released by the male and are fertilized as they sink. Apparently, dif-
ferentially-sized gravel on the stream bottom is important in insuring that
the eggs lodge in the bottom and are not carried away by the current.'
After completing a spawning pass, the female drifts downstream. She may
repeat the spawning act over the downstream portion of the spawning area, or
may move upstream to the head of the grounds before releasing more eggs. It
is not known how many spawning passes are required to extrude all the eggs,
but Alt (1969) noted one female which released eggs six times in a single pass.
Since large females may contain as many as 400,000 eggs, several passes are
undoubtedly needed to complete a spawning.
Development of the eggs is slow. The young hatch in late February to
April and are about 7 mm long. The yolk sac is absorbed in a week to 10 days
and the young inconnu then feed actively on plankton. Growth rates of the
early young are unknown. Eggs are easily handled under hatchery conditions,
but mortality of the young after absorption of the yolk sac has so far been
virtually 100% (LaPerriere, 1973; K. Alt, Alaska Dept. Fish Game, pers. comm.)
On the other hand, Russian workers apparently have had good success with
hatchery rearing of inconnu (Karzinkin, 1951; Nikolskii, 1961).
27
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The distribution of young fish appears to differ from one drainaqe to
another. In the Kobuk and Selawik rivers, and indeed in most areas, the}' are
probably swept downstream by spring floods. Alt (1969) found no fish younger
than 4+ in the Kobuk and Selawik rivers. On the other hand, he found young
of the year, as well as 1+ and 2+ fish, in the upper Yukon, and Fuller (1955)
believed the young remained for two years in rivers emptying into Great Slave
Lake.
Growth of the young is quite rapid for the first two years of life. One-
year-olds in Alaska are 110-150 mm long, with two-year-olds averaging about
200 mm. Growth subsequently slows down. At five years, Alaskan inconnu
average about 430 mm in total length; at 10 years, about 700 mm; and at 15
years, about 960 mm. The largest fish reported by Alt (1969) was a 19+ of
1,190 mm from the Kobuk River, His oldest specimen, a 20+ from the Selawik
varea, was 1,120 mm long.
The inconnu is known to hydridize with other coregonids, doubtless due
more to the broadcasting of eggs and sperm than to interspecific or inter-
generic pairing. Known hybrids include those with Coregonus ne1spni_ (Alt,
1971c), £. autumnalis (Kuznetsov, 1932; Dymond, 1943) and C. muksun
(Kuznets ov, 1932).
As with most fishes, the food of the inconnu changes with the age and
size of the individual, as well as with the time of year. The very younq
post-larvae feed almost exclusively on plankton of various sorts, but soon
graduate to insect larvae and larger zooplankton. In the upoer Yukon River
and in the Ob River of Siberia, inconnu begin to feed on fish in their first
year (Vork, 1948-, Alt, 1965), but in Great Slave Lake they apparently do not
do so until about four years old (Fuller, 1955).
Adults feed mostly on fish, especially the least cisco, Coregonus sardi-
nella. Also important are the isopod, Mesidotea entomon, and the mysid, Hysis
reli'cta. Other foods include king salmon finger!ings, several species of
coregonids, lampreys, charr, smelt, blackfish, suckers, burbot, chubs, stickle-
backs, sculpins and the nymphs and larvae of several orders of insects. The
last are eaten almost exclusively by the smaller fish (Fuller, 1955; Alt, 1965;
1969). The food habits vary with locality and time of year. In Selawik Lake,
C_. sardinella was the major food item of large inconnu in early spring, while
smaller fish fed heavily on small coregonids and invertebrates. In the latter
part of June, Mesi do tea and small coregonids became more important to inconnu
of all sizes, as did the rainbow smelt, Osnerus morel ax (Alt, 1969).
The inconnu of Alaska constitute five rather distinct populations. The
Minto Flats and the upper Yukon River groups are year-round residents,
wintering in the large rivers and moving relatively short distances to clear
streams to spawn. The Minto Flats population spawns chiefly in the Tolovana
and Chatanika Rivers, with a few entering the Chena River and other clear
streams. The fish of the upper Yukon apparently utilize almost every major
tributary.
28
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Fish of the Kuskokwim and lower Yukon rivers overwinter in the delta areas
of these streams, while the Kobuk-Selawik population spends the winter in the
brackish waters of Selawik Lake and Hotham Inlet. These fish may move as far
as 670 km up the Kobuk River, while those from the mouth of the Yukon travel
1,600 km to spawn in the Alatna River (Alt, 19715, 1973a).
Upstream migration from the wintering areas begins at ice breakup. The
early movements appear to be associated with feeding, but, the time depending
on the distance to be travelled, this soon becomes a definite migration to
the spawning area. The true spawning migration may last only a few weeks, or,
as with the lower Yukon fish, may take as long as four months. During this
spawning migration, the inconnu feed little, if at all, but, unlike salmon,
they are still in good condition when they reach the spawning grounds in late
September. They appear to utilize excess visceral fat during the migration
(Alt, 1969).
Spawning takes place in late September and early October, when the water
temperature is between,1.4 and 4.6°C. Following spawning, there is a fairly
rapid downstream migration to the wintering grounds, where the inconnu again
commence to feed, it is not certain whether inconnu spawn annually or at
longer intervals. Russian fish appear to spawn only every third or fourth
year (Nikolskii, 1961). Alt (1969) found two stages of gonad development in
mature fish at Selawik and suggested that this might indicate spawning every
other year. On the other hand, he felt that all mature fish in the Kobuk
River were spawners. It may be that non-spawners do not migrate very far
upstream.
IMPORTANCE TO MAN
Wherever it is found, the inconnu is prized for sport, subsistence or
commercial purposes. It is an excellent sport fish. Its fighting qualities
and size make it a trophy worthy of any angler's best endeavors. As food, it
is likewise excellent. The flesh is white, sweet, slightly oily. It has not
always enjoyed a high reputation, however, for Richardson (1823) wrote that
it was "disagreeable when used as daily food."
Subsistence fisheries of considerable magnitude exist along the Kobuk,
Selawik, lower Yukon, Koyokuk and Kuskokwim rivers. In the early 1960's,
the subsistence catch on the lower Yukon was on the order of 5,000 fish an-
nually, while the Kuskokwim produced about 1,600 - 1,700. No data are avail-
able for the Koyokuk, but statements from residents of Hughes suggest that the
subsistence take on that river is in excess of 2,000 fish a year. The great-
est subsistence fishery is in the Kobuk-Selawik area, where Alt (1969) esti-
mated a catch of 19,240-22,000 fish in 1965. This represents something like
50,000 kg, of which about 900 kg were sold commercially.
The only commercial fishery for Alaskan inconnu is in the Kotzebue area.
This fishery has produced up to 45,000 kg per year (Wigutoff and Carlson, 1950),
The total catch, both subsistence and commerical, from northwest Alaska in
1965 was estimated at 34,200 - 37,000 fish, or nearly 90,000 kg.
29
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Commercial fisheries for inconnu also exist in Great Slave Lake, with
annual yields on the order of 144,000 kg (Sinclair et al, 1967), and in the
rivers of Siberia. The annual catch from the latter region between 1936 and
1940 was reported as about 4.15 million kg per year (Nikolskii, 1961).
LEAST CI-SCO
Coregonus sardinella Valenciennes
DISTINCTIVE CHARACTERS
Adults of the least Cisco are easily distinguished from those of other
Alaskan ciscoes by the lower jaw protruding slightly beyond the upper when
the mouth is closed, and by the dusky to black pelvic fins. The latter
characteristic, however, does not appear until the fish are about 150 mm long,
so identification of the young is sometimes difficult.
DESCRIPTION
Body slightly to moderately compressed, its greatest depth 19-24% of
total length. Head rather short, less than 251 of total length. Snout about
25% of head length. Eye round, large, 26-32% of head length. Nostrils with
a double flap between the openings. Mouth moderate, lower jaw protruding
slightly beyond upper when mouth is closed. Maxillary reaching backward to
below anterior half of eye. No teeth on jaws, vomer or palatines, but a
patch of small teeth present on tongue. Gill rakers long and slender, 42-53
on first arch. Branchiostegal rays 8-9. Lateral line with 78-98 pored scales.
Fins
D. 12-14, rather high and falcate. Dorsal adipose fin present. A. 11-13,
P,. 14-17, narrow. P2- 8-12, a distinct axillary process present. Caudal
fin forked.
Scales
Moderately large, cycloid. Statements that "spawning males probably
develop tubercles along the sides" (McPhail and Lindsey, 1970) are in error.
At least in interior Alaska, spawning tubercles are not developed by either
sex.
Color
Brownish to dark greenish above, silvery below.
Size
The least Cisco is known to reach fork length of at least 413 mm in in-
terior Alaska (Kepler, 1973). Scott and Grossman (1973) gave 419 mm (total
length or fork length?) for Canadian fish, while Nikolskii (1961) indicated
a maximum size of 420 mm and 500 g for fish from Siberia.
30
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RANGE AND ABUNDANCE
Present in most streams and lakes of Alaska north of the Alaska Range,
from Bristol Bay to the arctic coast. The least cisco is present throughout
the Kuskokwim and Yukon drainages, ranges south in the Mackenzie to Fort
Simpson and eastward along the arctic coast of Canada to Bathurst Inlet and
Cambridge Bay. McPhail and Lindsey (1970) noted its presence on Victoria and
Banks islands in the Arctic Ocean. Westward, the range extends across the
Bering Strait and Siberia as far as the White Sea.
The least cisco is one of the more abundant freshwater fishes of Alaska,
but because of its migratory habits the abundance is seasonal.
HABITS
The least cisco spawns in late September and early October. Spawning
takes place at night, with peak activity between 2000 hours and midnight. The
spawning grounds in the Chatanika River, near Fairbanks, are confined to a
stretch of river from 16 km below to 12 km above the Elliott Highway bridge.
Individual spawning areas vary in size from 100-800 in in length and 15-22 m in
width. Water depth ranges between 1.3 and 2.6 m, with average velocity of
about 0.5 m/sec. Surface temperatures during the spawning period are between
0° and 3°C (Kepler, 1973). The stream bottom in the spawning areas is com-
posed of gravel with a little sand.
In the spawning act, a female swims almost vertically upwards, with her
ventral side upstream. She is joined by as many as five males (usually only
one or two), who swim upward close to the female. As the s'pawners approach
the surface, eggs and milt are released. A small cloud of milt can sometimes
be seen as the fish enter the upper 20-30 cm of water. The fish break the
surface, fall over backwards and swim back to the bottom of the pool. It is
not known whether a female deposits all her eggs in one night, or whether two
or more nights are required.
The eggs, somewhat less than 1 mm in diameter and cream to yellow in
color, sink to the bottom where they lodge in crevices in the gravel. They
are not adhesive. Fecundity varies between 9,800 and 93,500 eggs per female,
averaging about 54,350. Egg number is more closely related to age than to
size of the individual (Kepler, 1973). The Chatanika River ciscoes are more
fecund than those of Siberia, for Nikolskii (1961) gave only 23,600 as the
maximum egg count for fish from the Yenisei River.
The eggs over-winter in the gravel and hatch early in the spring. By
mid-June, large numbers of young of the year, which may be up to 40 mm-long,
»are moving downstream to deeper, slower water (Townsend and Kepler, 1974).
Males first become sexually mature at age 2+, with the majority maturing
at 3+. Females begin to mature at 3+ (rarely 2+), but most mature at 4+.
A large, mature female may weigh as much as 2,5 kg (5.5 Ib) (Alt, 1971 a.)
31
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Occasional hybrids between least Cisco and Alaska whitefish occur (UAFC
2173), but despite the fact that the two species spawn more or less simul-
taneously in the same areas and have similar breeding behavior, such hybrids
are rare.
The maximum age attained in most populations seems to be between 8 and 11
years (Cohen, 1954; Nikolskii, 1961; Kepler, 1973), although Scott and Cross-
man (1973) mentioned that ages approaching 26 years have been recorded in
specimens from Victoria Island, N, W. T.
Growth rates vary widely from place to place, and even within the same
habitat. Although stream-dwelling, migratory populations are generally stated
to grow faster and live longer than lake-dwelling, non-migratory fish (McPhail
and Lindsey, 1970; Scott and Grossman, 1973), Cohen (1954) found that the an-
adromous fish at Point Barrow were slower growing and thinner than a non-
migratory population in Ikroavik Lake. To further confound the situation, a
second population in Ikroavik Lake was slower growing and shorter lived than
the first two.
Kepler's (1973) data on spawning females from the Chatanika River yield
the following average fork lengths at ages from 2+ to 8+: 2+, 311 mm (2 fish);
3-4-, 316 mm (4); 4+, 333 mm (15); 5+, 370 mm (12); 6+, 383 mm (5); 7+, 413 mm
(1); 8+, 405 mm (2). Alt (1971a), on the basis of back-calculating from
scales, gave the following fork lengths at the end of each year of life for
Chatanika River fish: 1, 120 mm; 2, 208 mm; 3, 261 mm; 4, 304 mm; 5, 337 mm;
6, 364 mm; 7, 387 mm; 8, 410 mm.
Although lake-dwelling populations appear to be non-migratory, those
living in streams or reaching brackish water undertake considerable movements
to reach or leave their spawning grounds. As already noted, young of the
year move off the spawning grounds shortly after spring break-up. The up-
stream spawning migration of adults begins in early July in the Chatanika
River and is completed by late September.
Least Cisco feed primarily on various types of zooplankton, including
various small copepods, cladocerans, mysids and the adults and larvae of a
variety of insects. Plant material may also be eaten. The ciscoes normally
do not feed during the spawning run. (Nikolskii, 1961; Furniss, 1974; Morrow
et al, 1977).
Least cisco are preyed upon by many predators, including eagles, hawks,
kingfisher, pike, inconnu, lake trout, burbot, man, and, no doubt, any others
capable of catching them. The eggs may be eaten by grayling and Alaska
whitefish during spawning (Morrow et al, 1977).
IMPORTANCE TO MAN
In North America, the least Cisco is relatively unimportant. It is
taken by subsistence fisheries in Alaska and northern Canada, usually as an
incidental in nets set primarily for other whitefishes, pike or grayling. A
small sport spear fishery exists in the Chatanika River in interior Alaska.
32
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Least cisco is an important commercial fish in Siberia. Annual landings in
the late 1930's were in excess of 1,000,000 kg (2,200,000 Ib) (Nikolskii,
1961).
As a food fish, the least Cisco is generally considered somewhat in-
ferior to the humpback whitefishes. However, it is a very good fish, with
firm, tasty meat.
BERING CISCO
Coregonus laurettae Bean
DISTINCTIVE CHARACTERS
The pale pelvic and pectoral fins distinguish the Bering cisco from the
least cisco, while the lower number of gill rakers on the lower portion of
the first gill arch (18-25 vs 26-31) distinguish it from the arctic Cisco.
DESCRIPTION
Body rather elongate, slightly compressed. Depth about 2Q% of total
length. Head moderate, 22-25% of total length. Snout 20-25% of head length.
Eye about equal to snout, round. Nostrils with a double flap between the
openings on each side. Mouth moderate, terminals upper and lower jaws equal.
Maxilla reaching backward to middle of eye. Usually no teeth on jaws, but
weak teeth present on maxilla in young and a few small teeth rarely present
on lower jaw of adults. A small patch of teeth present on tongue. Gill
rakers 18-25 on lower portion of first gill arch, total count on first arch
35-39. Branchiostegals 8-9. Lateral line with 76-95 pored scales.
Fins
D. 11-13, rather high and falcate. Dorsal adipose fin present, A. 12-
14. PT. 14-17. P2. 10-12. Caudal fin forked.
Scales
Cycloid, fairly large.
Color
Generally brownish to dark green on back, silvery on lower sides and
belly. Anal, pelvic and pectoral fins pale, caudal and dorsal dusky.
Size
The largest known, recorded by Alt (1973b,c), was a female of 480 mm
fork length from the lower 500 m of Hess Creek, Alaska. The average size of
adults is about 300 mm (1 ft).
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RANGE AND ABUNDANCE
The Bering Cisco is found from Bristol Bay north and east to the mouth
of the Qliktok River on the arctic coast of Alaska. It is present in the
Yukon River as far upstream as Fort Yukon, and also in the Porcupine River.
It has also been found at the mouth of Ship Creek, Knik Arm, at Anchorage,
Alaska (SU 41858, now in the Ichthyological Collection of the California
Academy of Sciences,(McPhail, 1966}),in Tolugak Lake in Anaktuvuk Pass in the
Brooks Range (UAFC 617,618), and was found in 1972 in the Kenai River on the
Kenai Peninsula of Alaska (K, T. Alt, Alaska Dept. Fish Game, pers. comm.).
Throughout its range it is fairly abundant, at lease seasonally, for Alt
(1973b) reported up to 18 a day taken in a fish wheel at Rampart, on the
Yukon River, in September 1972.
HABITS
Very little is known of the biology of the Bering cisco.
following is derived from Alt (1973b,c).
Most of the
Spawning runs begin in the spring. Most of the Bering ciscoes apparently
winter in salt or brackish water near river mouths, but the presence of po-
tential spawners well up the Yukon and Kuskokwim rivers suggests that some
populations may spend the winter in fresh water, far from the sea. Bering
Cisco were first observed well inland in 1968 and 1969, when one (UAFC 2176)
was taken in the Chatanika River, near Fairbanks, and seven (UAFC 632) at
Rampart, on the Yukon River. Two more specimens (UAFC 617, 618) are from
Tolugak Lake in Anaktuvuk Pass. Subsequently, Alt (1973b) found Bering cisco
in the Porcupine River, 1,400 km from the mouth of the Yukon; in the Yukon
River at Fort Yukon; and in the South Fork of the Kuskokwim River, 840 km
from the ocean.
Spawning probably takes place in the fall, but spawning behavior and the
location of the spawning grounds are unknown. From the distribution of the
fish in June, it may be presumed that the spawning grounds are in clear-water
streams tributary to major rivers. Dymond (1943) mentioned C_. laurettae X
Stenodus 1eucichthys hybrids, but the location whence the specimens came, the
Mackenzie River delta, suggests strongly that they were actually hybrids be-
tween the arctic cisco, £. autumnal is and the inconnu.
Alt (1973b) stated that the majority of his specimens from Hess Creek
were 4+ to 6+ and were mature. His fish from Port Clarence and Grantley
Harbor were mostly 2+ and 3+ immatures, as well as a few adults, which showed
slower growth than the Hess Creek fish, possibly because of a shorter growing
season. Mean fork lengths at age were, for Hess Creek: 4+, 344 mm; 5+, 354
mm; 6+, 373 mm; 7+, 405 mm; 8+, 446 mm; and for the Port Clarence-Grant!ey
Harbor fish: 3+, 241 mm; 4+, 263 mm; 5+, 285 mm; 6+, 313 mm; 7+, 350 mm.
As noted above, Bering cisco undertake extensive spawning migrations.
Presumably they move downstream after spawning. The precise extent of the
migrations is unknown.
34
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The Bering Cisco, like other ciscoes, apparently does not feed on its
spawning runs. All the fish examined from the Yukon and Kuskokwim rivers
(June through September) had empty stomachs. By contrast, fish taken at
Port Clarence-Grant!ey Harbor had fed on invertebrates and small cottids
(Alt, 1973b,c). McPhail and Lindsey (1970) listed amphipods as food of the
Bering cisco.
IMPORTANCE TO MAN
The Bering cisco is little utilized. Small numbers are taken for sub-
sistence use by gill net and fish wheel in the Yukon and Kuskokwim rivers.
Similar utilization probably exists wherever subsistence fishing and the
Bering cisco coincide.
ARCTIC CISCO
Coregonus autumnal is (Pallas)
DISTINCTIVE CHARACTERS
Pale pelvic fins, a terminal mouth and 41-48 gill rakers on the first
arch serve to distinguish the arctic cisco.
DESCRIPTION
Body elongate, slightly compressed. Depth 20-23% of total length. Head
moderate, somewhat less than 25% of total length. Snout about 25% of head, a
little longer than eye diameter. Eye round, 20-24% of head. Nostrils with a
double flap between the openings. Mouth moderate, jaws toothless in adults
(a few weak teeth may be present in very small young), a patch of teeth on
tongue. Maxilla extends backward about to middle of eye. Gill rakers 41-48
on first arch. Branchostegals 8-9. Lateral line with 82-110 pored scales.
F i ns
D. 10-12, fairly high, slightly falcate. Dorsal adipose fin present.
A. 12-14.. P]. 14-17. ?2> "n-12- Caudal forked.
Seales_
Moderately large, cycloid, 82-110 pored scales in lateral line.
Col or
Brown to dark greenish above fading to silvery on sides and belly. Fins
pale.
S ize
Specimens up to 640 mm total length and up to 2,685 g weight have been
reported from the Lena River in Siberia (Berg, 1948). However, North American
35
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specimens generally are much smaller, averaging somewhere in the neighborhood
of 350-400 mm long (Roguski and Komarek, 1971} and perhaps a kilogram (2.2 1b)
in weight.
RANGE AND ABUNDANCE
In North America, the arctic cicso is found along the arctic coast of
Canada and Alaska from the southeastern end of Victoria Island westward to
Point Barrow and south in the Mackenzie tp the Liard River. It ranges
westward across arctic Siberia to the White Sea. Due to its migratory habits,
it is abundant seasonally.
HABITS
The arctic cisco is truly anadromous. It seems to be much more tolerant
of salt water than most coregonids, for it has been taken in water up to
22 ppt salinity (Roguski and Komarek, 1971). The upstream spawning run takes
place in the summer and may cover as much as 1,000 km (1,600 miles) (Nikolskii.
1961). Spawning occurs over gravel in fairly swift water. The eggs are
broadcast and left to fend for themselves. Mature females may produce as many
as 90,000 eggs (Berg, 1948). After spawning, the adults return downstream.
The arctic cisco occasionally hybridizes with the inconnu (Kuznetsov, 1932;
Dymond, 1943).
The young probably hatch in the spring and descend the rivers to
estuaries, as do other ciscoes.
Little is known of growth rates in North America. Roguski and Komarek
(1971) and Kogl (1971) have provided data on 65 specimens from the Beaufort
Sea and the Colville River delta, as follows: Age I, 114-115 mm fork length;
II, 136 mm, III,'233 mm; IV, 297 mm; V, 317 mm; VI, 325 mm; VII, 373 mm;
VIII, 414 mm. This is somewhat faster growth than was listed by Berg (1948)
for Yenisei River ciscoes. The fish become mature at about six years of age
and apparently do not breed every year (Nikolskii, 1961; Roguski and Komarek,
1971).
The arctic cisco shows a wide range in its feeding and has been found to
eat mysids, copepods, amphipods, isopods, chironomids and other insects and a
variety of small fishes (Nikolskii, 1961; Kogl, 1971).
IMPORTANCE TO MAN
In North America,, the Arctic Cisco is used primarily in subsistence
fisheries, and these may be of considerable magnitude. The fish are caught
"in abundance" during August at Barter Island, where it is preferred over
charr (Furniss, 1974). A small commercial fishery on the Colville River
delta takes about 11,000 kg (24,200 Ib) per year, about 601 of the total
catch of all species there (Winslow and Roguski, 1970). The arctic cisco is
an important component of the fisheries along the Mackenzie River (Wynne-
Edwards, 1952) and it is an important commercial fish in Siberia and the
eastern parts of northern USSR (Nikolskii, 1961).
36
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PYGMY WHITEFISH
Prosopium coulteri (Eigenmann and Eigenmann)
DISTINCTIVE CHARACTERS
The pygmy whitefish is distinguished by its bluntly rounded snout, less
than 70 pored scales in the lateral line and 14-33 pyloric caeca.
DESCRIPTION
Body elongate, round in cross-section. Depth about 17% of fork length.
Head moderate, its length about 22% of fork length. Snout short, broadly
rounded as seen from above. Eye large, round, diameter usually greater than
snout length, a notch present in lower posterior portion of adipose lid.
Nostrils with a single flap between the openings. Mouth small, subterminal,
maxillary reaches to anterior portion of eye. No teeth on jaws, vomer or
palatines, but a small patch of fine teeth present on tongue. Gill rakers
short, the longest 6.5-9% of head length, 12-21 on first arch. Branchi-
ostegal rays 6-9. Lateral line with 50-70 pored scales.
Fins
D. 10-13, a small adipose dorsal fin present. A. 10-14. P]. 13-18.
?2- 9-11, axillary process present. Caudal fin forked.
Scales
Large, round, cycloid.
Color
Brownish, often with greenish tints, above; sides silvery, belly white.
A row of about 11 large, round parr marks along sides of young, persisting in
all but the largest adults.
Size
This is the smallest of the Alaskan coregonids. The largest specimens
recorded were only 271 mm fork length (McCart, 1965) and most are scarcely
half that size.
RANGE AND ABUNDANCE
The pygmy whitefish has a most remarkably discontinuous distribution.
It is known from Lake Superior; from parts of the Columbia, Fraser, Skeena,
Alsek, Peace, Liard and upper Yukon systems in Washington, Montana, British
Columbia and the Yukon Territory; and from the Chignik, Naknek and Wood river
systems in southwestern Alaska. It is not known outside of North America.
Wherever it is found, it seems to be quite abundant.
37
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HABITS
Details of breeding behavior are unknown. Spawning takes place at night
in late fall and early winter (November-December) at water temperatures of
about 4°C or colder. Spawning grounds appear to be on gravel in lake shallows
and in streams. Presumably, the eggs are broadcast, settle into interstices
in the gravel and hatch the following spring. In the Bristol Bay region, the
eggs are about 2.4 mm in diameter (Heard and Hartman, 1966), somewhat larger
than the 2.0 mm reported for Lake Superior fish (Eschmeyer and Bailey, 1955).
Egg number varies between 103 and 1153 per female in Bristol Bay fish, a
greater average number than in Lake Superior fish.
Growth of the pygmy whitefish is very slow and shows considerable varia-
tion from one population to another. In general, females grow faster than
males, although in some populations the males grow faster during the first
year (Eschmeyer and Bailey, 1955). In Brooks Lake, Alaska, the average three-
year-old fish was about 70 mm fork length, while in South Bay, Naknek Lake,
the same aged fish averaged about 116 mm. The maximum age recorded for fish
in the Naknek system was 5+ (Heard and Hartman, 1966), but ages up to 9+ have
been noted in Maclure Lake, British Columbia (McCart, 1965).
The pygmy whitefish does not make extended migrations. However, it does
move on to the spawning areas in the early winter and presumably back into
deeper water after spawning.
Food of the pygmy whitefish includes a rather wide variety of items.
Listed as most important in the Naknek system were cladocerans, dipteran
(chiefly Chironomidae) larvae and pupae, adult Diptera, and nymphs of
Plecoptera. Other items included diatoms and other algae, pelecypods, nerria-
todes, arachnids and fish eggs (Kendall, 1921; Heard and Hartman, 1966). In
Lake Superior, ostracods and amphipods were the principal foods (Eschmeyer
and Bailey, 1955): The pygmy whitefish feeds almost exclusively during day-
light hours. In its feeding, the pygmy whitefish makes "short distinct jabs
or darts, apparently at specific food items, such as insect larvae, when pick-
ing up mouthfuls of bottom material." They may also rise off the bottom and
take specific items from the current (Heard and Hartman, 1966).
There is a distinct positive correlation between the diet and the average
size of individuals in a population. Groups in which insects are the dietary
mainstay average much larger than those in which zooplankton is the chief
food (Heard and Hartman, 1966). McCart (1970) found high and low gill raker
count forms in Aleknagik, Naknek and Chignik lakes in Alaska and suggested
that these represented sibling species. The high count form was found almost
exclusively in deep water, fed on plankton and grew more slowly. By contrast,
the low count form was found in both shallow and deep water, ate mostly in-
sects and grew faster. However, the difference in growth rates through age
IV in Chignik Lake does not appear to be significant. More careful analyses
of larger samples are needed.
As noted above, the pygmy whitefish is found in both deep and shallow
water. In Lake Superior, it was reported as most abundant at depths of
38
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46-71 m (Eschmeyer and Bailey, 1955; Dryer, 1966). In the Naknek system,
the species was found at depths to 168 m but was also abundant in the shallows
(Heard and Hartman, 1966). McCart (1970) found that, in Chignik Lake, beach
seine samples were composed entirely of the low gill raker count form, while
in gill nets set at 30 m or deeper, the high count form accounted for 36.2%
of the fish taken.
IMPORTANCE TO MAN
The pygmy whitefish is of no direct importance to man. It is too small
and scarce and of too limited distribution to be a profitable object for any
kind of fishery. However, it is undoubtedly fed upon by predatory fishes
such as charr, pike, burbot, etc., and may contribute to the overall scheme
of competition. In this respect, it is interesting to note that the pgymy
whitefish attains its greatest size in waters where there is no competition
from other coregonids (McCart, 1965).
ROUND WHITEFISH
Prosopiurn cylindraceum (Pallas)
DISTINCTIVE CHARACTERS
The narrow, rather pointed snout, more than 70 pored scales in the
lateral line, and 50 or more pyloric caeca distinguish the round whitefish.
DESCRIPTION
Body elongate, cylindrical, slender. Depth 15-201 of fork length. Head
relatively short, its length averaging about 201 of fork length. Snout short,
about 22% of head, pointed as seen from above. Eye round, its diameter equal
to or less than snout length, notch present in membrane below posterior edge
of pupil. Nostrils with a single flap separating the openings on each side.
Mouth small, upper jaw overhanging lower, maxilla reaches about to anterior
margin of eye in adults, a little farther back in young. Teeth restricted to
a small patch of embedded teeth on tongue, also present on bases of gill
rakers. Gill rakers short, 14-21. Branchiostegal rays 6-9 on each side.
Lateral line with 74-108 pored scales. Pyloric caeca 50-130.
Fins
D. 11-15, an adipose dorsal
Caudal fin forked.
Scales
fin present. A. 10-13. PI. 14-17. Pg. 9-11
Fairly large, cycloid, nuptial tubercles prominent on lateral scales of
breeding males, but only feebly developed in females.
39
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Color
Bronze dorsally, sometimes with a greenish tinge. Sides silvery, belly
silvery white. Fins of most Alaskan'specimens are more or less colorless or
slightly dusky. Scott and Grossman (1973) reported the pectoral, pelvic and
anal fins to be amber colored, becoming orange at spawning time. The young
are marked with three rows of rather well-defined parr marks. One row lies
along the lateral line, a second row (sometimes ill-defined) just above the
first, and a third row just below the mid-line of the back. The spots of
this third row often coalesce across the mid-dorsal line.
Size
The largest on record is a specimen of 561 mm total length from Great
Slave Lake (Scott and Grossman, 1973). The round whitefish is known to reach
a weight of 2 kg (Keleher, 1961) and has been reported as reaching "about 5
pounds" (2.27 kg) in Lake Superior (Koelz, 1929).
RANGE AND ABUNDANCE
The round whitefish is found throughout mainland Alaska from the Taku
River, near Juneau, north to the arctic coast. It ranges eastward across
Canada to the western shores of Hudson Bay. A discontinuity in range exists
in Manitoba and northern Ontario, and the species is again present in the
Great Lakes (except Lake Erie), eastward to New Hampshire and Maine, south
to Connecticut and north to the arctic coast of Labrador. In Asia, the round
whitefish ranges west to the Yenisei River, south to Kamschatka.
The round whitefish is fairly abundant wherever it is present, although
it usually does not occur in such large numbers as some of its relatives.
HABITS
Spawning occurs in the late fall, late September through October, in
interior Alaska, but not unil November or December in more southern parts of
the range. Spawning appears to be an annual affair, with many fish breeding
in successive years, even in the arctic (McCart et al, 1972). The spawning
beds are located on gravelly shallows of rivers and the inshore areas of lakes.
Inshore and upstream migrations have been observed (Harper, 1948; Normandeau,
1969) at spawning time and are probably characteristic. However, fish in
interior Alaska do not seem to show the concentrated migrations characteristic
of ciscoes and humpback whitefish. According to Normandeau (1969), the fish
swim in pairs during spawning, a single male with each female. Details of
spawning behavior have not been described, but probably resemble those of the
pygmy whitefish, Prosopium coulteri. In that species, the fish contact each
other and rest on the bottom for 2-4 seconds, emitting eggs and milt, then
separate (Brown, 1952). The eggs of the round whitefish are known to be
broadcast and to receive no parental care. Females produce between 1,000 and
12,000 eggs, with the average between 5,000 and 6,000 (Bailey, 1963; Norman-
deau, 1969; Furniss, 1974). The size of ovarian eggs varies with locality.
In New Hampshire, unfertilized eggs averaged 2.7 mm in diameter (Normandeau,
40
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1969), but Furniss (1974) found ovarian eggs of Alaskan fish to be only 1.0-
1.8 (average 1.2) mm in diameter. The eggs absorb water after fertilization
and may reach diameters of 3 to almost 5 mm in a few hours. The eggs are
yellow to orange, demersal, though not sticky, and settle into crevices in
the rocks and gravel of the bottom. Time of development has been reported
as about 140 days at 2.2°C in New Hampshire (Normandeau, 1969), and pre-
sumably is not much different in Alaska. The young hatch out as sac fry. In
two to three weeks, the yolk has been absorbed and the young have left the
spawning grounds.
Growth rates vary from one locality to another. Lake Michigan fish grow
very rapidly, reaching a total length of about 500 mm in seven years (Mraz,
1964a). By contrast, in Elusive Lake, in the Brooks Range of Alaska, this
length is not achieved until the age of twelve. The oldest known round white-
fish is one of 16+ from Shainin Lake, Alaska (Furniss, 1974), Sexual maturity
is reached at about five years in the southern parts of the range, but not
until age seven in the Brooks Range of Alaska (Furniss, 1974).
Except for the spawning movements already mentioned, the round whitefish
apparently does not migrate.
Food of the round whitefish is primarily the immature stages of various
insects, especially Diptera and Trichoptera. Adult Trichoptera are also im-
portant, as well as gastropods, Daphnia and fish eggs (Martin, 1957; Loftus,
1958; Normandeau, 1969; Furniss, 1974). In some areas, the round whitefish is
considered a serious predator on the eggs of lake trout (Martin, 1957; Loftus,
1958).
IHPORTANCE TO MAN
The round whitefish was formerly taken in considerable quantities in
the Great Lakes. In the late 1920's, annual catches from northern Lake Michi-
gan were on the order of 90,900-163,200 kg (Mraz, 1964a), but present day
catches are much smaller, primarily because of the relatively small size of
the fish and an uncertain supply. In Alaska, the round whitefish is of some
importance in freshwater subsistence fisheries. It is occasionally smoked in'
strips and sold as "squaw candy."
BROAD WHITEFISH
Coregonus nasus (Pallas)
DISTINCTIVE CHARACTERS
The broad whitefish is set off by its short gill rakers, which are less
than 1/5 as long as the interorbital width, and the rounded to flat profile
of the head.
DESCRIPTION
Body elongate, compressed, especially in large specimens, sides rather
flatter than in most other whitefishes. Depth of body 23-31 £ of fork length
41
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in adults, more slender in young. Head short, 15-20% of fork length, dorsal -
profile rounded to flat (may be slightly concave in large specimens). Snout
blunt, rounded (sheep-nosed) in profile, short, its length about equal to or
less than diameter of eye. Eye small, 12-16% of head length. No notch in
adipose lid. Nostrils with a double flap between openings. Mouth small,
upper jaw overhanging lower, maxilla reaching posteriorly about to below an-
terior edge of eye. No teeth, except for a small patch of weak teeth on base
of tongue. Gill rakers 18-25, blunt, short, longest less than 1/5 (13-19%) of
interorbital width. Branchiostegals 8-9. Lateral line with 84-102 pored
scales.
Fins
D. 10-13, dorsal adipose fin present and fairly large. A. 11-14. P-j.
16-17. PZ- 11-12. Caudal fin forked.
Scales
Large, cycloid. Males develop prominent breeding tubercles on lateral
scales at spawning time, these only weakly developed in females.
Color
Olive brown to nearly black on back; sides silvery, often with a gray
cast; belly white to yellowish. Fins usually grayish in adults, pale in
young.
Size
This species is the largest of the Alaskan whitefishes. It is reported
to reach weights up to 16 kg in the Kolyma River of Siberia (Berg, 1948), but
most mature fish run around 2-5 kg. One of 715 mm weighing 5.7 kg from the
Yenisei River is mentioned by Berg (1948). The largest Alaskan specimen known
is a fish of 670 mm fork length from the Colville River at Umiat (Alt and
Kogl, 1973).
RANGE AND ABUNDANCE
The broad whitefish is found throughout Alaska from the Kuskokwim River
north to the arctic coast. It is present in the Yukon River from the mouth to
the headwaters. In the Tanana River drainage it is known from Minto Flats
and the Tolovana, Chatanika and Chena rivers and probably occurs further up-
stream as well. It is present in most, if not all9 of the rivers draining
into the Bering, Chukchi and Beaufort seas. The range extends eastward to
the Perry River, Northwest Territories, westward across Siberia to the
Pechora River, south to the Bay of Korf and to the Penzhina River on the Sea
of Okhotsk. It is fairly abundant seasonally, though apparently not as
numerous as some of its relatives.
42
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HABITS
Little is known of the biology of the broad whitefish. Although the
adults are more or less anadromous, those reaching the sea apparently do not
venture far from brackish water. Upstream spawning runs begin as early as
June and may extend into September or even later (Kogl, 1971; Alt and Kogl,
1973; Kepler, 1973; Townsend and Kepler, 1974). Spawning actually takes
place from September through October, possibly even into November. Wynne-
Edwards' (1952) statement that "The broad whitefish spawns in the rivers in
August..." is probably based on a misinterpretation of the timing of the
spawning runs. Except that spawning takes place in streams with gravel
bottoms, nothing is known of the breeding habits. Presumably they are similar
to those of other coregonids. The ovarian eggs are pale yellow to milky
white in color and up to 4 mm in diameter (Berg, 1948; Nikolskii, 1961).
Young hatch in the spring and move downstream. Adults apparently move down-
stream after spawning and over-winter in deep parts of the rivers or in
estuaries.
Growth is relatively slow, especially in the arctic. Berg (1948)
mentioned lengths of 500-530 mm at eight years for fish from the Kara and
Kolyma regions, but in the Colville River, Alaska, the%average length of
eight-year-olds was under 400 mm (Kogl, 1971). Broad whitefish from the
Minto Flats area grow at about the same rate as the Siberian fish (Alt and
Kogl, 1973). Maximum age recorded is 15 years (Alt and Kogl, 1973), al-
though Nikolskii (1961) stated that "The age limit of this fish exceeds 15
years."
The broad, whitefish appears to be mainly a bottom feeder. It is known
to eat chironomids, snails, bivalve molluscs (Kogl, 1971), mosquito larvae
(Berg, 1948) and crustaceans (Scott and Grossman, 1973).
IMPORTANCE TO MAN
The broad whitefish is an unimportant commerical species in Siberia,
where pre-World War II catches averaged 40,000 kg per year. In North America,
however, it is utilized almost exclusively in subsistence fisheries. A com-
mercial fishery in the Colville River delta takes about 7,000 kg per year.
Despite its lack of utilization, the broad whitefish is an excellent food fish.
43
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HUMPBACK WHITEFISHES
"Coregonus clupeaformls complex"
This group of three closely related species forms a most confusing
assemblage, for almost the only means by which they can be separated one
from another seems to be the modal number of gill rakers in large samples.
The form here called Co reg on us pidschian has average gill raker counts of
21-23, with range from about 17 to 24 or 25 in individual specimens.
Coregonus nelsoni averages 24 or 25 (mode usually 25) with a range of 22-27,
while Coreoionus clupeaformis has modal counts of 26 or more, with range of
individual counts from 24 to 33, Fisheries biologists in Alaska have applied
one or another of these names to humpback whitefish throughout the state, all
too often without adequate samples for proper identification. Hence, dis-
tributional records are often of little value.
There appear to be some differences in ecological relationships among
the three species. C_. clupeaformis is primarily a lake-dwelling form. C_.
nelsoni is mostly a stream dweller, only rather rarely being encountered in
lakes. It seems, also, to be intolerant of salt water. C_. pidschian appar-
ently is truly anadromous, at least in some areas, and may over-winter in the
sea near river mouths.
ALASKA WHITEFISH
Coregonus nelsoni Bean
DISTINCTIVE CHARACTERS
Gill rakers longer than 201 of interorbital width, 22-27 total gill
rakers on first arch (modal counts 24 or 25) and a pronounced hump behind the
head in adults are the distinctive marks of the Alaska whitefish.
DESCRIPTION
Body moderately compressed, sides rather flat. Depth of body usually
25% or more (up to 33%) of fork length in adults, increasing in larger fish.
Head short, less than 25% of fork length. Dorsal profile of head distinctly
concave behind eyes in adults due to the prominent nuchal hump. Snout
27-35% of head length. Eye small, its diameter 20-25% of head length, no
notch in lower posterior part of membrane. Nostrils with a double flap be-
tween openings. Mouth rather small, upper jaw over-hanging lower, maxilla
reaching backwards to below anterior third of eye. A few weak teeth present
on premaxilla in young, no teeth on jaws in adults. A few small teeth present
on tongue. Gill rakers 22-27, average total counts around 24 or 25. Longest
44
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raker longer than 20%
line with 77-95 pored
of interorbital
scales.
space. Branchiostegals 8-10. Lateral
D. 11-13. Dorsal adipose fin well developed, often larger in males than
in females. A. 10-14. P] . 15-17. P2. 11-12. Caudal fin forked.
Scales
Cycloid, fairly large, well-developed nuptial tubercles on lateral
scales of males, but less strong in females.
Color
Dark brown to midnight blue above, paling to silver on sides and white
beneath. No parr marks in young.
Size
Up to at least 532 mm fork length in the Chatanika River, Alaska (Alt,
1971a).
RANGE AND ABUNDANCE
The precise distribution of Alaska whitefish is uncertain, primarily be-
cause of the difficulty of identifying the three species of humpbacked
coregonids which occur in Alaska. However, as far as can presently be de-
termined, the Alaska whitefish seems to be pretty well confined to the Yukon
and its tributary drainages, where it is to be found all the way from Nulato
to the Canadian border. It is present in the Tanana River and the Koyokuk
River and their tributaries and in Lake Minchuniina. Specimens which may be
of this species have been reported from the Unalakleet and Wulik rivers.
Possible £. nelsoni are known from the Alsek, Copper and Susitna systems, ,
the upper parts of the Yukon River in Canada, the lower reaches of the
Mackenzie River and several lakes in western Canada (Lindsey, 1963a,b;
Lindsey et al , 1970; McPhail and Lindsey, 1970).
The Alaska whitefish is
sunnier and fall. Throughout
disperse widely.
HABITS
locally and seasonally abundant during the
the rest of the year, the fish apparently
The Alaska whitefish spawns from late September through October in in-
terior Alaska. Spawning areas are in clear, moderately swift streams with
fairly clean gravel bottom. In the Chatanika River, these areas are from
100 to 800 m long, 15-22 m wide, 1.3-2.6 m deep, with water velocities of
about 0.5 m per second. Water temperatures at spawning are between 0° and
3°C (Kepler, 1973). Average fecundity of 20 mature females five to ten
years old and 395-520 mm fork length was about 50,000 eggs. Fecundity was
not closely correlated with age or size of the fish (Townsend and Kepler, 1974)
45
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The spawning act is similar to that of the least cisco. A female begins
to swim vertically towards the surface, belly upstream. She is joined by a
male (sometimes two, rarely three). Eggs and milt are extruded as the fish
approach the surface of the water. The fish break the surface, fall away from
each other and return to the bottom of the pool. In contrast to the least
cisco, the Alaska whitefish spawns actively both at night and in the daytime.
The yellow to orange eggs, with an average diameter of 2.1-2.3 mm, drift down
to the bottom where they lodge in crevices in the gravel. The exact time of
incubation is unknown. However, young of the year have been taken in June and
July, so presumably the young fish hatch out in late winter or early spring.
The Alaska whitefish of the Chatanika River grow rapidly during their
early years. A one-year-old averages about 120 mm fork length. By the age
of five, the fish average between 350 and 400 mm fork length, and at ten
years about 485 mm. The oldest so far recorded is a 12+ of 532 mm fork length.
Sexual maturity is reached between the ages of three and five years (Alt, 1971 a;
Townsend and Kepler, 1974).
The Alaska whitefish of the interior undertakes fairly extensive upstream
and downstream movements. Upstream migration, apparently the beginning of the
spawning run, may start as early as late June. The migration seems to be
rather indefinite at first, but becomes marked as the season progresses and
more fish approach breeding condition. By September, schools of up to several
hundred fish are on or close to the spawning areas. Following the completion
of spawning, the majority of the fish move downstream, but a few may over-
winter in deep pools near the spawning grounds. The young of the year move
down stream in their first year of life and as a rule do not return to the
spawning areas until they are sexually mature.
Alaska whitefish generally return to the same spawning grounds year after
year. Townsend and Kepler (1974) found that five fish tagged in 1972 were
present on the same grounds in 1973. On the other hand, these same investi-
gators noted far fewer tag returns that were expected and suggested that this
might indicate either increased mortality of tagged fish or non-consecutive
spawning. The recent (October 1975) recovery at Nenana of an Alaska whitefish
tagged in the Chatanika River in 1974 (Townsend, pers. comm.) suggests that
some fish may wander far from their natal streams. Apparently all fish do not
return each year to the same spawning areas.
Hybrids of Alaska whitefish and sheefish (inconnu) are known (Alt, 1971c).
The two species spawn at the same time and in the same places. Because of the
differences in breeding behavior, hybridization is probably not the result of
inter-generic pairing but due rather to simultaneous broadcasting of sex pro-
ducts in the same area. Occasional hybridization occurs also between the
Alaska whitefish and the least cisco (UAFC 12173).
Alaska whitefish feed primarily on immature stages of insects, notably
Diptera and Trichoptera. Although they generally do not feed during the latter
part of the spawning run, this is not always so. On occasion they will feed
heavily on eggs of the least cisco (Morrow et al, 1977).
46
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IMPORTANCE TO MAN
The Alaska whitefish is an excellent food fish but is virtually un-
utilized. Its major importance lies in the subsistence fisheries, but even
here it falls far short of the various salmons. A small sport fishery in the
Chatanika River is estimated to take up to 500 fish yearly by spearing at
night (Kepler, 1973), and similar small spear fisheries exist at other loca-
tions in Alaska. Small commercial fisheries have operated in some of the
lakes of the Copper River drainage, but the take has not been large (Williams,
1968). The young are utilized by predatory fishes such as pike and burbot
(Alt, 1968), and by other predators such as kingfishers, mink, otter, etc.
HUHPBACK WHITEFISH
Coregonus pidschian (Gmelin)
DISTINCTIVE CHARACTERS
Gill rakers longer than 20% of interorbital width, 19 tp 24 or 25 gill
rakers with average counts of 22 or 23, and a pronounced hump behind the head
in adults distinguish the humpback whitefish.
DESCRIPTION
See description of £. nelsoni. Except for the gill raker counts, there
are no known morphological differences of any significance. It is our impres-
sion, as well as that of several fisheries biologists in the Fairbanks Office
of the Alaska Department of Fish and Game, that pearl organs are far fewer in
number and are less well developed in C_. pidschian than in C_. nelsoni. Speci-
mens from the Kobuk River (author's observation) and from Highpower Creek and
the Kolitna River in the Kuskokwim system {K. Alt, pers. comm.), all taken in
early October, had few pearl organs.
RANGE AND ABUNDANCE
The humpback whitefish is to be found in most of the Alaskan rivers that
empty into the Bering, Chukchi and Beaufort seas. It ranges throughout the
Kuskokwim River drainage, and well above Umiat in the Colvilie. Alt and
Kogl (1973) found it at Umiat in July, whence it is presumed that the spawning
grounds in that river must be much farther upstream. In the Yukon, on the
other hand, it apparently is confined to the lower reaches, where it has been
recorded from Marshall, Its range extends eastward along the Arctic coast at
least to the Sagavanirktok River, westward across Siberia to the Kara Sea.
Throughout its range, it is quite abundant during the spawning concentrations,
but the fish apparently disperse at other times of the year.
HABITS
Humpback whitefish appear to be truly anadromous, though it is not known
how far the over-wintering fish move from the river mouths. They have been
taken in the Beaufort Sea several miles offshore off the Colvilie and Sagava-
nirktok rivers, as well as in Kotzebue Sound, off Nome, and around the mouths
47
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of the Yukon and Kuskokwim rivers. In the Kara Sea of western Siberia, they
have been taken well out in the northern parts "which are characterized by
high salinities" {Berg, 1948). Upstream spawning migrations may be extensive.
Fish tagged in the Kuskokwim River below Bethel have been recovered at Medfra
and Telida on the North Fork, the latter representing a migration of not less
than 1,280 km. Possible C_. pidschian have been found in the Yukon River at
Fort Yukon and in the Porcupine River (K. Alt, Alaska Dept. Fish and Game,
Fairbanks, pers. comm.), but their origin remains unknown. Other populations
seldom venture far upstream and still others may never go to sea at all (Berg,
1948).
The spawning run generally begins in June and spawning usually occurs in
October. However, these fish have been found spawning under the ice in the
Kuskokwim River near Bethel as late as 15 November (Alt, pers. comm.) and
similar phenomena have been recorded in Siberia (Berg, 1948). Spawning behav-
ior has not been described, but presumably is similar to that of the Alaska
whitefish. Sexual maturity is attained at 4-6 years of age. Ovarian eggs
are reported as 1.2 mm in diameter in Siberian fish (Nikolskii, 1961).
Fecundity of females varies from one population to another and with the size
of the fish. The general range is from about 8,000 to nearly 50,000 eggs per
female. The young presumably hatch in the late winter and spring, subse-
quently moving downstream, to return as mature adults four to six years later.
The young feed mainly on zooplankton, but adults feed mostly on molluscs,
crustaceans and chironomid larvae (Nikolskii, 1961).
Growth rates vary greatly from place to place and even in different
sections of the same river (Nikolskii, 1961). In Alaska, fish in arctic rivers,
such as the Colville, Kobuk and Agiakpuk, grow much more slowly than do those
in the Kuskokwim and lower Yukon drainages. Huraback whitefish from the first
three rivers average about 267 mm fork length at 5 years and 405 mm at 10 years,
while those from the latter areas average 347 and 445 mm at the same ages (K.
Alt, Alaska Dept. Fish and Gamej Fairbanks, pers. comm.).
IMPORTANCE TO MAN
The humpback whitefish of Alaska is of little direct importance except
in local subsistence fisheries. A commercial operation on the Colville River
delta takes about 1,000 fish annually (Alt and Kogl, 1973). However, this
fish is an important commercial species in Siberia (Berg, 1948; Nikolskii,
1961).
LAKE WHITEFISH
Coregonus clupeaformis (Mitchi11)
DISTINCTIVE CHARACTERS
The lake whitefish is differentiated from the other two humpbacked white-
fishes of Alaska by its higher gill raker count, 26-33 total.
48
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DESCRIPTION
See description of C_. nelsoni. Except for the gill raker counts, there
are no known morphological differences of any significance.
RANGE AND ABUNDANCE
The lake whitefish is widely distributed across Canada and northern
United States, from the upper Yukon and Northwest Territories south to Montana,
Minnesota and the Great Lakes, east to New England, Quebec and Labrador.
Knowledge of its distribution in Alaska is uncertain, due to the confusion
with closely related species. However, it has been recorded with reasonable
certainty from Paxson and Crosswind lakes in the Copper River drainage and
from Lake Louise and the Tyone lakes in the Susitna drainage (Williams, 1968;
Van Wyhe and Peck, 1969). Lindsey et al (1970) showed a possible record of
lake whitefish from Old John Lake at the head of the Sheen jek River, but it was
based on only two specimens (Lindsey, pers. comrn.). Wherever it is found,
the lake whitefish is quite abundant, especially when schooled up for spawning.
HABITS
Breeding behavior of the lake whitefish is similar to that of the Alaska
whitefish, except that spawning generally takes place in inshore regions of
lakes. Stream populations, of course, utilize the rivers and creeks. Spawn-
ing takes place over rocky or gravelly bottom in depths of 1 to 3 m. A female
and one or more males rise to the surface, extrude eggs and milt, then descend
separately towards the bottom. Spawning occurs at night (Bean, 1903; Hart,
1930; Everhart, 1958). Adults breed annually in the southern parts of the
range, but apparently only every other year or even every third year in the
arctic and sub-arctic (Kennedy, 1953).
Fecundity varies greatly from one population to another, averaging
around 50,000 eggs per female, with a reported range from less than 6,000
to more than 150,000. Spawning occurs from October to December, depending
on locality, and seems to be associated with water temperatures of about
6°C or less. Hatching normally occurs in late April. Development of the
eggs takes 140 days at 0.5°C and this seems to be the optimum temperature
for the eggs. In laboratory studies, none survived at 0° or at 12°C. Mor-
talities through hatching increased from 27% at 0.5° to 411-421 at 2-6°, 81%
at 8° and 99% at 10°C. Abnormalities also increased from 0 at 0.5-2°C. to
at 10°C (Price, 1940).
The larvae are 11-14 mm long at hatching and grow rapidly during the
summer. In Lake Huron, the larvae are close inshore from soon after break-up
to the end of the summer (Faber, 1970), often associated with emergent vege-
tation. They stay at or near the 17°C isotherm (Reckahn, 1970), descending
with it to the metalimnion. Van Wyhe and Peck (1969) found similar movements
of young-of-the-year "lake whitefish" in Paxson Lake, Alaska.
Growth slows abruptly in September so that by the end of October the
larvae are about 120 mm long. This slowing of growth is associated with de-
scent into the colder water of the hypolimnion.
49
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Growth rates vary with locality and population. Average total lengths
at age, covering a wide variety of localities in the U. S. and southern Canada,
are: 1 year, 130 mm; 2, 216 mm; 5, 386 rum; 10, 511 mm; 15, 627 mm (Carlander,
1969). By contrast, "lake whitefish" in Paxson Lake, Alaska, had the follow-
ing age-length relationships: 1, 40 mm; 2, 77 mm; 5, 202 mm; 10, 373mm;
15, 460 mm (Van Wyhe and Peck, 1969). Maximum age reported is a fish of 28
years from great Slave Lake (Kennedy, 1953), while the largest is a fish of
19 kg taken in Lake Superior in 1918 (Van Oosten, 1946). If the length-weight
relationship given by Dryer (1963) for Lake Superior whitefish can be applied
to this specimen, then the fish must have been on the order of 1,350 mm total
length. The next largest known weighed just over half as much, 10.9 kq
(Keleher, 1961).
The lake whitefish appears to be a rather sedentary fish, at least in
the Great Lakes. Tagging studies (Budd, 1957; Dryer, 1964} indicate that the
majority of fish stay within 16 km of their spawning grounds, although one
fish in Lake Huron was recaptured 240 km from the point of release. There
seems, also, to be a tendency towards movement in definite directions, although
no well-defined routes were determined (Budd, 1957). In general, movement of
lake whitefish in large lakes consists of (a) travel from deep to shallow
water in the spring; (b) movement back into deep water during the summer as
the shoal water warms; (c) migration back to the shallow water spawning areas
in the fall and early winter; and (d) post-spawning movement back to deeper
water.
Within each of the Great Lakes, and probably in most large lakes, the
lake whitefish forms more or less discrete populations. These are usually
characterized by different growth rates rather than by morphological differ-
ences (Budd, 1957; Roelofs, 1958; Dryer, 1963, 1964; Mraz, 1964b). It is
not known whether these populations are genetically distinct or are environ-
mentally produced. In any case, the lack of migratory habits probably tends
to keep them separate.
Food of the lake whitefish varies with size and age of the fish, the
location and the type of food available. The initial food of the young
consists of copepods, later on of cladocerans. By early summer they begin
to feed on bottom organisms, but Cladocera, especially Bosmina, remain a
dominant food item for some time (Reckahn, 1970). Adults feed mainly on
benthic organisma, but pelagic and semi-pelagic forms also are important.
Kliewer (1970) found a significant negative correlation between gill raker
length and the proportion of benthic food, but a strong positive correlation
between the number of gill rakers and the amount of benthic food. He listed
the following food items for lake whitefish from the Cranberry Portage area
in northern Manitoba: Pelecypods, gastropods, amphipods, Diotera (tendipedid
larvae and pupae, culicid and ceratopogonid larvae), Ephemeroptera, Trichop-
tera, Megaloptera, plant material, fish eggs, Hirudinea, Cladocera, Conepoda,
mysids, Hemiptera (Corixidae), Hymenoptera, fishes. In Paxson Lake, Alaska,
adult whitefish were seen to prey upon young sockeye salmon until the fry
grew too big for the whitefish (Van Wyhe and Peck, 1969).
Although extensive hatchery programs for the propagation of lake white-
fish have been carried on for years on the Great Lakes and other places,
50
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there is no evidence to show that these programs have ever influenced the
strength of year-classes (Koelz, 1929; Christie, 1963). Weather seems to be
the most important factor. Cold water temperatures, below 6°C, at spawning
time, followed by, a steady (non-fluctuating) decrease to 0.5°C, and warm
temperatures at hatching time produce the strongest year-classes (Christie,
1963; Lawler, 1965a).
IMPORTANCE TO MAN
The lake whitefish has long been one of the most valuable freshwater
species in North America. Environmental deterioration, depletion of the
stocks and other factors led to a decline in yield from the 5,500,000 kg per
year of the 1880's to the 700,000 kg per year of the 1920's (Kqelz, 1929),
but in the late 1960's the catch was increasing. In 1970, the U.S. and
Canada landings from the Great Lakes and the international lakes amounted to
1,690,500 kg (Anonymous, 1973). In addition, there are considerable Canadian
fisheries in the northern lakes, such as Lake Winnipeg and Great Slave Lake.
In Alaska, however, the lake whitefish is virtually unutilized. Attempts at
commercial fishing for lake whitefish have been made in Crosswind Lake in the
upper Copper River drainage, and in Lake Louise and Tyone Lake in the upper
Susitna. These have not been especially successful (Williams, 1968).
CUTTHROAT TROUT
SaTtnp clarki Richardson
DISTINCTIVE CHARACTERS
The cutthroat trout is distinguished by having a red to orange mark on
the underside of each lower jaw, numerous small black spots over most of the
body, and a patch of small teeth (basibranchial teeth) behind the tongue be-
tween the gills.
DESCRIPTION
Body elongate, terete, little to moderately compressed, greatest depth
about 25% of FL, deeper in large specimens. Head about equal to body depth.
Snout rounded to slightly pointed, longer than eye diameter, especially in
spawning males. Eye round, its diameter about 22% of head length. Mouth
terminal, large, upper jaw reaching well behind eye in adults. Teeth canini-
form, well developed, present on both upper and lower jaws, tongue, head and
shaft of vomer, and palatines. A patch of minute teeth usually"present be-
tween gills behind base of tongue. Gill rakers 14-22. Branchiostegals 10-12.
Lateral line decurved anteriorly, 116-230 pores. Pyloric caeca 27-57.
Fins
D. 8-11. Adipose dorsal present. A. 11-13. P-j. 12-15. Pp. 9-10, a
distinct axillary process present. Caudal slightly forked.
51
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Scales
Cycloid, small, 116-230 (usually 120-180) along lateral line.
0
Color
Body dark greenish on back and upper sides, olivaceous on middle sides
fading to silvery below. Often a pinkish sheen on gill covers. Numerous
small black spots on back and sides, these extending to or almost to belly.
Spots also present on dorsal, anal and caudal fins. Adipose fin usually
with several spots, edge of adipose with an incomplete black border. Fins
of a uniform ground color, no white or colored borders. Underside of lower
jaw with a yellow to red (usually red) slash in the skin folds of each side.
Freshly sea-run fish usually more bluish, with silvery sides, inconspicuous
spots, and the red jaw marks inconspicuous or absent.
Size
The largest cutthroat known was a fish of the interior subspecies
(S_- clarki lewisi) taken in Pyramid Lake, Nevada, in 1925. This specimen
was 991 mm (39in") long and weighed 18.6 kg (41 Ib). However, the average
coastal cutthroat (_S. clarkl clarki) of 5 or 6 years of age will be only
250-380 mm (10-15 in) long.
RANGE AND ABUNDANCE
The coastal cutthroat ranges from the northern parts of Prince William
Sound, Alaska, south to the Eel River in Northern California, and is to be
found in most of the streams emptying into the Pacific within those limits.
It has been widely introduced into various streams and lakes within its
natural range, as well as into a few lakes in eastern North America.
Its abundance is irregular. In some streams it may be the most
numerous sport fish present, while other streams support only small popu-
lations. Its willingness to strike a lure, coupled with its low fecundity,
leads to adverse response to heavy fishing pressure.
HABITS
Coastal cutthroat spawn early in the spring, from April to mid-f4ay in
southeastern Alaska (Baade, 1975). In some parts of its range, the cutthroat
may spawn as early as February (Scott and Grossman, 1973).
Spawning takes place in small gravel-bottomed streams, as a rule
(Sumner, 1953, 1962; Baade, 1957; Lowry, 1965; Dewitt, 1954), but may go on
in the main stream of small drainages (Jones, 1975). At spawning time, the
male courts the female by nudging her and coming to rest beside her and
quivering. Having chosen the site for her redd or nest, the female begins
to dig out the egg pit in typical salmonid fashion. Heading upstream she
turns on her side, presses her tail against the bottom and gives five to
eight vigorous upward flaps of the tail. This sucks silt, debris and pebbles
out of the bottom whence they are carried downstream by the current. Digging
52
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in this fashion continues every few minutes, with occasional interruptions,
until the pit is deep and clean enough for egg laying. Such a pit is usually
100-150 mm (4-6 in) deep and not quite as long as the fish. Two to four hours
seems to be the usual time required to dig the redd.
Meanwhile, the male takes no part in the digging. He is very attentive
to the female, courting her and driving away intruders.
When the pit begins to take shape, the female tests it frequently by
sinking down into the pit and feeling around the stones with the tip of her
anal fin. Finally, she settles into the pit, with her anal fin pushed into
the gravel and her head and tail arched upwards. The male joins her at once,
both fish open their mouths and become rigid, quiver slightly, and eggs and
milt are extruded.
The fish now leave the pit, the female moves to its upstream end and
begins to dig again. The gravel she disturbs is carried down into the pit,
covering the eggs. She continues digging until the entire pit is filled and
the eggs well covered, a process requiring 40 minutes to an hour (Smith, 1941).
The new pit, upstream of the first, may also be used for spawning, or the
female may seek out a new spot and repeat the entire process. Both male and
female cutthroat may spawn with one or more members of the opposite sex.
Spawning goes on both in daylight and at night.
Fecundity varies with the size of the individual and with locality. Egg
numbers up to 6,500 have been reported from Wyoming, but Alaskan fish average
something less than 1,000 eggs per female (Baade, 1957; Jones, 1975).
The eggs, 4.3-5.1 mm in diameter, hatch in six to seven weeks and the
fry remain in the redd for one or two weeks longer before they become free-
swinming. The young are only about 15 mm (0.6 in) long at hatching, but grow
quite rapidly. Growth rate varies tremendously with stock and locality, but
overall, the young average 50-75 mm (2-3 in) long by the end of September.
By the end of the first year of life, most of the fish are around 100-150 mm
(4-6 in) long. At age 3, Alaskan fish average about 179 mm (7 in); 4, 210 mm
{ 8 1/4 in); 5, 254 mm (10 in); 6, 309 mm (12 in); 7, 332 mm (13 in); and 8,
380 mm (15 in) (Jones, 1973, 1975). Farther south, growth rates are faster.
About 90% of the years growth occurs between April and September (Coooer,
1970). Most young cutthroat do not go to sea until they are two or three
years old. Most populations of coastal cutthroats are dominated by fish 4-7
years old. Sexual maturity is reached in the second or third year of life,
with males generally maturing earlier than females. The young fish usually
stay in the stream for one or two years before going to sea, but some popula-
tions may never go to sea at all. In sea-going populations, the freshwater
life may be as long as eight years for rare individuals (Sumner, 1962; Jones,
1975).
The outmigration to the sea takes place in late spring and summer (mid-
April to July), with the peak of movement in late Hay and early June (Baade,
1957; Jones, 1973, 1975), but in the southern part of the range the migration
may be several months earlier (Lowry, 1965). Most movement goes on at night
on moderate stream flows. Extreme high or low water inhibits migration. In
general, the larger fish migrate to sea earlier in the season than do the
smaller ones. Cutthroat stay in the sea for 12 to 150 days in southeastern
53
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Alaska, those migrating earliest staying out the longest. It is possible
that some individuals may stay at sea for a year or more, as seems to be true
of some southern populations. However, most of the Alaskan cutthroats seem
to go to sea annually and to stay there a relatively short time (Baade, 1957;
Armstrong, 1971a; Jones, 1973, 1975). Most of the fish seem to stay fairly
close to the home stream, but tagged fish have been recovered as far as 70 km
(44 miles) from the mouth of the home stream (Jones, 1973).
Mortality of smolts (fish going to sea for the first time) is extremely
high, 98% or more between the beginning of the outmigration and their return
(Sumner, 1953, 1962).
The return migration from the sea to fresh water takes place in the sum-
mer and fall, with peak movement in September (Armstrong, 1971a; Jones, 1973,
1975). In Alaskan waters, this in-migration takes place mostly at night on mod-
erate stream flows, but in Oregon streams the cutthroat appear to move most ac-
tively during periods of high water and later in the year (Sumner, 1953; Lowry,
1965). It is apparent that this migration pattern is not firmly associated with
spawning, for in Petersburg Creek, Alaska, over half the incomers are immature
(Jones, 1973).
Post-spawning mortality in northern Oregon averaged 68.4% over a four-
year period, and ranged from a low of 54% to a high of 89%. Of the upstream
migrants, 68.4% had not spawned previously, 26.6% had spawned once, 4.5%
twice, and only 0.51 had spawned three times (Sumner, 1962). In Blue Lake,
northern California, only 5-10% survived to a second spawning, and these fish
produced significantly fewer eggs than did first-time spawners (Calhoun, 1944).
Post-spawning mortality has not been studied in Alaskan cutthroats.
The food of the cutthroat varies with locality and time of year. Fish
from Eva Lake on Baranof Island, Alaska, had fed chiefly on insects and young
salmon during the late summer; on sticklebacks, insects and gastropods in
winter; mainly on insects during the outmigration; and on amphipods and young
salmon while at sea." Near Ketchikan, Alaska, cutthroats fed on salmon eggs
and fry, insect larvae and sculpin eggs in the spring; on insects, sculpins,
coho fry and leeches in summer; and chiefly on salmon eggs in the fall.
Oregon fish were reported to eat aquatic and terrestrial arthropods, frogs,
earthworms, crayfish, small fishes and fish eggs (Baade, 1957; Lowry, 1966;
Armstrong, 1971).
Cutthroat trout prefer relatively small streams (watershed less than
13 km2), with gravel bottoms and gentle gradients (Hartman and Gill, 1968).
However, they may also be present in the sea, estuaries, lakes, and in alpine
lakes and streams to over 4,375 m altitude. The residents of these high
altitude habitats, however, belong to the inland subspecies.
IMPORTANCE TO MAN
The cutthroat trout is an important sport fish wherever it occurs. Al-
though generally considered inferior to the rainbow trout, it is nevertheless
a hard fighter, but does not jump as much as the rainbow. The cutthroat will
take a wide variety of lures - spinners, spoons, flies both wet and dry, and
small plugs. "Cutthroats are raised commercially in southwestern United States,
mainly for introduction into private ponds. The flesh is orange-red and of ex-
cellent flavor.
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RAINBOW TROUT
SaljTio gairdneri Richardson
DISTINCTIVE CHARACTERS
Black spots on the sides, back and on dorsal and anal fin, a reddish
band along the side (absent in sea-run fish) and no basibranchial teeth on
the base of the tongue serve to distinguish the rainbow trout.
DESCRIPTION
Body elongate, moderately compressed, greatest depth varying, according
to locality and size of fish, from about 21% to nearly 30% of standard length
(19-28% of fork length). Head about 20% of FL, larger in breeding males,
especially steelhead. Snout round, a little longer than eye diameter, but
much extended in breeding males. Eye round, about 20% of head length in fe-
males and non-breeding males. Mouth large, terminal, slightly oblique,
maxilla reaching to or well behind posterior edge of eye. Teeth caniniform,
well developed on both jaws, head and shaft of vomer, palatines and tongue.
No teeth on basibranchials at base of tongue. Gill rakers 15-22. Branchi-
ostegals 8-13 on each side. Lateral line with 100-150 pored scales, slightly
curved anteriorly, straight on most of sides and tail. Pyloric caeca 27-80.
Fins
D. 10-12, dorsal adipose fin present. A. 8-12. P-]. 11-17. Po. 9-10,
axillary process present. Caudal broad, slightly indented in smalt, but
quite square-edged in large specimens.
Scales
Small, cycloid, the size and number variable in different populations.
Color
Extremely variable according to locality, sexual condition and size of
fish. In general, top of head, back and upper sides dark blue to greenish or
brownish. Lower sides silvery, whitish or pale yellow. Belly silvery white
to grayish. Cheek and gill cover pinkish, sides with a band of bluish pink
to rose red, not present in fish fresh from the sea (steelhead). This band
reddest in spawning or recently spawned out fish, especially males. Back
and upper sides with many small, black spots, which may extend well down on
lower sides. Dorsal and caudal fins profusely spotted with black. Adipose
fin with black spots and a black border. Other fins with few spots, dusky or
unmarked. In some populations inhabiting Briston Bay drainages, red to orange
marks present on lower jaw, resembling cutthroat trout. Stream resident fish
may retain parr marks as adults.
55
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Si ze
Although in most areas a rainbow of 1 or 2 kg (2-5 Ibs) is considered a
very good fish, they are known to attain much greater size. The largest taken
by angling was a fish of 16.8 kg (37 Ibs) from Lake Pend Oreille, Idaho. A
fish of 19.1 kg (42 Ibs) was recorded from Corbett, Oregon (Field and Stream
Magazine, March, 1960) and one of 23.6 kg (52 Ibs) from Jewel Lake, B. C.
(McPhail and Lindsey, 1970).
RANGE AND ABUNDANCE
The original native range of the rainbow is from the Kuskokwim River in
Alaska south to northern Mexico near Ciudad Durango, about 24°N, westward in
Alaska to Port Holler on the Alaska Peninsula (160° 34' 30" W). The entire
range is west of the continental divide except for the headwaters of the
Peace River in British Columbia and the Athabasca River in Alberta.
Rainbow and steel head have been introduced into every continent and most
major islands of the world. Rainbow trout have been planted virtually all
over North America, with the result that the species is now present in every
state of the U. S. except Louisiana, Mississippi and Florida, and in every
province and territory of Canada except the Northwest Territories (MacCrimmon,
1971).
The rainbow is not native to interior Alaska (north of Alaska Range), but
has been planted in a number of lakes, ponds and gravel pits in the Fairbanks
area and near Big Delta, as well as Summit Lake. Lost Lake, about 48 km from
Big Delta, was rehabilitated in 1951 and received the first plant of young
rainbows in 1952. Alaska was thus the last state to practice planting rain-
bow trout. These first plants were made with fish raised from eggs imported
from Idaho and Montana, as this was less expensive than taking eggs of native
fish. (Marvich, 1952; Marvich and McRea, 1953; Marvich et al, 1954.)
HABITS
The rainbow trout is basically a spring spawner, with the majority
breeding between mid-April and late June (Lindsey et al, 1959; Hartman, 1959;
Hartman et al, 1962). However, various populations may spawn as early as
November or December (Dodge and MacCrimmon, 1970) or as late as August, and
fish have been bred in Idaho hatcheries to spawn in the fall. When these
latter were transplanted to warm temperatures (up to 17°C) in California,
they were able to spawn twice a year, summer and winter (Hume, 1955).
Spawning takes place in streams, usually on a riffle above a pool. Most
spawning occurs at temperatures between 10° and 13°C, but early spawners in
the north may encounter temperatures as low as 5.50C, while southern fish
have been known to spawn when water temperature was 17°C. Breeding behavior
is typically salmonid. The male courts the female by coming in contact with
her, sliding over her back, rubbing her with his snout, vibrating beside her
and pressing against her. When the female has selected a site for the redd,
she turns on her side and gives several strong, upward flips of her tail.
This displaces sand and gravel which is washed downstream by the current.
56
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Repeated digging soon results in a pit somewhat longer and deeper than her
body. Meanwhile, the attendant male has been courting her and driving off
rival males, although a subordinate male may also be in attendance. These
processes may go on both in daylight and at night.
When the redd is finished, the female drops down into the pit. The
dominant male joins her, the bodies of the two parallel and close together.
Both fish open their mouths, quiver, and extrude eggs and milt for a few
seconds. The subordinate male often participates in the spawning also. The
eggs fall into the interstices in the gravel, where they are fertilized.
From one or two hundred to as many as 1,000 eggs may be dropped in a single
nest. Fecundity per female ranges from as low as 200 (Needham, 1938) to as
high as 12,749 eggs, the latter from specially selected stock (Buss, 1960).
Average fecundity is on the order of 3,250 eggs per female, with younger and
smaller fish producing fewer eggs than larger, older fish. The number of eggs
produced depends in part on adequate nutrition. Insufficient food, such as
may be effected by overly large populations and excessive intra-specific
competition, leads to follicular atresia of the eggs, although the eggs which
survive are of normal size (Scott, 1962). As soon as the spawning act is
completed, the female moves to the upstream edge of the redd and digs again.
The displaced gravel is carried downstream and covers the eggs. The whole
process is repeated, either with the same or'other males, until the female's
egg supply is exhausted.
The eggs average between 3 and 5 mm in diameter and are pink to orange in
color. Time of development varies with temperature, ranging from as little as
18 days at 15.5°C to 101 days at 3.2°C (Embody, 1934; Wales, 1941; Knight,
1963), Under most natural conditions, development to hatching takes 4 to 7
weeks. The alevins require 3 days to more than 2 weeks to. absorb the yolk sac
completely, but normally begin to feed about 15 or 16 days after hatching, even
though some yolk may still be unabsorbed. The young emerge from the redds in
mid-June to mid-August for spring (April-May) spawners.
Survival of eggs is directly correlated with the velocity of ground water
passage through the redd and with the amount of dissolved oxygen in this ground
water (Coble, 1961). It has been found that wood fibers in the water do not
affect the survival of eggs but have pronounced adverse effects on growth and
survival of young fish (Kramer and Smith, 1965).
Subsequent life history varies greatly according to environmental con-
ditions and the genetic make-up of the population. Stream-resident fish
generally stay in the natal stream. By contrast, the young of lake-resident
fish usually move up or down stream to the lake in a few days to several
months. However, rainbows introduced into the Finger Lakes of New York have
been found to stay as long as 4 years (usually 1 or 2) in the streams before
moving down to the lake (Hartman, 1959). Movement of the young appears to be
associated with water temperature. In cold water, less than 13°C, the fry
rarely contact the bottom during the hours of darkness, and are carried down-
stream. By contrast, in warmer waters (over 14°C), the young fish make fre-
quent contact with the bottom and tend to stay nrore or less in one area. Up-
stream movements are associated with rapid increases in water temperature
(Northcote, 1963).
57
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Growth rates and growth patterns vary tremendously. Genetic composition,
type and availability of food, temperature, type of habitat and geographical
area all come into play. Sexual maturity is usually reached at two years or
older (rarely in the first year by some males) or as late as six years of age
in some females. In general, age at maturity is between 3 and 5 years, with
males usually maturing a year earlier than females. The size at sexual matur-
ity is equally variable. Mature individuals from small streams may be only
about 150 mm (6 in) long, while steelhead or fish in large lakes may be as
long as 400 mm (about 16 in). Maximum age seems to be about 9 years (Sumner,
1948).
Many rainbow trout spawn more than once. Particularly if environmental
conditions are good, rainbows may spawn annually for up to five successive
years (Hartman, 1959). The percentage of repeat spawners in a population
varies widely, from 5% to 57%. In general, larger and older females are less
likely to survive spawning than are smaller, younger ones, and males less
likely to survive than females (Hartman, 1959; Hartman et al, 1962; Withler,
1965). Survival to subsequent spawning seems to be inversely related to the
number of spawners entering a stream (Hartman, 1959).
Although the rainbow is generally considered a cold-water fish, with a
preferred temperature of about 13°C (Sarside and Tait, 1958), it can tolerate
fairly warm water, and indeed seems to grow best at about 21°c. The upper
lethal temperature is 24°C (Black, 1953).
The migratory patterns of rainbows are as varied as the populations and
the areas in which they are found. In general, stream-dwelling rainbows tend
to be non-migratory and to spend their entire lives in relatively short sections
of the stream. Lake-dwelling fish migrate into the spawning streams in the
spring, those going to inlet streams a month or more later. Both groups move
back into the lake 3 to 6 weeks after leaving it. This return seems to be
triggered by rising temperature. In Loon Lake, B. C., the return to the lake
occurred when the water temperature reached 10°C (Hartman, et al, 1962).
Steelhead undertake the greatest movements of any form of rainbow trout.
After 1-4 years (usually about 2) of stream life, the steelhead run down
stream in the spring and summer and enter the sea. Here they may stay from
a few months to as much as 4 years before returning to the natal stream to
spawn. Most of the freshwater movement, both up and downstream, goes on at
night (French and Wahle, 1959). At the time of the first downstream migration,
the young fish lose their parr marks and become silvery, the smolt stage.
This change in color is the result of increased activity of the thyroid gland
(Robertson, 1949). The younger fish adapt readily to sea water, but the longer
the period of freshwater life the longer the period of adaptation which is
required (Houston, 1961). The return migration may take place in spring-summer
(May-August) or in the winter (December-March). The fish of the latter group
are nearly ripe and spawn that same spring. By contrast, the spring-summer
fish stay almost a year in fresh water and spawn the following spring. At
spawning, these fish are typical rainbows, with the usual red band along the
side and the males with a well-developed kype. By contrast, winter fish show
little, if any, sexual dimorphism (Smith, 1960). Spring-run fish in California
have been reported to spawn in the late autumn (Briggs, 1953; Shapovalov and
58
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Taft,'1954). This division into spring-summer runs and winter runs appears
to be genetically controlled. It is probable that the evolution of the two
types has come about-in response to stream barriers which are passable only
during periods of high water (Withler, 1965).
Rainbows resident in fresh water generally do not migrate very far.
Stream-resident fish, in particular, may spend their entire lives within a
few kilometers of the place where they were hatched. Lake residents generally
do not move about much in the lake, but nay move up to 60 to 70 km, especially
up inlet streams. Steelhead, however, move extensively, undertaking upstream
migrations of hundreds of kilometers in large rivers such as the Columbia,
Steelhead introduced into Great Lakes drainages of Michigan have been known
to travel as much as 1320 km in 8 months through the lakes. Regardless of
strain, rainbows exhibit a high degree of homing. Better than 90% of sur-
viving migrants return to the home stream (Taft and Shapovalov, 1938; Lindsev,
et al, 1959).
Food habits of rainbow and Steelhead seem to depend as much on size and
availability of food items as on any preference on the part of the fish. In
general, fish in fresh water feed on various invertebrates, especially the
larvae and adults of dipteran insects, and crustaceans such as Gammarus, but
plankton, various insects, snails, and leeches also may be eaten. Other
fishes may be eaten by adult rainbows, and the availability of fish may be
necessary for the rainbow to reach maximum size. Steelhead in the sea are
known to feed heavily on squid and fishes (Reimers, et al, 1955; Le Brasseur,
1965; Scott and Grossman, 1973).
Because of its popularity as a sport fish, the rainbow has been widely
introduced, and many populations are maintained at levels suitable to anglers
only by continued stocking. Planted hatchery fish, especially in fast-moving
streams, have a regrettable tendency to die off in a relatively short time,
yielding but poor returns to the angler (Randle and Cramer, 1941; Burns and
Calhoun, 1966). At least some, and perhaps a major, part of the die off re-
sults from the inability of the fish to cope with the continued vigorous
swimming necessary in fast water. As a result, glycogen reserves are used up
faster than they can be replaced, blood lactate reaches very high levels (four
or more times above the level found in surviving fish), blood pH drops to 6.9
or less, and death ensues (Black, et al, 1962; Hochachka and Sinclair, 1962;
Jonas, et al, 1962; Miller and Miller, 1962).
IMPORTANCE TO MAN
It is safe to say that Salmo gairdneri is one of the most sought after
sport fishes in North America, and probably the most important of all sport
fish west of the Rockies. It takes both flies and lures, fights hard and
leaps frequently. The flesh is of high quality, both in texture and flavor.
Rainbows are raised for eating purposes in a'number of hatcheries in the U. S.,
as well as in Europe and Japan, and a few thousand cases of Steelhead are
canned annually from fish taken by commercial salmon fishermen. The economic
value of the sport fishery is not to be sneezed at. Anglers spend vast sums
for tackle, bait, lodging, travel, etc., in order to fish for rainbows and
59
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steelhead. In the Copper River (tributary to Lake Iliamna), Alaska, sport
fishermen in 1972 spent $125,552.18 to catch 3,621 rainbows, for an average
cost of $35.50 per trout; (Siedelman, et al, 1973.)
BROOK TROUT
Salvelinus fontinalis (Mitchill)
DISTINCTIVE CHARACTERS
The combination of dark green marbling on the back and dorsal fin and
the red spots with blue halos distinguish the brook trout.
DESCRIPTION
Body elongate, only slightly compressed, its greatest depth 20-28% of
total length, generally deeper in larger, sexually mature individuals. Head
longer than in most other American charrs, 22-28% of total length. Snout
more or less rounded, longer than eye diameter, 25-38% of head length. Eye
round, its diameter 15-22% of head length. Mouth large, terminal, maxilla
reaching well behind eye in adults, only to posterior edge of eye in young.
Breeding males often develop a kype. Teeth caniniform and well developed on
upper and lower jaws, head of vomer (but absent from shaft), palatines and
tongue. Gill rakers 14-22. Branchiostegals 9-13, usually one less on right
side than on left. Lateral line with 110-130 pored scales.
Fins
D. 10-14, adipose dorsal fin present. A. 9-13. PI. 11-14. P2. 8-10,
an axillary process present. Caudal fin nearly straight (hence the old popu-
lar name "squaretail") or with a shallow indentation.
Scales
Cycloid, minute, in about 230 rows in a mid-lateral series.
Color
Varies, but generally more or less as follows: Back greenish to brownish,
marked with paler vermiculations or marblings which extend onto the dorsal fin
and sometimes the caudal also. Sides lighter, marked with numerous pale spots
and some red spots, each of the latter surrounded by a blue halo. Anal, pel-
vic and pectoral fins with a white leading edge followed by a dark stripe, rest
of fins reddish. In spawning fish, especially the lower sides and fins become
red. Sea-run fish are dark green above, with silvery sides, white belly and
very pale pink spots.
Size
The largest known is a fish of 6.6 kg which was 700 mm long, taken in
the Nipigon River, Ontario, in 1916. In most areas, fish as large as 1-1.5 kg
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are considered large. Occasionally, fish over 7 kg have been reported, but
these have invariably turned out to be arctic charr.
RANGE AND ABUNDANCE
The native range of the brook trout is from Ungava Bay in northern
Quebec to New Jersey and southern Pennsylvania in the lowlands, still farther
south to northern Georgia in the Appalachian highlands; from the easternmost
tip of Newfoundland west to eastern Minnesota and Manitoba, where it reaches
north along the west coast of Hudson Bay to the Churchill River. It has been
widely introduced into the western states and provinces, as well as to other
parts of the world such as South America, New Zealand, Asia and Europe. In
Alaska, brook trout were introduced into southeastern in 1920 (MacCrimmon and
Campbell, 1969), with plantings continuing into the 1950's (Anonymous, 1953).
Unfortunately, it hybridizes readily with the native Dolly Varden and the
resultant hybrid is inferior to both the parent species (Baade, 1962). It
also has a distressing tendency to overpopulate in some situations, resulting
in large numbers of stunted fish which are of no interest to anglers.
HABITS
The brook trout spawns in the fall of the year, from late August in the
far north to as late as December in the southern part of the range, at water
temperatures between about 3° and 9°C. Maturation of sex products appears to
be stimulated by lessening day length and falling temperatures (Henderson,
1963). Artificial manipulation of day length, beginning in mid-January, in
hatchery situations has resulted in fish becoming ripe as early as the first
week of August. Fish subjected to this treatment produced smaller than normal,
but more numerous, eggs. The treatment was highly successful with two year
old fish, less so with older ones (Corson, 1955).
Spawning takes place over gravelly bottom, most often in fairly shallow
streams, but sometimes also in lakes at locations where there is upwelling of
ground water. The presence of springs seems to be essential and may be the
most important factor in the choice of a spawning site (White, 1930; Hazzard,
1932; Webster, 1962). Gravel size on spawning beds has been noted to vary
from coarse sand to stones 75-100 mm in diameter, and apparently is unimpor-
tant as long as ground water seepage is present.
Courtship begins with a male attempting to guide or drive a female towards
suitable spawning gravel. j\ ripe* receptive female makes close visual inspec-
tion of the gravel and, having chosen a spot, begins to dig the redd. She
turns on her side and, with powerful upthrusts of her tail, lifts gravel, silt,
etc., from the bottom. The smaller and lighter particles are swept away by the
stream current, but the larger ones settle at once to the bottom, building" a
mound just downstream from the pit. Repeated digging forms a depression from
100 to 150 mm (4-6 in) deep. The female tests the pit from time to time with
her anal, caudal and pelvic fins. Meanwhile, the male continues his courtship
activities, darting alongside of her and quivering, swimming over and under
her and rubbing her with his fins. He spends most of his time, however, in
driving off other males, in which the female assists.
61
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When the pit suits the female, she drops her anal fin deeply into it and
arches her tail. The male comes beside her, toth fish open their mouths and
quiver, and eggs and milt are deposited in the pit. This process takes about
one second and up to about 800 eggs may be dropped at one time (Greeley, 1932).
Due to the depth of the pit, quiet water and gentle eddies are present at the
bottom, which hold the eggs and the cloud of sperm in place or spread the
latter upstream and laterally (Smith, 1941; Needham, 1961).
After spawning, the female at once begins to cover her eggs, but the
method is very different from that used by most salmonids. She goes to the
mound of gravel which has formed at the downstream edge of the redd and begins
a sinuous movement, using the tips of her anal and caudal fins to sweep small
pebbles upstream into the nest. The eggs are quickly covered, after which
the female circles the pit, continuing to sweep in this rranner for about half
an hour. Only after the eggs are well covered does she go to the upstream end
of the nest and again begin the characteristic salmonid digging of a new redd
(Smith, 1941; Needham, 1961).
In Alaska, the brook trout may hydridize with the native Dolly Varden,
with results which are a credit to neither parent species (Baade, 1962). The
brook trout has been experimentally crossed with kokanee and with brown, rain-
bow and lake trout. Survival of offspring is virtually nil in the first three
crosses. Brook trout X lake trout crosses produce viable offspring called
"splake", but survival and fertility are lower than for either parent species
(Buss and Wright, 1956; 1958; Grossman and Buss, 1966). It is not known
whether the brook trout X Dolly Varden hybrids are fertile.
The eggs are large, 3.5-5.0 run in diameter after spawning. The number
of eggs produced is directly related to the size of the female. The average
number varies from 100 eggs for a fish of about 150 mm to 5,000 for a fish of
565 mm (Vladykov, 1956; Wydoski and Cooper, 1966).
Hatching time varies with temperature, but usually occurs in late winter.
At 5°C, 100 days are required, but normal, though slower, development occurs
at temperatures as low as 1.7°C. The upper limit appears to be about 11.7°C
(Embody, 1934; Bigelow, 1963).
The young fish remain in the gravel until the yolk sac has been absorbed.
They emerge in April or May when they are about 38 mm long. They stay on the
redd for a few days and may burrow into the gravel if frightened, but soon
disperse. Scales begin to form at a length of about 50 mrr. Survival to this
stage is good, averaging about 16.4% (3.6-42.4%) in one series of observations
(Smith, 1947).
Growth is rapid during the first summer. By the end of September, the
young may be as much as 100 mm long and weigh about 10 grams. Subsequent
growth rates vary with locality, temperature, food supply and the genetic
composition of the population. The most rapid growth takes place in May, slows
through the summer and goes on at a minimal level from September until spring
(Cooper, 1951).
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Sexual maturity may be reached as early as one year of age, but most
usually in the second year. All fish are mature at three years (Wydoski and
Cooper", 1966).
The brook trout is a relatively short-lived fish. Few wild individuals
survive more than five years, although there are reports of introduced fish
reaching 15 years of age in California (McAfee, 1966).
- Most brook trout are more or less stable residents in streams or lakes,
where they prefer temperatures below 20°C. On the east coast of North America.
general upstream movements have been observed in early spring, summer and late
fall, downstream movements in late spring and fall (White, 1941, 1942; Smith
and Saunders, 1958, 1967). Some fish, popularly known as "salters", run to
sea in the spring as stream temperatures rise. They stay at sea for up to
three months, feeding and growings but never venturing more than a few kilo-
meters from the river mouths. Their return to fresh water apparently is re-
lated to freshets in the main river and there is a definite homing tendencv
to the tributaries of origin within the drainage (White, 1941, 1942; Bigelow,
1963).
Brook trout eat a most amazing variety of food organisms, ranging from
copepods, eaten by the young, to small mammals (mice, voles and shrews) by
large adults. The most important items are insects of all sorts, both aquatic
and terrestrial. Other foods include Cladocera, spiders, worms, leeches,
crustaceans, molluscs, fishes, amphibians and mammals (Clemens, 1928; Ricker,
1930, 1932; Wadman, 1962). The most intensive feeding and best growth occur
at temperatures of about 13°C (Baldwin, 1957).
IMPORTANCE TO MAN
The brook trout is one of the most important sport fishes in North
America. -It is angled for with all types of sport gear by people from all
walks of life. In addition, it is amenable to pond culture and large numbers
are raised in private ponds for commercial sale, either for food or for stock-
ing.
LAKE TROUT
Sal ve 1 in us narnaycush_ (Walbaum)
DISTINCTIVE CHARACTERS
The lake trout is easily distinguished by its color, whitish or yellowish
spots on a dark green to grayish background; its deeply forked tail; and the
numerous (93-208) pyloric caeca.
DESCRIPTION
Body elongate, more or less terete. Depth usually 18-261 of TLS but much
deeper in some populations, especially those which feed largely on molluscs.
Head fairly long, 21-28% of TL. Snout long, 26-36% of head length. Eye
63
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rather small in adults, 12-20% of head length. Mouth terminal, large, maxilla
reaching well behind eye in adults, snout protruding beyond lower jaw when
mouth is closed. Teeth cam'niform, present on both jaws, on head but not
shaft of vomer, on palatines, tongue and basibranchials. Gill rakers 16-26.
Populations of lake trout which feed mostly on plankton tend to have more gill
rakers than those which are mainly piscivorous. Branchiostegals 10-14, usually
more on left than on right. Lateral line with 116-138 pores, slightly curved
anteriorly. Pyloric caeca numerous, 93-208, mostly 120-180.
Fins
D. 8-10. Adipose dorsal present. A. 8-10, P-j. 12-17. P£. 8-11, a small
axillary process present. Caudal deeply forked.
Seale s
Small, cycloid,
CoT_or
Back and sides usually dark green liberally sprinkled with -whitish to
yellowish (never pink or red) spots. Ground color varies from light green to
gray, brown, dark green or nearly black. Belly white. Pale spots present on
dorsal, adipose and caudal fins and usually on base of anal. Sometimes orange-
red on paired fins and on anal, especially in northern populations. Anterior
edge of paired and anal fins sometimes with a white border. At spawning time,
the males develop a dark lateral stripe and become paler on the back (Royce,
1951).
Size
The lake trout is the largest member of the genus Salvelinus. The
angling record is a fish of 29.5 kg (65 1 b) from Great Bear Lake, N.W.T.
The largest known came from Lake Athabasca and weighed 46.4 kg (102 Ibs)
(Canadian Dept. Fisheries, 1961).
RANGE AND ABUNDANCE
The Take trout is widely'distributed in northern North America and has
been introduced into other parts of the world such as Scandinavia, South
America and New Zealand. Its natural range in North America corresponds
closely with the limits of Pleistocene glaciation (Lindsey, 1964). It is
generally absent from lowland regions such as the Yukon-Kuskokwim valleys in
Alaska and the Hudson Bay-James Bay lowlands of Canada. The range extends
from the Alaska peninsula east to Nova Scotia, south to northern New York and
Pennsylvania and the Great Lakes, north to the islands of the Canadian arctic.
The species is intolerant of salt water (Boulva and Simard, 1968) and has made
its way only across narrow stretches of the marine environment. However, it
has been found on Southhampton Island (Manning, 1942), and Baffin, King William,
Victoria and Banks islands (Scott and Grossman, 1973).
64
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In the more isolated parts of its range, the lake trout is abundant, but in
many other areas, especially the Great Lakes, it has become rather scarce,
except where maintained by artificial propagation and careful management.
HABITS
Like most charrs, the lake trout spawns in the fall. The exact time varies
with latitude and temperature. Over the entire range, spawning activity occurs
as early as late August and early September in the far north, e.g., Great Bear
Lake and certain lakes in northern Alaska (Miller and Kennedy, 1948a; McCart
et al, 1972) to as late as December in the southern parts of the range (Royce,
1951).
Spawning takes place over clean, rocky bottom in depths from as little as
a meter or even less (Merriman, 1935; Martin, 1957', Rawson, 1961) to as deep
as 36 m (Eschmeyer, 1955). Virtually all lake trout spawn in lakes, but a few
populations are known to spawn in streams tributary to Lake Superior (Loftus,
1958). Conversations with residents of Kobuk, Alaska, suggest that a few lake
trout may spawn in the Kobuk River. However, this has not been verified.
Spawning takes place mostly at night, with peak activity between dusk and
9 or 10 p.m. (Royce, 1951; Martin, 1957). During the day, the fish are more
or less dispersed away from the spawning beds, but return in numbers in the
late afternoon. Males reach the spawning beds first and spend time cleaning
the rocks. This they do by twisting their tails and bodies over the bottom
and by rubbing the rocks with their snouts. This activity by a group of males
cleans an area of several tens of square meters. When the females arrive, a
few days after the males, they are courted by the males. A male butts the
side of the female, follows her, and may swim beneath her, brushing her vent
with his dorsal fin. At this time, the males become pale on their backs and
develop a dark stripe on each side and dark areas around the head (Royce, 1951;
Martin, 1957).
During and following courtship, the males attempt to spawn with the females.
One or two males approach a female, press against her sides with vents close
together and then quiver. Both sexes have their mouths open and the male's
dorsal fin is erect. On occasion, as many as seven males and three females
may engage in a mass spawning act (Royce, 1951). The spawning act lasts only
a few seconds and is probably repeated many times before a female has voided
all her eggs. Fecundity varies from a few hundred to more than 17,000 eggs
per fema.le, the number of eggs increasing with the size of the fish (Eschmeyer,
1955).
Spawning is an annual affair in southern areas, but evidence indicates
that it occurs only every other year in Great Slave Lake and only every third
year in Great Bear and some other lakes of'the high arctic (Miller and Kennedy,
1948a; Kennedy, 1954; Furniss, 1974).
The large eggs, 5 to 6 mm in diameter, fall to the bottom and lodge in
crevices and crannies among the'rocks. Incubation requires 15-21 weeks or
mores depending on temperature, and hatching normally occurs between mid-
February and late March (Eschmeyer, 1955; Martin, 1957). The newly hatched
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young are about 16 mm long-, with an exceptionally large yolk sac. Buds of the
pelvic fins and elements of the anal and caudal are already present (Fish,
1932). Growth is usually good during the summer and by September or October
the young fish are about 60-70 mm long. Subsequently, growth rates vary widely
from place to place, according to water temperature, altitude, type and amount
of food available, and the genetics of the fish. Most growth occurs during
the summer, with some northern populations growing only from June to September
(Kennedy, 1954). Overall, age I fish average about 200 mm TL; age V, 450 mm,
874 g; age X, 728 mm, 3,240 g; age XV, 964 mm, 7,631 g. The slowest growing
lake trout are those of Great Bear Lake, N.W.T., and of Itkillik and Campsite
lakes in Alaska. Ten-year-olds from Great Bear Lake average only 406 mm TL, -
age XC, 528 mm, age XX, 678 mm. A 37 year old fish was 1,046 mm TL and weighed
15.4 kg (Miller and Kennedy, 1948a). What appears to be the oldest known was
a fish of 927 mm and 12.5 kg estimated to be 42 years old taken in Chandler
Lake, Alaska, in July 1973 (Furniss, 1974). Another aged at 41 years was re-
ported from the west coast of Hudson Bay (Sprules, 1952). Fish from the two
Alaskan lakes had even slower growth rates when calculations were based on
otoliths. However, when calculations were based on scales, growth rates of
Itkillik Lake fish were quite comparable to those from Great Bear Lake (McCart,
et al5 1972). Growth of lake trout in other Alaskan lakes is generally better
than in Great Bear Lake but below the general average (Roguski and Spetz, 1968;
Van Wyhe and Peck, 1969; Furniss, 1974).
Sexual maturity is generally reached around the seventh year, but may be
achieved as early as age 5 or as late as 13. Males usually mature a year
earlier than females.
Although the lake trout can hardly be called migratory, in that whole
populations do not undertake movements in definite directions, it is definitely
a solitary wanderer. Tagged fish in Lake Superior have been recovered up to
408 km (255 mi.) from the point of release. In other areas, lake trout wander
throughout the lake, the extent of their movements apparently limited chiefly
by the size of the body of water. Smaller fish tend to move shorter distances
than do larger fish (Eschmeyer et al, 1953; DeRoche and Bond, 1957; Rawson,
1961). Dispersal begins shortly after spawning. By spring, the fish are wide
spread, and as the water warms above 10°C., they tend to go into deeper water
and to congregate below the thermocline during the summer. (Martin, 1952;
Rawson, 1961; Dryer, 1966). With the approach of fall, the fish return to
the spawning beds. They show a marked tendency to return to their natal
spawning areas, although stocked fish may utilize artificial spawning beds
(Eschmeyer, 1955; Hacker, 1957; Loftus, 1958, Martin, 1960). The newly
hatched fish move off the spawning areas and into deep water a month or so
after hatching, and apparently stay there for some time.
As with most fishes, food habits vary with age and size of the fish,
locality and the food available. Total diet includes a wide variety of such
items as zooplankton of various sorts, insects, spiders, clams, snails, many
kinds of fish, plant material such as Nostoc, sponges, amphibians, worms,
shrews, mice. The very young fish subsist almost entirely on small crustacean
plankters of various sorts. As size increases, larger organisms such as Mysis,
Pontpporeia and insect larvae become of major importance. Mature lake trout
66
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in most areas feed almost exclusively on fishes (Hildebrand and Towers, 1927;
Dimbar and Hildebrand, 1952; Hacker, 1957; Kimsey, 1960; Rawson, 1961; Martin,
1970). Coregonids appear to be particularly important in most places and to
provide superior nourishment for the'lake trout. In Lake Opeongo, Ontario, a
shift to coregonids from other kinds of fish resulted in faster growth* greater
weight, older age at maturity and larger and more numerous eggs (Martin, 1970).
Some populations of lake trout feed on plankton throughout their lives.
This may be due to unavailability of forage fish, or perhaps to a genetic
characteristic of the trout. Such fish tend to have more numerous and longer
gill rakers, and often develop accessory rakers on the medial side of the
gill arch (Martin and Sandercock, 1967). Plankton-feeding lake trout grow
more slowly, mature earlier and at smaller size, die sooner, and attain smaller
maximum size than do their fish-eating counterparts (Martin, 1966). Seasonal
changes in food habits, probably in response to availability, have been ob-
served in several areas. In Lake Superior, coregonids made up the major food
in fall and early winter, but were replaced by smelt in February and March,
with cottids, mysids and insects most common in late summer (Dryer, et al,
1965). A similar situation prevailed in certain lakes in Ontario, where again
fishes were eaten more commonly in winter (Martin, 1954).
Lake trout are not known to produce natural hybrids with other species of
Salvelinus, perhaps because of their unique spawning habits. However, artifi-
cial crossing of female lake trout and male brook trout yields viable offspring,
called "splake." These splake grow more rapidly than either parent species,
are fertile and are known to reproduce naturally (Martin and Baldwin, 1960).
Lake trout are highly susceptible to pollution, especially from such in-
secticides as DDT. This substance accumulates in the yolk and fatty tissues
and concentrations of less than 3 ppm produce mass mortalities of the alevins
following yolk-sac absorption (Burdick et al, 1964).
IMPORTANCE TO MAN
The lake trout was formerly one of the mainstays of the Great Lake fish-
eries, with landings up to nearly 7,000,000 kg in 1940 (Fiedler, 1943). How-
ever, with the entrance of the sea lamprey, Petromyzon marinus, into these
waters, and the extensive use of insecticides such as DDT, the stocks declined
drastically and annual catches now amount only to about 190,000 kg (Anonymous,
1975). It is still an important commercial species in Canada, especially in
the Northwest Territories, but even here, free of the lamprey, landings have
declined by a third in recent years (Scott and Grossman, 1973).
As a sport fish, the lake trout is well regarded. It fights hard, though
not spectacularly, and the large size attained makes it a trophy worthy of any
angler's skill. The flesh is usually of a yellowish creamy color, but may be
anything from white to orange, depending, at least in part, on diet. Regard-
less of color, lake trout is excellent as human food.
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ARCTIC CHARR
Salveljnus alpinus (Linnaeus)
DISTINCTIVE CHARACTERS
The arctic charr is distinguished by the presence of 23 to 32 gill rakers
and 35 to 75 pyloric caeca, and the presence of pink to red spots, the largest
usually larger than the pupil of the eye, on the sides and back.
DESCRIPTION
Body elongate, more or less terete, compressed posteriorly. Depth about
20% of TL, but highly variable, depending on time of year, sex, size, and
state of maturity. Head moderate, 22-25% of TL. Snout rounded, its length
usually equal to or longer than eye diameter. Eye more or less round, large,
diameter averaging about 23% of head length, but highly variable. Mouth large,
terminal, maxilla reaching nearly to posterior margin of eye in small specimens,
to well behind eye in large fish, especially in spawning males. Teeth canini-
form, in an irregular single row on each jaw and palatines. A patch of teeth
present on head of vomer. Teeth on tongue 10-24, in two rows. Gill rakers
23-32 on first arch. Branchiostegal rays 10-13. Lateral line with 123-152
pored scales, slightly decurved anteriorly. Pyloric caeca 35 to 75.
Fins
D]. 12-16 (10-12 major rays). Adipose dorsal fin present. A. 11-15
(8-11 major rays), P-j. 14-16. Pg. 9-11, axillary process present.
Scales
Small, cycloid.
Color
Highly variable, depending on location, time of year and degree of sexual
development. In general, the back is dark, usually brownish but sometimes
with a greenish cast. The sides are lighter colored, belly pale. Sides and
back are liberally sprinkled with pink to red spots. The largest spots along
the lateral line are usually larger than the pupil of the eye. Leading edges
of pectoral, pelvic arid anal fins, and sometimes the caudal, with a narrow
white margin. Fins pale in young fish, dorsal and caudal dark in adults,
but other fins light. Spawning adults, especially males, are brilliant
orange-red to bright red on the ventral side and on pectoral, pelvic and anal
fins. Young Arctic charr have about 11 dark parr marks on each side.
Size
The largest arctic charr known is one of "over 27 pounds (12.2 kg)" from
the Tree River, N.W.T. (Scott and Grossman, 1973). However, average size is
much less than this. Arctic charr in Lake Aleknagik, Alaska, run about 1 to 2
kg. Fourteen fish taken in a small gill net in Campsite Lake at the head of
68
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the Kuparuk River, Alaska, in August 1973, averaged less than 1 kg, but others,
taken on hook and line in the same lake, ran up to 5 kg. Thirty four charr
from Chandler Lake, Alaska, taken by gill net in July 1973, ranged in weight
from 569 to 2196 grams (Furniss, 1974).
RANGE AND ABUNDANCE
Distribution of the arctic charr is circumpolar in arctic and sub-arctic
regions, with isolated relict populations in cold lakes south of the normal
range. In North America, the species ranges from the Aleutian Islands, the
Alaska peninsula and Kodiak Island north to the Arctic Ocean as far north as
Discovery Harbor, Lady Franklin Bay, Ellesmere Island (82°42'N according to
GUnther, 1877, but more recently listed as 81042'N, 65°20'W in the Gazeteer
of Canada, 1971), and thus is the most northerly of all salmonids. The general
distribution is circumpolar reaching south to New England, the British Isles,
isolated locations in the Alps and Siberia, and the islands of Japan, In
most areas where it is present, the arctic charr is abundant, though this
abundance is often highly localized.
HABITS
Spawning occurs in the fall of the year, usually over gravel shoals in
lakes, but sometimes in quiet pools in streams close to a lake. In lakes of
the North Slope of the Brooks Range, Alaska, spawning may begin as early as
late August or early September, while farther south it may not take place until
November or December. Water temperatures range from about 3°C to nearly 13°C.
The male fish are territorial, guarding their areas and driving off in-
truding males with considerable vigor. However, as soon as the females begin
to show spawning behavior, the males pair with them and lose interest in their
territories. The female invades the male's territory and at first may be
attacked by the male. She persists, however, and makes visual examination of
the bottom. With head and tail bent downwards and eyes directed towards the
bottom, she swims slowly about until she finds a suitable spot. During this
period, she may attack nearby fish of either sex. Hhen a suitable spot has
been found, the female begins to dig the redd. This may start with powerful
swimming movements, pushing water backwards and carrying away sand and loose
debris. More often, however, digging begins in the typical salmonid fashion.
The female turns on her side and gives a few strong, upward flaps of her tail,
thus displacing the lighter particles, sand and small gravel. After each spell
of digging, the female swims forward, circles around and returns to about the
same spot to dig again. Digging occurs about once a minute. Depending largely
on the size of the bottom particles, the completed redd varies from a fairly
deep pit to a clean spot on large stones.
While the female digs the redd, the male courts her by circling around
her, then gliding along her side and quivering. In intensive courtship, the
verticle and pelvic fins are erected and the male opens his mouth. During
courtship, the male develops a dark band along the side and across the top of
the head, while the back becomes pale.
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When the redd is completed,- the female tests it with her erected anal
fin, bends head and tail upward, opens her mouth and quivers. The male joins
her, assuming a like-position, and eggs and sperm are ejected. The fish then
swim forward out of the nest, often still ejecting sex products. This act may
be repeated up to five times before the female begins to cover the eggs. She
does this by swimming about the redd, sweeping the gravel with her anal and
caudal fins. This sweeps the eggs into the nooks and crannies of the gravel.
The female then digs at the edge of the pit, covering the eggs and beginning
the next redd.
Spawning apparently takes place at almost any time of the day or night,
although most apparently occurs during the day. Males often mate with more
than one female, taking the second mate after the first has exhausted her
eggs. Occasionally a female may mate successively with two or more males
(Fabricius, 1953; Fabricius and Gustafson, 1954).
Several days are usually required for a female to deposit all her eggs.
Depending upon the size of the fish, a female may produce as few as a couple
of hundred to as many as four or five thousand. The eggs are fairly large,
3.5 to 5.5 mm in diameter, yellowish to orange in color. Time of development
seems to vary widely according to conditions and stock. Under hatchery con-
ditions at 4,4°C, hatching may occur in 60-70 days (Bigelow, 1963), while in
the northwestern part of the Hudson Bay area "eggs are still visible at
break-up in the following spring, probably hatching soon after open water
appears." (Sprules, 1952). The situation in Alaskan waters is not known.
The young are about 17 mm long at hatching. Subsequent growth rates
depend largely on temperature and the abundance of food. In Swedish hatcheries
average lengths of 97 mm at the end of the first year, 123 mm at the end of
the second, and 137-216 mm at the end of the third have been reported (Bigelow,
1963). Growth under wild conditions is much slower. In the eastern Canadian
arctic, 5 year old fish averaged from 130 to 382 mm fork length; 10-year-olds
from 363 to 584 mm; and 15-year-olds from 553 to 671 (Sprules, 1952; Grainger,
1953; Andrews and Lear, 1956). In Chandler Lake, in Alaska's Brooks Range,
10 year old fish averaged 424 mm FL, 14-year-olds 537 mm (Furniss, 1974). The
arctic charr is thus a very slow growing fish. It is known to achieve an age
of at least 24 years (Grainger, 1953).
Some populations of arctic charr, particularly in Europe, Siberia and
eastern Canada, are anadromous, running to sea shortly after break-up in the
spring and returning to fresh water in late summer and early fall. Other'
populations, especially lake-dwelling fish, remain in fresh water all their
lives and do not migrate. In Alaska, it appears that all arctic charr are of
the fresh water, lake-dwelling type. The species has been reported as being
anadromous in most rivers from Bristol Bay to Barter Island. However, exami-
nation of numerous specimens and published data shows that these anadromous
fish are the northern form of the Dolly Varden, SalyeTinus. malma (Morrow, un-
published) .
The food of the arctic charr is varied, mostly according to what is
available. In Chandler Lake, Alaska, insects, especially Diptera, were most
70
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important. Arctic charr feed heavily on young sockeye salmon in the Karluk
and Wood River lakes systems in Alaska (DeLacy and Morton, 1943; Thompson,
1959; personal observations). Other foods reported from various parts of the
world include fishes, crustaceans, molluscs, nereid worms and their own young.
IMPORTANCE TO MAN
The arctic charr is an important sport fish in the Canadian Arctic and
in Scandinavia, as well as in the Wood River lakes of Alaska. Canadian
eskimos rely heavily on it for food in some regions. It is fished commercially
in various locations in northern Canada, Greenland, the Scandinavian countries
and Siberia. The flesh is of excellent quality and commands a good price.
Red-meated fish are preferred over those with pink or white flesh, and are
priced accordingly. Frozen arctic charr are shipped to gourmet restaurants
in most of the larger cities of the world. Canned charr is another high
priced delicacy. However, the arctic charr is slow-growing and there is
some doubt as to whether many populations can withstand heavy exploitation.
There are a number of examples of serious depletion in only a couple of
years (Andrews and Lear, 1956).
DOLLY VARDEN
Salyelinus malma (Walbaum)
DISTINCTIVE CHARACTERS
The Dolly Varden is identified by the lack of worm-like marbling on the
back, the presence of pink or red spots on the sides, 11-26 gill rakers, and
13-35 (rarely up to 39) pyloric caeca.
DESCRIPTION
Body elongate, rounded. Depth about 20% of FL. Head moderate to fairly
long, 15-28% of FL, longest in breeding males. Snout rounded, about equal to
eye in small fish, longer in adults, especially breeding males. Eye round or
nearly so, about 20% of head length. Mouth large, terminal, upper jaw reaching
about to vertical through hind margin of eye in small fish, well behind it in
large ones. Well developed caniniform teeth present on both jaws, head (but
not shaft) of vomer, palatines and tongue. Zero-44 small teeth on base of
tongue (basibranchials). Gill rakers 11-26, average numbers 17-19 in the
southern form, 21-23 in the northern. Branchiostegals 10-15, usually one
more on left than on right. Lateral line with 105-142 pored scales. Pyloric
caeca 13-35, rarely to 39.
Fins
D, 13-16 (10-12 major rays). Adipose dorsal present. A, 11-15 (9-12
major rays). P]. 14-16. ?2- 8-11, usually 9, a small axillary appendage
present. Caudal fin emarginate.
71
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Scales
Small, cycloid.
Color
Extremely variable according to locality, size of fish and degree of
sexual maturity. Very young parr are usually brownish with 8-11 (12) dark
parr marks, the sides and back showing small red spots, the fins pale to
dusky. Fish in salt water or just entering fresh water may be dark above,
with silvery sides sprinkled with pale or pink spots. Breeding fish become
dark greenish black above and on the upper sides. Males in breeding condition
turn bright orange to red on the lower sides and belly. Pectoral, pelvic and
anal fins usually have white or creamy leading edges followed by a black or
red line. Dorsal and caudal fins usually dusky or brownish. Largest spots
on sides usually smaller than pupil of eye.
Size
The largest Dolly Varden known was 103 cm (40.5 inches) long and weighed
14.5 kg (32 Ibs).
RANGE AND ABUNDANCE
Definition of the range of the Dolly Varden is difficult because it has,
in the past, been confused with at least two other species, the arctic charr,
Salvelinus a1 pinus, and the bull trout, S^ confluentus (McPhail, 1961; Caven-
der, pers. cornm.). The range in North America, as presently known, extends
from the arctic coast of Alaska and probably of western Canada as well, south
to northern California, eastward into Nevada, Idaho, Montana and Alberta. At
least some of the eastward records will undoubtedly turn out to be $_. con-
fluentus. The Dolly Varden is widely distributed throughout Alaska, where
both anadromous and apparently non-anadromous populations are to be found in
clearwater streams. The presence of northern type Dolly Varden in the lower
Susitna drainage and of southern type in the upper Tanana has undoubtedly
been brought about by headwaters exchange between the Nenana and Susitna in
the one case, the Slana and the Little Tok in the other (Morrow, unpublished).
On the western side of the Pacific, Dolly Varden are found from the
Cnukhotsk Peninsula south to Japan and Korea.
Wherever found, the Dolly Varden is usually abundant, although, because
of its migration patterns, the abundance may be seasonal.
HABITS
The life history of the Dolly Varden is fairly complex and the account
is not simplified by the fact that different populations seem to behave dif-
ferently. The vast majority of Dolly Varden are anadromous, but some popula-
tions, especially in interior Alaska, apparently spend their entire lives in
freshwater streams.
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Spawning takes place in the fall, from late August to the latter part of
November (Armstrong, 1967; Yoshihara, 1973). Spawning behavior is similar to
that of other members of the genus. The male courts the female by swimming
around and beside her, pressing against her and quivering. The female selects
the nest site on clean gravel, the stones being from 6 to 50 mm in diameter.
The redd is located in fairly strong current, usually near the center of the
stream in water 0,3 m or more deep. Preferred water temperature in south-
eastern Alaska seems to be around 5.5°-6.5°C (42°-44°F). She digs the redd in
typical salmonid fashion, turning on her side and giving several strong upward
flips of her tail, thus dislodging the gravel and debris which are carried
downstream. While the female is preparing the redd, the male continues to
court and drives off intruding males. When the redd is finished, both fish
drop into it, press against each other with arched backs and open mouths,
quiver and expel! the sex products. This procedure lasts about 5 seconds,
and may be repeated several times before the female begins to cover the eggs.
Covering is accomplished by the female who swims along the edge of the redd,
sweeping small pebbles and other particles into it with her tail and anal fin.
Later she may dig again in typical salmonid fashion and further cover the eggs
while preparing a new nest (Needham and Vaughan, 1952; Blackett, 1968). Sexual
dimorphism is apparent at this time. Females develop an extended and swollen
ovipositor. Males, at least of anadromous populations, develop a pronounced
kype, which begins to return to normal along in November.
The mature eggs are 4.5-6.0 mm in diameter, orange-red and demersal. Egg
number per female varies with stock and location. Southern Alaskan Dolly Varden
produce from 739 to 5,968 eggs. Fish in streams of the Arctic coast may sur-
pass 6,000 eggs (Nagata, 1967; Blackett, 1968; Yoshihara, 1973). Development
to hatching requires about 4 months (129-136 days) at 8.5°C (Blackett, 1968),
but as long as 7 months along Alaska's north slope {Yoshihara, 1973). The
newly hatched young, about 15-20 mm long, remain in the gravel until the yolk
sac has been absorbed,.a period of 60-70 days. On emergence they are about
25 mm long.
The young begin to feed actively as soon as they emerge from the redd.
The major foods are various winged insects and the larvae of mayflies and
midges, but other insects and various small crustaceans are also consumed
(Karzanovskii, 1962). These newly emerged fry tend to stay on the bottom in
pools or eddies, relatively inactive except when feeding. Growth rates vary
tremendously from area to area and population to population. Thus, in Eva
Creek, in southeastern Alaska, young fish reach only about 63 mm at 1 year,
78 mm at 2, and 100 mm at 3 (Heiser, 1966; Blackett, 1968). By contrast,
young fish at nearby Hood Bay, on Admiralty Island, and on Prince of Wales
Island, achieve 90 mm at 1 year, 154 mm to 182 at 2, and over 200 mm at 3
years (Reed, 1967; Armstrong, 1967). Dolly Varden in rivers of the north
slope grow at intermediate rates (Roguski and Komarek, 1971; Yoshihara, 1973).
The young Dolly Varden usually spend three or four years in the creek before
going to sea. These young fish going to sea are typical smelts. The parr
marks have disappeared and the fish are silvery.
Subsequently, greatest annual growth takes place between May and September
of each year, while the fish are in salt water. Some of these fish may double
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their weight in this short time (Revet, 1962; Heiser, 1966). The Dolly Varden
of southeastern Alaska appear *o grow more slowly than those of the Arctic
rivers. At six years of age, arctic fish run about 50 mm longer than those
from southeastern areas, and at 10 years the difference is nearly 80 mm.
Typical lengths at various ages for northern fish are 3 years, 164 mm; 5 years,
329 nrn; 8 years, 470 run; 10 years, 517 mm (Reed, 1967b; Armstrong, 1967;
Rogushi and Komarek, 1971; Kogl, 1971; Yoshihara, 1973). Whether this differ-
ence reflects food conditions or a genetic difference between the northern and
southern forms of Dolly Varden is not known.
Sexual maturity is achieved at 4 to 6 years in the southern form, but not
until 7 to 9 in the northern (Blackett, 1968; Yoshihara, 1973).
The migration pattern of the Dolly Varden is rather complex. In south-
eastern Alaska, a migration of smelts and adults out of the over-wintering
lakes begins in early spring and continues into July. After a stay in the sea
of as little as a couple of weeks or as long as seven months, they return to
spawn in the stream and over-winter in the lake. Thus, some fish are still
moving out while others are already returning to freshwater. Some may enter
non-lake streams to spawn, then leave and enter a lake-stream system to over-
winter. Homing to the over-wintering lake is on the order of 40-501, with the
remaining over-wintering fish of unknown origin (Revet, 1962; Armstrong, 1965a).
Migrations of the northern form are not at all well known. Such data as
are available indicate that the outmigration of smelts, mostly 3 or 4 year olds,
occurs immediately after break-up. In arctic rivers, adults probably over-
winter either in deep pools or springs in the river, or in the estuary. It is
possible that at least some may spend the winter in the sea. The return mi-
gration back into the streams begins in August, with peak movement of spawners
in the last week of that month, but continuing at least well into September.
Non-spawners tend to enter freshwater a few weeks later than spawners. The
northern Dolly Vardens apparently spawn only about every third year in these
arctic rivers, whereas in southeastern Alaska spawning is probably an annual
affair (Roguski and Komarek, 1971; Kogl, 1971; Yoshihara, 1973; Furniss, 1974).
The Dolly Varden is known to hybridize with the brook trout in some lakes
of southeastern Alaska where the latter species has been introduced. The hy-
brids are said to be inferior to both parent species.
Movement in salt water is usually more or less coastwise in-southeastern
Alaska. However, this impression may well be due to the arrangement of islands
and channels in the area. Within these confines, however, tagged Dolly Varden
have been recovered as much as 115 km (72 miles) from the point of release
(Armstrong, 1965a). Northern Dolly Varden of unknown origin have been taken
in the Bering Sea at least as far west as 170° W (Hanavan and Tanonaka, 1959).
Dolly Varden in southeastern Alaska reach 10 to 12 years of age, although
there is .a rapid reduction in numbers after age 5, perhaps because of post-
spawning mortality (Heiser, 1966). Northern fish, especially those along the
Arctic coast, tend to live longer. The majority of spawners are 8 or 9 years
old, and one of 18 years has been recorded (Yoshihara, 1973). A hatchery fish
in California was reported to have reached its 20th year (Shebley, 1931).
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The diet of the Dolly Varden varies with the size and age of the fish,
location, time of year and local availability of food items. As already noted,
the young fry feed chiefly on insects and small Crustacea. Stream food of both
rearing fish and adults also consists largely of various insects, spiders and
annelids, as well as snails, clams, fish eggs and various small fishes. In
estuaries and at sea, the food of the adults consists largely of small fishes
and invertebrates, including smelts, herring, young salmon, sand lance,
greenlings, sculpins, flounder larvae, cods, as well as amphipods, decapods,
mysids, euphausiids, brachiopods, polychaetes, megalopids, and isopods. In
the high Arctic, mysids seem to be the most important food, with fish second
(Townsend, 1942; Roos, 1959; Lagler and Wright, 1962; Narver and Dahlberg,
1965; Armstrong, 1965b, 1967, 1971b; Yoshihara, 1973).
IMPORTANCE TO MAN
The Dolly Varden has long been reviled by salmon fishermen, canners and
others (some of whom should have known better) as an eater of salmon spawn
and a most serious predator on young salmon, especially sockeye (see, for
example, Ohmer 1929a, b; Rounsefell, 1958). Despite the fact that a few bold
biologists began to refute this shortly after the turn of the century (Chamber-
lain, 1907), it was not until the 1940's that it became apparent that the
Dolly Varden did not deserve its bad reputation and that, at least in some
areas, the real culprit was the closely related arctic charr (DeLacy and
Morton, 1943). Dolly Varden do eat salmon eggs and young salmon on occasion.
However, the eggs are "drifters", washed out of the redd or dug up by later
spawners, which would not survive anyhow (Chamberlain, 1907; Reed, 1967a;
Armstrong, 1967). With respect to young salmon, especially sockeye, other
predators appear to be much more important than Dolly.Varden. Removal of more
than 20,000 predaceous fishes from Cultus Lake, B.C., led to a 1 1/3 times
increase in sockeye survival (Foerster and Ricker, 1941). However, the Dolly
Varden made up less than 5% of the predators, so could scarcely have accounted
for much of the increase. In other places, such species as coho salmon, rain-
bow and cutthroat trout, squawfish and sculpins have been shown to be much
more serious predators on salmons than the Dolly Varden (Chamberlain, 1907;
Pritchard, 1936b; Ricker, 1941; Armstrong, 1965b; Reed, 1967a). Dolly Varden
certainly will eat young salmon when they are abundant and concentrated (Bower
and Fassett, 1914; Pritchard, 1936b; Ricker, 1941; Hartman, et al, 1962^ Arm-
strong, T965b; Thompason and Tufts, 1967; McCart, 1967), but in most places
the damage has been grossly exaggerated.
For many years a bounty was paid in Alaska for Dolly Varden tails in the
hope of decreasing the Dolly Varden population and thus enhancing the survival
of young salmon. While this practice may have served to enhance the income
of native fishermen, it did little, if anything, towards accomplishing its
stated purpose. The bounty was finally eliminated in 1941.
As a sport fish, the Dolly Varden will take many artificial lures,
especially shiny spoons and spinners, and also flies. They fight gamely,
though not spectacularly, seldom jumping, and are not as strong as rainbow
trout of comparable size. However, Dollies generally are quite a satisfactory
sport fish.
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For many years, Dolly Varden were taken incidental to the salmon fisher-
ies of southeastern and western Alaska and in some years as much as 91,528 kg
(over 201,000 Ibs) were marketed fresh, frozen, canned or pickled (Bower,
1919). In recent years, however, landings have been mim'seule, only two or
three thousand kilograms (5,000 Ibs) a year (Anonymous, 1975). The Dolly
Varden is an important part of the subsistence fisherey in many areas of
northern Alaska.
ANGAYUKAKSURAK CHARR
Sal veljjnus anaktuvukensis Morrow
DISTINCTIVE CHARACTERS
The angayukaksurak is distinguished from its closest relative, the Dolly
Varden, by the more numerous rays in the dorsal, anal, pectoral and pelvic
fins; and in having more lateral line pores and parr marks. The angayukak-
surak is easily separated from the arctic charr by the fewer gill rakers and
pyloric caeca.
DESCRIPTION
Body elongate, sub-cylindrical anteriorly, somewhat compressed posteriorly.
Depth 18-24% of SL. Head moderately large, its length 20-29% of SL. Snout
longer than eye diameter, 27.6% (24-31%) of head length. Eyes nearly round,
the horizontal axis a little longer than the vertical, 191 (16-23%) of head.
Nostrils double, the anterior one with a flap around its posterior margin.
Mouth large, upper jaw reaching nearly to posterior margin of eye in young,
well behind eye in adults. Teeth small, caniniform, in a single, irregular
row on each jaw and palatine. Vomerine teeth in a V-shaped patch on head of
vomer, ends of patch usually separated by a wide edentate space from anterior
palatine teeth. Tongue teeth 4-13, in two widely separated rows. Gill rakers
about 21 (18-23). Branchiostegals 9-12 on left, 9-11 on right, usually one
more on left than on right. Lateral line straight, with 135 (127-152) pores.
Pyloric caeca 28 (24-32),
Fins
D. 14-17, adipose dorsal present. A. 13-15. PI. 13-15. Pg. 10, rarely
9, small axillary process present. Caudal only slightly indented, 19 principal
rays.
Scales
Small, cycloid.
Color
Deep, velvety black on back and sides, paler below. Small, fiery red
spots on back and sides, spots smallest and most numerous on back. Largest
spots smaller than pupil of eye. Some or all spots may be surrounded by a
76"
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blue to pale halo. Leading edges of dorsal, anal and pelvic fins cream to
yellow. All rayed fins dark, without spots or marks. Adipose fin paler, with
a dark posterior margin.
Size
The angayukaksurak does not reach large size. The largest known is the
holotype, only 278.5 mm FL.
RANGE AND ABUNDANCE
Known only from headwaters situations in the Brooks Range, Alaska. Here
it is distributed somewhat discontinuously from Howard Pass eastward to the
Romanzoff Mountains, specifically: Western part of Howard Pass; Killik River;
Ikiakpuk (Ekokpuk) Creek, Giant Creek, Contact Creek, Loon Lake Creek, all in
the John River drainage; Tolugak Lake, Kanayut (Shainin) Lake, Willow Creek in
the Anaktuvuk River drainage; Chandler River; North Fork, Koyukuk River; Hula
Hula River; Aichilik River (Walters, 1955; S. Paneak, pers. comm.; personal
obvervations).
HABITS
Virtually nothing is known of the biology of the angayukaksurak. It is
found only in small headwater streams. In the winter, it congregates in
springs, where there is a constant flow of water, and is reported to spawn in
the springtime (Simon Paneak, Anaktuvuk Pass, pers. comm.). This report is
borne out by the appearance of the gonads of the topotype series, which was
taken in December. The angayukaksurak is extremely slow-growing. Five speci-
mens taken in October, 1968 were 230 to 275 mm FL and were 6 to 9 years old.
Dolly Varden of the same age from the same general area are about twice that
size (Winslow, 1969).
IMPORTANCE TO MAN
At present, the angayukaksurak is important only as a rare species, a
zoological curiosity. In the old days, however, it was important to the in-
land Eskimos as an emergency source of winter food (S. Paneak, pers. comm.).
PINK SALMON
Oncprhynchus gorbuscha (Walbaum)
DISTINCTIVE CHARACTERS
The pink salmon is distinguished from its relatives by the presence of
large, black spots on the back and on both lobes of the caudal fin. The young
have no parr marks.
DESCRIPTION
Body elongate, fusiform, slightly compressed. Depth about 221 of FL in
females and non-breeding males. Breeding males develop a great hump behind
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the head, so that body depth is about 1/3 of FL. Head moderate, about 20% of
FL, but much longer in breeding males, which develop an elongate, hooked snout.
Snout about 30% of head in females, nearly 50% in breeding males. Eyes rather
small, round, in advance of posterior end of maxilla. Mouth terminal, slightly
oblique, breeding males with a pronounced kype. Teeth caniniform, well developed
on both jaws, head and shaft of vomer, on palatines and tongue. No teeth on
basibranchialIs. Gill rakers moderate, 24-35 on first arch. Branchiostegals
9-15 on each side. Lateral line with 147-205 pored scales. Pyloric caeca 95-
224.
Fins
D. 10-15, a large adipose dorsal fin present. A. 13-19. P]. 14-17.
Pg. 9-11, axillary process present.
Scales
Cycloid, small.
Color
Pink salmon in the sea are steely blue to blue-green on the back, silver
on the sides and white on the belly. Large, oval black spots present on back,
adipose fin and on both lobes of caudal fin. Breeding males become dark on
the back, the sides red with brownish-greenish blotches; females similarly but
less distinctly colored.
Size
The pink is the smallest of the Pacific salmons. The greatest size
achieved is about 750 mm (30 in) and 6.4 kg (14 Ibs), but most run from 1-3 kg,
with the average a bit less than 2 kg (about 4 Ibs).
RANGE AND ABUNDANCE
The pink salmon is found from La Jolla, California (Hiller and Lea, 1972)
north to the Arctic Ocean. The range extends eastward to Kidluit Bay in the
HcKenzie River delta, westward across northern Siberia to the Lena River,
south along the Asian coast to Korea and Kyushu, Japan. It is most abundant
in the central parts of the north-south range.
Pink salmon have been introduced along the Atlantic coast from Maryland
north to Newfoundland. The northern introduction apparently survived for some
years, but seems to have disappeared recently (Hart, 1973). An accidental
introduction into a tributary of Thunder Bay, Lake Superior, in 1956, seems to
have survived. Introductions have been made into several European areas and
one stock is said to have survived (Scott and Crossman, 1973).
HABITS
Adult pink salmon move into streams from the sea any time from June to
late September, depending largely on location. As a rule they do not go far
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upstream and in many areas spawning occurs in the lower tidal areas. Even
though the eggs and alevins are unable to adapt to salt water (salinity
31.8o/oo), they can withstand exposure to it for a day or more without serious
harm (Weisbart, 1968). This is, of course, much longer than the periods of
high tide during which intertidal redds are exposed to sea water. However,
some-groups may ascend rivers such as the Kushokwim and Yukon for as much as
160 km (100 mi), while in Siberia pink salmon ascend the Amur River as much as
600 km (375 mi). Spawning takes place in mid-July in the lower Yukon, but
generally in late August to October in areas to the south.
Spawning behavior is typically salmonid. The female prepares the redd by
turning on her side, pressing her caudal fin against the bottom and giving
several strong flaps with her tail. This dislodges silt, debris and small
gravel which are carried downstream by the current. Repeated digging results
in a pit somewhat longer than the female and perhaps as deep as the depth of
her body, but may be as long as 915 mm (3 ft) and up to 457 mm (1.5 ft) deep.
The male takes no part in digging the redd, but spends most of his time driving
off intruding males. When the redd is finished, nale and female drop into it,
erect their fins, open their mouths, vibrate and release eggs and sperm. Pair-
ing is usually on a 1:1 basis, but up to six males have been seen to spawn with
a single female (Wickett, 1959). All pink salmon die after spawning.
The large (about 6 mm diameter) orange-red eggs fall into the interstices
of the gravel and are fertilized by the milt. The eggs are covered with gravel
as the female subsequently digs a new nest at the upstream edge of the previous
one. Each female may produce as few as 800 or as many as 2,000 eggs. In gen-
eral, larger fish have more eggs, but fish from small runs are said to be more
fecund than those of the same size from large populations (Nikolskii, 1952).
Development requires 61 to about 130 days, depending largely on temperature.
The young hatch out in late December through February and remain in the gravel
until spring. Mortality within the redd occurs chiefly before the eyed stage,
and survival from spawning to emergence from the gravel shows a strong inverse
correlation with the number of adult spawners (Hanavan, 1954; Hunter, 1959).
The alevins have a large yolk sac. The yolk sustains them while they remain
buried in the gravel through the winter. Usually in April or early May the
yolk is fully absorbed and the young fry, now 30-45 mm long, wriggle up through
the gravel. Almost immediately afterward, they begin to move downstream to the
sea. Most of the fry swim downstream at night, staying at or close to the sur-
face of the water. Migrations of up to 16 km (10 miles) may be accomplished in
a single night. In short streams or from spawning areas near the river mouth,
the migration to the sea may be accomplished in a single night. If not, the
fry hide in the gravel during the day and resume their downstream movement the
next night (Neave, 1955). Sometimes the fry become daylight-adapted, school
and no longer hide during the day. This is particularly apparent in fry which
have been migrating for several days (Hoar, 1956). In southeastern Alaska,
this outmigration begins in March, peaks in mid-May, and ends by mid-June. The
young fish are about 30 mm long at this time. They stay in the estuary for
about a month, growing to a length of about 40 mm, then follow the salinity
gradient within the estuary (Thorsteinson, 1962; Mclnerney, 1964), and move
off, generally staying fairly close to shore. Larger fry tend to migrate first,
so schools farthest from the river mouth tend to be made up of larger indivi-
duals than schools closer to the river. Within a school, the larger fry are
79
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usually found on the offshore side of the school. When the fry have reached
a length of 60-80 nwi (2.4-3.2 in), they move offshore (Le Brasseur and Parker,
1964).
The life-cycle normally takes two years, though rare three-year fish
have been found (Turner and Bilton, 1968). It is probable that these fish are
sterile. Mortality during early sea life is fairly high, 2-4% per day for the
first 40 days. In the subsequent 410 days, this drops off, resulting in an
average sea mortality of 0.4-0.81 per day. These rates are highly variable,
however, with sea survival computed at about 2-22% and probably averaging
about 5% (Neave, 1953; Parker, 1966).
During their oceanic life, pink salmon from the southern part of the
range tend to remain in a broad band of ocean more or less parallel to the
coast. Alaskan pinks, on the other hand, may be found in most of the north-
east Pacific, from Bering Strait southwestward to about Kiska Island in the
Aleutians, thence more or less southeasterly to the California coast (Hart,
1973).
In the sea, the young feed chiefly on copepods and Larvacea (QikDpleura).
As the fish grow, there is a shift towards amphipods, euphausiids, and fishes.
Other items include ostracods, decapod larvae, cirripeds and tunicates (Levani-
dov and Levanidova, 1957; Le Brasseur, 19665; Manzer, 1969).
After spending about 18 months in the sea, the adults return to spawn in
their natal stream. The pink salmon is less certain in its homing than the
other Pacific salmons, and there is a certain degree of wandering. Pink salmon
have been found spawning in streams as much as 640 km (400 miles) from the
natal stream.
In general, the upstream run seems to be triggered by high water. A
significant correlation has been found between the number of fish entering a
stream each day, the daily water level in the stream and the daily rainfall in
the area. But if water levels are too high and the current too strong, the
fish stay in eddies and wait for a drop to suitable current strength (Pritchard,
1936a). (
The males are usually larger than the females and tend to precede the
females in entering fresh water. Early running females tend to be larger than
late arrivals (Skud, 1955). Due to the two-year life cycle, the runs in odd-
numbered and even-numbered years are genetically separated, often with observ-
able, though minor, morphological differences. Thus, in British Columbia, the
adults of the odd-year cycle are consistently larger than those of the even-
year runs (Godfrey, 1959).
The pink salmon occasionally hybridizes in nature with the chum. Artifi-
cially produced hybrids show characteristics of both parent species rather than
being intermediate between the two. The hybrids exhibit the size of the chum,
but have a short (two or three year) life cycle. Most of the hybrids returned
as two-year-olds, a group in which males outnumbered females by as much as
4.2:1. In those returning as three-year-olds, the sex ratio was reversed.
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The pink-chum hybrid is fertile,vbut data are not available to indicate
whether or not self-sustaining populations could be established (Simon and
Noble, 1968).
IMPORTANCE TO MAN
Until World War I, the pink salmon was of negligible importance in North
America, either for subsistence or commercial use. Demands for food during
the war led to rapid growth of the fishery. The pink salmon is the most
numerous of the Pacific salmons and the annual pack in North America runs into
several millions to tens of millions of kilograms annually. The U. S. catch
in recent years has ranged from a low of 23,509,545 kg (51,721,000 Ibs) in
1967 to a high of 53,528,181 kg (117,762,000 Ibs) in 1970. Alaska's share of
these catches has run between 88.1% and 99.96%, but the per pound price paid
to Alaskan fishermen is usually somewhat less than the U. S. average. The
pink salmon is considered only slightly better than the chum and is priced
accordingly. Thus, while pinks have, since 1967, made up from 23.9-41.9% of
the total pack, they have accounted for only 12.9-26.81 of the total value.
Virtually all the catch is canned.
About twenty years ago it was discovered that pink salmon in the sea will
take a trolled lure. Since that time a small sport fishery for them has
developed (Scott and Grossman, 1973).
CHINOOK SALMON
Qncorhynchus tshawytscha (Walbaum)
DISTINCTIVE CHARACTERS
The small black spots on the back and on both upper and lower lobes of
the caudal fin, and the black gums of the lower jaw serve to distinguish the
Chinook salmon.
DESCRIPTION
Body elongate, fusiform, laterally compressed in adults. Depth about 25%
of fork length. Head moderate, its length about 20% of fork length, longer in
breeding males. Snout blunt, less than 33% of head in females and immature
males, but elongate and turned down in breeding males. Eye small, nearly round,
Mouth terminal, slightly oblique, upper jaw reaching well behind eye. In
breeding males, both jaws elongate and curve anteriorly, forming a kype. Teeth
canine-like, present in both jaws, on head and shaft of vomer, on tongue and on
palatines; anterior jaw teeth notably enlarged in breeding males. Gill rakers
16-30, wide spaced and rough. Branchiostegals 13-19. Lateral line with 130-
165 pored scales. Pyloric caeca numerous, 90-240, usually about 140-180.
Fins
D. 10-14, adipose dorsal fin present. A. 13-19. ?]. 14-17. PZ- 10-11,
axillary process present. Caudal emarginate.
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Seales
Cycloid, small.
Color
Fish in the sea are dark greenish to blue-black on top of head and back,
the lower sides and belly silvery to white. Numerous small, dark spots present
along back and upper sides and on both lobes of caudal fin. Gum line of lower
jaw black. In fresh water, with the approach of breeding condition, the fish
change to olive brown, red or purplish, the color change more marked in males
than in females.
Size
This is the largest of the Pacific salmons. The largest officially re-
corded came from Petersburg, Alaska, and weighed 57.3 kg (126 Ibs), but another
has been reported at 61.4 kg (135 Ibs). In most areas, however, 90-104 cm
(35-41 in) and 12-18 kg (26-40 Ibs) would be the maximum size range. The
average Cook Inlet chinook is about 750 mm long for males, 855 mm for females,
and weighs 7-9 kg (Yancey and Thorsteinson, 1963).
RANGE AND ABUNDANCE
Spawning populations of chinook salmon are to be found a-11 the way from
the Ventura River in southern California north to Point Hope, Alaska. It is
rare in streams south of San Francisco Bay, but has been taken in the sea, as
strays, as far south as La Jolla (Hubbs, 1946). At the northern end of the
range, strays have been reported from the Coppermine River emptying into the
Arctic Ocean at about 115°W. The marine adults range all across the northern
Pacific Ocean and through the Bering Sea. Along the Asian coast, chinooks are
known from the Anadyr River of northern Siberia south to Hokkaido in northern
Japan (Schmidt, 1950). Asian populations are much smaller than those of North
America (Nikolskii-, 1961). Within its North American range, the chinook is
most abundant from the Sacramento River to the Yukon River. However, because
of its anadromous habits, this abundance is highly seasonal.
Chinook have been introduced into many areas outside their normal range,
including the Great Lakes, eastern United States and Canada, Central and South
America, Hawaii, northern Europe, Australia, Tasmania and New Zealand. Some
of the Great Lakes transplants seemed to thrive at first, but sooner or later
died out. Of all the introductions attempted, only those to New Zealand have
resulted in self-sustaining populations (Davidson and Hutchinson, 1938).
HABITS
Adult chinook salmon return from the sea and move into freshwater streams
to breed. The time of arrival at the main river mouth varies with geographic
location and stock. In many streams in the southern part of the range (e.g.,
Columbia River, Fraser River) there are fairly well-defined early and late
(spring and fall) runs, while farther north (e.g., Yukon River) there is only
a single run, .although it may be spread over a period of several months. The
chinooks enter the Yukon 'in June and have reached the Canadian border by
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mid-to-late July. In Cook Inlet, chinooks may appear in the Susitna River as
early as May, but most appear in late June and July (Yancey and Thorsteinson,
1963). As a rule, the fish entering fresh water earliest are those which will
travel farthest upstream. Spawning takes place in July to early September in
the Yukon drainage, but the season may continue into November or even December
in the southern part of the range (Briggs, 1953; Scott and Grossman, 1973).
Chinook utilize remarkably few streams for major spawning areas. In British
Columbia, 50% of the spawning occurs in only 14 streams. In the United States,
some 380 streams from California to Alaska are utilized, although this figure
does not include the numerous tributaries of the Yukon and Kuskokwim rivers
(Aro and Shepard, 1967; Atkinson et al, 1967).
On reaching the spawning area in the natal stream, the female selects the
spot for her nest. She turns on her side and gives several powerful thrusts,
up and down, with her tail. The gravel, sand and debris thus loosened from
the bottom are carried downstream by the current. Repeated digging at the
same spot at intervals of five to ten minutes produces a pit, the redd, which
may be more than 3.5 m long and 35 cm deep, although usually it is only about
1.3 m long. The female drives off other females during the period of nest
digging, but pays little, if any, attention to males. She is usually ac-
companied by a dominant and one or more subordinate males. The dominant male,
in particular, drives off intruding males, although subordinates may undertake
this activity if a new intruder arrives while the dominant male is engaged.
During nest-digging, the male may court the female by coming to rest beside
her and quivering; by swimming about over her, touching her dorsal fin with
his body and fins; and occasionally by nudging her side gently with his snout
(Schultz, 1938).
When the pit is finished, the female drops into it, pushing her vent well
down toward the bottom of the pit. She is immediately joined by the dominant
male, sometimes by one or more subordinate males as well. The fish open their
mouths, quiver and eject the eggs and sperm. The female at once swims to the
upstream edge of the nest and digs. The eggs are quickly covered with the
gravel displaced from the new pit, and the whole process is repeated until the
female has voided all her eggs, a process which may take several days. The
male then leaves and may mate with another female. The female continues to
dig for a week or more, the digging becoming haphazard and aimless as she
weakens. She finally dies, as do the males.
Fecundity of females varies with the population and the size of the
individual. Average number of eggs per female varies from about 2,600 to
8,500, with northern fish generally producing more eggs than southern. In
Alaska, egg numbers range from 4,242 to 13,619 (Rounsefell, 1957; Yancey and
Thorsteinson, 1963).
The eggs are large, 6-7 mm in diameter, orange-red in color and demersal.
They hatch in 7-9 weeks in the southern part of the range, probably up to 12
weeks, or perhaps a little longer, in interior Alaska. The alevins remain in
the gravel until the yolk sac has been absorbed, usually about 2-3 weeks after
hatching, then work their way up through the gravel to become free-swimming,
feeding fry.
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The fry may go to sea after only three months In fresh water (fall-run
fish in the Puget Sound area) (Mason, 1965), or may stay for as long as two
(rarely three years) (Yukon River). Most chinooks stay one year in the stream
before migrating. During this period, they feed chiefly on terrestrial insects,
chironomids, corixids, caddis flies, mites, spiders, aphids, small crustaceans -
virtually anything available to them, although they do not appear to eat fishes
during their freshwater life. Growth is moderate during the freshwater life. .
By the end of the first year, the young chinooks are 100-150 mm long, and by
the end of the second year they may exceed 200 mm.
As the time for migration to the sea approaches, the young fish "smolt-up",
lose their parr marks and become silvery. They tend to seek deeper water and
avoid light, and the major downstream movements occur at night (Meehan and
Siniff, 1962). In the sea, the young fish remain for a time near shore, where
they feed mostly on herring, sand lance, terrestrial insects, crustaceans and
molluscs (Foskett, 1951). In some areas, there are seasonal changes in the
diet, with euphausiids important in early spring with a shift to fishes in the
summer (Prakash, 1962).
Some of the chinooks may remain close inshore throughout their life (Milne
and Ball, 1958), but others undertake extensive migrations. Fish from Califor-
nia to British Columbia reach the outer waters of southeastern Alaska. Others
from the same populations and from Alaskan streams go even farther, entering
the Gulf of Alaska gyre and moving extensively across the north Pacific. In
the spring of the year, they seem to be scattered across the north Pacific and
in the Bering Sea, while during the summer their numbers increase in the
Aleutian - western Gulf of Alaska region (Manzer et al, 1965). Many of the
inshore fish in southeast Alaskan waters appear to be of local origin (Parker
and Kirkness, 1951; Mason, 1965; Royce et al, 1968). During ocean life, growth
is rapid, often averaging 0.5 kg per month. The chinooks feed largely on her-
ring and sand lance, as well as on pilchards, walleye pollock, Pacific cod,
tomcod, smelts, sand fish, rockfishes, sticklebacks, shiner perch, wolffish,
anchovies, squid, euphausiids, amphipods and crab larvae (Pritchard and Tester,
1939, 1940, 1942). Sexual maturity is reached at ages of 2-7 years. Fall run
fish from the Columbia River are mostly 3 and 4 year olds, but the spring run
is dominated by 5 year olds. In the Yukon River, 6 year olds predominate.
The return from the sea begins in the winter, so that the first fish are
near the river mouths by spring. Oust how this oceanic movement is accomplished
is not understood, but it may well be on the basis of inherited responses to
electro-magnetic cues (Royce et al, 1968). The final year in the ocean is
very important for the chinook to attain full growth, so much so that maximum
harvest can be achieved only by harvesting mature fish (Parker, 1960).
Once in fresh water, the home stream is picked out through olfactory
stimuli (Hasler, 1966). Despite the amazingly acute sense of smell, chinooks
apparently do not respond to a wide variety of pollutants and are not deterred
by them. On the other hand, minute amounts of extracts of mammalian skin
(human hands, bear paws, deer feet, dog paws and sea lion meat) produce an
immediate alarm reaction and a temporary halt to upstream migration (Brett
and MacKinnon, 1952, 1954),
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IMPORTANCE TO MAN
The chinook is a most important part of the native subsistence fisheries
of Alaska and British Columbia, and in former times held a similar position in
Washington, Oregon and northern California.
Commercially, most of the chinook catch is taken by trolling and is sold
fresh or frozen. Some are taken by purse seines and gill nets, also, and most
of these go to canneries. In terms of poundage, the chinook, along with the
coho, is a relatively small part of the Alaska salmon catch. In recent years,
Alaska has produced an average of about 5,319,432 kg (11,702,750 Ib) per year,
representing between 3.3 and 5.51 of the total Alaskan salmon catch. However,
chinook is the highest priced of the salmons so the value of the catch is on
the order of 10% of the total.
Chinooks are usually red-meated, but some are white-meated. Flesh color
is probably genetically controlled. About 70% of the chinook of southeastern
Alaska are of the red-meated form, the remaining 30% white. (Finger and Arm-
strong, 1965). The red chinooks command a higher price.
As a sport fish, the chinook is highly regarded wherever it is found. Its
large size, fighting ability and eating qualities all make it a much-sought-
after species. In the Strait of Georgia region of northern Washington and
southern British Columbia, as many as 93,000 adult chinooks have been taken by
anglers in a single year (Scott and Grossman, 1973). In southeastern Alaska,
the Juneau and Ketchikan areas' sport catches account for 2,000 to 2,500 fish
a year. Herring is the most popular bait in this region, with strip cut most
successful followed by plug cut and whole herring. Spinning from drifting or
anchored boats is the most successful method (Finger and Armstrong, 1965).
Various artificial lures are also quite popular.
The viscera of chinook salmon, normally discarded, has a quite high vita-
min A content and has been used successfully as food for hatchery fish. In
this respect, it is said to be superior to beef liver (Butler and Miyauchi,
1947; Burrows and Karrick, 1947).
CHUM SALMON
0_nc_orhynchus keta (Walbaum)
DISTINCTIVE CHARACTERS
Chum salmon are distinguished by the lack of distinct black spots on back
and tail and the presence of 18-28 short, stout, smooth gill rakers on the
first arch.
DESCRIPTION
Body elongate, fusiform, slightly compressed. Depth about 25% of fork
;h, usually slightly deeper in mature males than in females. Head about
)f fork length, but much longer in breeding males due to elongation of
length. ---,----.
22% of fork length, but
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snout and lower jaw. Snout usually blunt, moderately long, about 35% of head
length in females, but much longer in breeding males. Eye round, rather small.
Mouth large, terminal. Jaws of breeding males develop noticeable hook or kype.
Teeth caniniform, well developed on both jaws, head and shaft of vomer, pala-
tines and tongue. Gill rakers short, stout, smooth, rather widely spaced, 18-
28 on first arch. Branchiostegals 12-16 on each side. Lateral line with 124-
153 pored scales. Pyloric caeca numerous, 163-249.
Fins
D. 10-14. Adipose dorsal present. A. 13-17. P-j. 14-16. Pg. 10-11, an
axillary process present. Caudal truncate to slightly emarginate.
Scales
Small, cycloid.
Color
Large fish in the sea are dark metallic blue dorsally, becoming silvery
on sides and belly. Fine dark speckling may be present, but no definite, large,
black spots. At spawning, the fish become dirty red on the sides and dusky be-
low, with greenish bars or mottlings on the sides. Females are less brightly
colored than males.
Size
Known to reach 102 cm and 15 kg (40 in and 33 Ib), but average size of
mature fish in most places is about 60-70 cm and 4.5-6 kg.
RANGE AND ABUNDANCE
The chum salmon ranges from the Sacramento River in California (and as a
stray as far south as Del Mar), north to the Arctic, east at least as far as
the McKenzie and Anderson rivers, west along the arctic coast of Siberia to
the Lena River, and south on the Asian coast to Korea and Japan. In North
America, it is most abundant from the Columbia River to Kotzebue Sound, The
Noatak and Kobuk rivers support runs estimated at about 1,000,000 fish each
(Smith et al, 1966), but north of this area the runs are too small to be of
more than incidental value.
HABITS
The chum salmon is typically a fall spawner, with the greatest spawning
activity occurring in September and October in most localaties. However, the
first spawning run in the Yukon appears in the lower river in June or even
earlier, often before the ice goes out. A ripe male was found near Fairbanks
in February, and spawning has been recorded as late (or early) as April on
Vancouver Island (Wickett, 1964). In interior Alaska, the first spawners
appear in the Chena River, near Fairbanks, in late July and the run is at its
height about a month later. In the upper Yukon Ri%ver, the major portion of the
86
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run passes Eagle in late August and early September. The Yukon River run of
chums reaches to the headwaters in Teslin Lake, some 3,200 km from the mouth.
In most rivers, however, chums do not travel far, seldom going more than 160 km
or so upstream.
Spawning usually occurs over gravel 2-3 cm in diameter, but they have also
been seen to use coarser stones, even bedrock covered with small boulders. In
the upper Yukon drainage, and also in the Amur River in Siberia, at least some
of the fish select spawning sites in springs or ground water seepages, pre-
sumably a protection against freezing of the redds (Birman, 1953; Kogl, 1965).
In preparing the nest, the female faces upstream, turns on her side and gives
several powerful flaps of her tail, thus displacing the smaller particles of
the bottom which are carried away downstream by the current. Continued digging
at the same spot produces an elongate pit somewhat longer than the fish and up
to about 40 cm deep. The female drifts into the pit, which apparently stimu-
lates the attending males. She is joined by the dominant male, both fish open
their mouths, quiver and extrude the sex products into the nest. The female
may then proceed to dig a new pit at the upstream end of the first one, cover-
ing the newly laid eggs as she does so. The process is repeated until all
her eggs are shed. The males take no part in nest digging, but are aggressive
towards each other. One male usually dominates the rest, driving them away
from the nest site. However, a female may spawn with several males, and a
male may mate with more than one female. After spawning is completed, the
female remains near the nest and may continue digging, although in a haphazard,
unorganized way. Fish of both sexes die a few days after spawning.
In the Amur River of Siberia, "summer" and "fall" runs are recognized.
The "fall" fish are larger (average length 72-75 cm vs 58-61 cm) and more
fecund (average 3,366 eggs per female vs 2,468) than the summer fish (Nikolskii,
1961). The chum of the Yukon and Kuskokwim Rivers, and possibly of other
northern Alaskan streams, appear to correspond to the "fall" chum, while the
fish utilizing streams of southeast Alaska and farther south resemble the
Asian "summer" chum (Lovetskaya, 1948).
The eggs are large, up to 7 mm in diameter before fertilization (McPhail
and Lindsey, 1970) and swell even further, to 8-9.5 mm after being fertilized
(Mahon and Hoar, 1956). Major mortality of eggs seems to occur between ferti-
lization and the eyed stage (Hunter, 1959). Hatching occurs from Demember to
February in the more southerly parts of the range, depending largely on temper-
ature. Time of hatching in interior and northern Alaska is not known, as it
must take place while the streams are still ice-covered. However, eggs ferti-
lized on 30 September and placed in artificial redds in the Chena River, near
Fairbanks, hatched between 17 November and 31 December (Kogl, 1965). The
alevins remain in the gravel until the yolk sac is absorbed, 60 to 90 days
after hatching (Nikolskii, 1961). They then make their way up through the
gravel and begin the migration to the sea. The young chums can stand tempera-
tures up to about 23.8°C, but, with pinks, are the least resistant of the
Pacific salmons to prolonged exposure to high temperatures (Brett, 1952).
Young chum with only a short distance to travel probably do not feed until
they reach the ocean. However, those which must spend several days to weeks
on their journey feed actively on chironomid larvae, daphnids and bosminids
and may attain significant growth before reaching the sea (Sparrow, 1968).
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Most of this seaward movement is accomplished at night, the little fish hiding
in the stream bottom during the day (Neave, 1955; Hoar, 1956). However, when
the migrations are long, they may travel in daylight as well. The young chum
may form schools while still in fresh water and always do so upon reaching the
estuaries.
In the sea, the young chum, now up to 70 mm long, remain close to shore
for several months before dispersing into the open ocean. Alaskan chum salmon
occupy the Chukchi and Bering seas, westward along the Aleutian chain to about
178°E, south in the Gulf of Alaska to about 43°N, and eastward to the North
American coast. Chums from the southern part of the North American range tend
to stay in the Gulf of Alaska and regions along the coast to the south, while
Asian chums may sp.read out through virtually the entire North Pacific Ocean.
The fish are found from close to the surface down to at least 61 m. There is
some indication of vertical diurnal movement, tending toward the surface at
night and deeper during the day (Manzer, 1964). This is probably a response to
movements of food organisms.
Growth is fairly rapid in the sea and the chums attain weights of 3.6-
5.5 kg (8-12 Ib) in three or four years. A large portion of this growth is
accomplished during the final year in the sea, so much so that it has been
estimated that high seas fishing for chum results in a net loss of total yield
of more than 50% (Ricker, 1964). After three or four years in the sea, the
adult chums return to spawn in the natal stream. The average time spent in
the sea varies to some extent with geographical location. In general, fish
from the southern part of the ranqe return in their third or fourth year of
life, while those from the Yukon (and probably other far northern rivers as
well) are mostly in their fourth and fifth years (Gilbert, 1924; Chatwin,
1953). Fish which have only a short freshwater journey ahead of them may
begin to show pale flesh, less oil, sexual dimorphism and spawning colors
even before they enter the river. By contrast, fish facing a long freshwater
trip are still silvery, the flesh red and oily, and the males have not yet
developed a kype. The closer to the spawning grounds the fish are, the more
pronounced are the secondary sex characters, spawning colors and loss of fats.
Upon reaching the sea, the young chum salmon feed largely on small
crustaceans of many varieties, terrestrial insects and young herring (Foskett,
1951). In an experimental situation, copepods between 1.6 and 4.5 mm long-
were the preferred food, to the almost total exclusion of other items (Le
Brasseur, 1966b). Farther at sea, the diet consists largely of copepods and
tum'cates (Larvaceae) {Manzer, 1969) as well as euphausiids, pteropods, squids
and a variety of fishes. Upon reaching fresh water on the spawning run, the
adults cease feeding. The digestive tract may undergo considerable shrinkage
and degeneration at this time.
IMPORTANCE TO MAN
Although generally regarded as one of the less desirable species of
salmon, the chum has always been an important source of food for native peoples,
as well as for their dogs. Large numbers are still taken with gill nets and
fish wheels and are smoked and/or dried for winter subsistence. The commercial
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fishery for chum began about 1893 on the Columbia River when scarcity of
other species made the chum profitable (Craig and Hacker, 1940). However,
the yellowish color and low fat content of the flesh make for a product less
desirable than sockeye. The price for chums is scarcely higher than that for
pinks. Still, the species is abundant and Alaskan landings in recent years
have ranged between 10,300 and 25,416 metric tons (22.6 to 55.9 million Ib)
with values to the fishermen on the order of 10 to 13 cents a pound (Anony-
mous, 1971, 1972, 1973, 1974). Fluctuations in landings are in response to
several factors, including abundance, price, and abundance of higher priced
species. The chum salmon is not considered a sport fish. Nevertheless, chums
will often take small spinners and the angler equipped for Dolly Varden or
grayling who hangs a chum will find he has tangled with a worthy opponent.
COHO SALMON
Oncorhynchus kisutch (Walbaum)
DISTINCTIVE CHARACTERS
The coho is characterized by the presence of small, black spots on the
back and on the upper lobe only of the caudal fin, and by the lack of dark
pigment along the gum line of the lower jaw.
DESCRIPTION
Body elongate, streamlined, somewhat compressed. Depth 22-25% of FL,
deepest in adult males. Head about equal to body depth, longer in breeding
males. Snout bluntly pointed, about 331 of head in adult females, about 40%
in breeding males. Eye small, round. Mouth terminal, gape reaches well be-
hind eye; upper and lower jaws of breeding males hooked (kype). Teeth sharp,
well-developed, present on both jaws, head and shaft of vomer, palatines and
tongue, but not on basibranchials. Gill rakers 18-25 on first arch, rough,
widely spaced. Branchiostegals 11-15, Pyloric caeca 45-114. Lateral line
nearly straight, 112-148 pored scales.
Fins
D. 9-13, a slender adipose dorsal present. A. 12-17, first ray longest
in young, resulting in a curved outer margin. P-j. 13-16. Pg. 9-11, axillary
process present.
Scales
Cycloid, small.
Color
Fish in or fresh from the sea are dark metallic blue or greenish on the
back and upper sides, brilliant silvery on mid- and lower sides, white below.
Small, black spots present on back and upper sides and on upper lobe of caudal
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fin. Fish in breeding condition turn dark to bright green on head and back,
bright red on the sides, the belly often dark. Females are less brightly
colored than males.
Size
The size record was long held by a fish of 14.1 kg (31 Ib) caught in
1947 in Cowichan Bay, B. C. However, a fish of 15 kg (33 lb) was taken re-
cently in the Manistee River, Michigan (Scott and Grossman, 1973). Usual
weights of adults are between 2.7 and 5.4 kg (6-12 lb). Such fish are usually
between 630 and 900 mm (2-3 ft) long.
RANGE AND ABUNDANCE
The natural range of the coho is from Monterey Bay, California (and
apparently as strays in the sea as far south as Chamalu Bay, Baja California)
north to Point Hope, Alaska, westward across the Bering Sea to the Anadyr
River in Siberia and south along the Asian coast to Japan. In addition to
the Great Lakes introductions mentioned above, echoes have also been planted
in east coast waters from Maine to Maryland. A few populations apparently
have survived in New Hampshire. California, Oregon, Washington, Alaska and
Alberta have planted echoes in lakes to provide sport fishing. These plants
are strictly "put and take", for the planted fish do not spawn and the stocks
must be renewed periodically. Despite the fact that the returns to anglers
represent but a small percentage of the number of young fish planted, the
echoes provide a significant portion of the sport catch in these waters,
Cohoes are nowhere as numerous as are chums, pinks and sockeyes. Never-
theless, in most areas the coho probably should be considered as at least
moderately abundant. Abundance is, of course, seasonal, depending on the
arrival of the spawning run. At other times of the year, adult anadromous
coho are scarce to absent.
HABITS
Coho salmon may enter fresh water on their spawning run any time from
mid-summer to winter. As a rule the winter (December-January) entries occur
in the southern parts of the range, with appearance in fresh water becoming
progressively earlier to the north (Briggs, 1953; Shapovalov and Taft, 1954;
McHenry, 1973). The preferred streams are usually short, coastal streams, but
coho salmon are known to spawn in Yukon River tributaries several hundred
kilometers from the sea. Initial entrance of the fish into fresh water seems
to be triggered by a rise in river level, and possibly also by a rise in water
temperature (Briggs, 1953; McHenry, 1973).
On arrival at the spawning area, the female selects a suitable spot and
begins to dig the redd. She turns on her side, facing upstream, and gives
several powerful flips of her tail. The currents and suction so generated
raise debris, silt, sand and pebbles off the bottom to be carried downstream
by the current. Repeated digging in the same spot at intervals of 2-5 minutes
produces a pit somewhat longer than the fish and usually 200-250 mm deep.
The female is aggressive towards other females, driving them away vigorously.
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The attendant male takes no part in nest digging, but usually stays slightly
downstream and to one side of the female. He courts her by occasionally
swimming close to her side, sometimes coming to rest beside her and quivering;
by swimming over her and touching her dorsal fin with his body and fins; and
by gently nudging her side with his snout (Schultz, 1938). The male drives
off other males from the nest site.
When the pit is finished to the satisfaction of the female, she drops
into its deepest part. The male joins her at once. The two fish, side by
side in the pit, open their mouths, quiver, and expel! the eggs and sperm.
The eggs drop to the bottom of the pit, lodging in crevices between the
stones, and are fertilized by the cloud of milt. The female immediately moves
to the upstream edge of the nest and starts digging a new pit. The gravel so
removed is carried downstream and covers the eggs just deposited.
Nest digging and spawning continue at intervals for several days to a
week, until the female has deposited all her eggs. The male then leaves and
may seek another female. The spent female usually continues to dig, but
weakly and irregularly, until she dies.
The number of eggs a female deposits varies with the size of the fish,
the stock and sometimes the year. Numbers have been reported from 1,440 to
5,700; the average is probably between 2,500 and 3,000.
The eggs are fairly large, 4.5-6.0 mm in diameters orange-red in color.
Development time varies with temperature, but usually takes 6-7 weeks. De-
velopment periods up to 115 days have been reported from Asia (Nikolskii, 1961).
The young remain in the gravel until the yolk sac has been absorbed (2-3 weeks
or more), then emerge as free swimming fry and begin feeding.
The coho fry feed mainly on terrestrial insects, especially Diptera and
Hymenoptera. Hemiptera (aphids) and Thysanoptera (thrips) may also be im-
portant. The diet may also include mites, beetles, collembola, spiders and
a little zooplankton. Feeding at this period involves shore-oriented cruising
by individuals and schools, at least when .the fish are in a lake (Mason, 1974a).
As the young fish grow, they take larger food items and often become serious
predators on young sockeye salmon. In Chignik Lake, Alaska, young coho have
been found to eat seven times as many sockeyes as do Dolly Varden, and in other
localities may be equally serious predators (Pritchard, 193€b; Ricker, 1941;
Roos, 1960; Reed, 1967a).
The young coho normally spend a year in fresh water before going to sea,
although some may go to sea at the end of the first summer and others may stay
two or even three years in fresh water. Fish which stay more than two years in
fresh water may become sexually mature without ever going to sea. However,
these "residuals" never spawn, hence all residuals are the offspring of anadro-
mous parents. Early migration may be the result of over-crowding (Mason, 1974b)
Summer carrying capacity of a stream can be greatly increased by supplemental
feeding, but this benefit is lost if winter capacity cannot be increased as
well (Mason, 1974c).
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During their life in freshwater streams, the young coho stay almost en-
tirely in pools, avoiding riffle areas. They soon become strongly territorial,
defending their space against other coho and against other salmonids (Hartman,
1965).
After a year in fresh water, the young begin to turn silvery ("srnolt up"),
avoid light and seek deeper water. They move downstream, usually at night,
and reach the sea in the spring to early summer.
When the young fish reach the sea, they tend to stay fairly close to
shore at first, feeding chiefly on planktonic Crustacea of various sorts. As
they grow larger, the fish move farther and farther from the home river and
feed on larger organisms. Most coho in the sea feed chiefly on herring, with
sand lance second in importance. Fishes make up 70 to 80% of the food, the
remainder being invertebrates (Pritchard and Tester, 1943; Prakash, 1962).
Some populationss however, remain on the crustacean diet. Such fish generally
do not get as big as those that change to a fish diet (Prakash and Milne, 1958)
The oceanic movements of coho in the southern part of the range seem to
be chiefly coastwise, and some fish apparently never venture far from land
(Milne and Ball, 1958; Allen, 1965). By contrast, northern fish, particularly
those from Alaskan streams, spread out all across the north Pacific and into
the Bering Sea. It has been shown that these fish travel "downstream" in the
Alaskan gyre and other major current systems, making one complete circuit per
year (Manzer et al, 1965; Royce et al, 1968).
Having spent two or three years in the ocean and reached full adulthood,
the coho now return to their natal stream to spawn. As a rule, 85% or more
of the fish return to the proper stream. The mechanism by which they make
the return journey through the open ocean is not known, but it has been sug-
gested that inherited responses to electro-magnetic cues are involved (Royce
et al, 1968). Once in the river, the fish home, apparently on the basis of
olfactory cues, to the tributary from which they migrated. The place of re-
turn apparently is impressed on the fish at the time of smoltation, so that
fish transplanted as yearlings will return to the transplant location rather
than to the place where they were reared (Donaldson and Allen, 1958).
IMPORTANCE TO MAN
Although the coho does not rival the sockeye, pink or chum in terms of
numbers, it is nevertheless an imporatnt part of the fisheries of the Pacific
coast of North America. Alaskan landings alone in recent years have averaged
about 4.8 million kg per year, with an annual value to the fishermen of about
$2,700,000. The fish are taken by purse seine, gill net or trolling and are
canned, mild-cured, smoked, or sold fresh or frozen. Frozen coho is a luxury
item on the east coast.
In addition to its commercial value, the coho is highly regarded as a
sport fish. It is taken in the sea by anglers using trolling lures of various
sorts. It is also taken in fresh water on large flies, spoons, spinners and
various sorts of bait.
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The coho was introduced into Michigan waters of the Great Lakes in the
spring of 1966. The fall of the same year, Michigan commercial fishermen
started catching jacks of up to 1 kg or heavier, and an angler caught a coho
of 3.2 kg. The fish spread rapidly and successfully, aided by further intro-
ductions in other lakes. In the first nine months of 1970, U. S. anglers in
the Great Lakes took 320,000 kg of coho salmon. The coho of the Great Lakes
grow rapidly, utilizing the enormous populations of smelt and alewife for food,
and by the time they are three year old adults they average about 2.5 kg in
weight (Scott and Grossman, 1973).
SOCKEYE SALMON AND KOKANEE
Oncorhynchus nerka (Walbaum)
DISTINCTIVE CHARACTERS
The sockeye is distinguished by having 30-40 long, fine, serrated, closely
spaced gill rakers on the first arch, and by lacking definite spots on back and
tail.
DESCRIPTION
Body elongate, fusiform, somewhat compressed laterally. Depth of body
about 20% of FL, somewhat deeper (about 25% of FL) in spawning males. Head
about 221 of FL, longer in breeding males due to elongation of the jaws and
development of the kype. • Snout rather long, about 40% of head in females and
immature males, considerably longer in breeding males. Eye round, small,
about 10% (8-161) of head-length. Mouth large, terminal, reaching well be-
hind eye. Teeth canine-like, well developed, present on both jaws, palatines,
head and shaft of vomer, and on tongue. Gill rakers long, slender, serrated
and close set, 30-40 on first arch. Branchiostegals 11-16. Lateral line
straight, with 120-150 pored scales. Pyloric caeca 45-115.
Fins
D. 11-16, a dorsal adipose fin present. A, 13-18. P1. 11-21. Pg. 9-11,
axillary process present. Caudal fin emarginate,
Scales
Small, cycloid, the margins often resorbed in spawning fish.
Color
Pre-spawning fish are dark steel-blue to greenish blue on the top of
head and back, the sides silvery, belly white to silvery. There are no defi-
nite spots on the back, although some fish may have dark speckling and there
may be some irregular marks on the dorsal fin. At spawning, the head of the
males becomes a bright green to olive green, with black on the snout and upper
jaw, lower jaw whitish, and the whole body turns brilliant red. The dorsal
adipose and anal fins turn red, the paired fins and tail generally grayish to
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green or dark. Various populations may show less brilliant colors, and a few
turn dull green to yellowish, with little if any red. Females are generally
less brilliantly colored than males.
Size
The sockeye is known to reach at least 840 mm and 7 kg (33 in and
15.5 Ib), but the average fish in most areas will be in the neighborhood of
650 mm and 3-5 kg (25 in and 6-11 Ib). The kokanee is much smaller, averaging
about 350 mm and 0.5 kg (14 in and 1 Ib) or less.
RANGE AND ABUNDANCE
The sockeye salmon ranges from the Klamath River in California and Oregon
north to Point Hope, Alaska. Stragglers have been reported from Bathurst In-
let in the Canadian Arctic Ocean. The .most northerly population of any size
is that of Salmon Lake on Alaska's Seward Peninsula. Sockeye have been taken
in the Yukon River as far upstream as Rampart (UAFC #2023). On the Asian side
of the Pacific, sockeye are known from the Anadyr River in Siberia south to
Hokkaido, Japan. The abundance of sockeye varies tremendously from place to
place. The greatest centers are in the Fraser River system in British Colum-
bia and in the Bristol Bay system - the Kvichak, Naknek, Ugashik, Egegik and
Nushagak rivers of Alaska. In good years, the runs to these areas may be in
the tens of millions of fish. The kokanee is known from Japan and Siberia
on the Asian side, and in North America from the Kenai Peninsula in Alaska,
south to the Deschutes River in Oregon. It has been introduced widely into
the Great Lakes, New England, various western and mid-western states, and
California. Kokanee may be extremely abundant in some lakes, while in others
the populations are small. There seems to be no particular explanation.
HABITS
The adult sockeyes return to their natal streams during the summer and
fall, July to October in most areas, but as late as December in the southern
part of the range. Spawning occurs almost exclusively in streams with a lake
connection, although a few populations spawn in lakes and a very few in streams
with no lake. Lake spawners tend to breed later in the year than do stream
spawners, but this is by no means universal.
As with other salmons, the female selects the site and digs the nest.
Nest digging is accomplished by the female facing upstream, turning on her
side on the stream bottom and giving several powerful strokes with her tail.
This disturbs debris, sand and gravel, which is carried downstream by the
current. Repeated digging at the same spot produces a pit somewhat longer
than the fish and as much as 400 mm (16 in) deep (Hanamura, 1966). The nest
is usually constructed where the bottom is of fine gravel, but may be over
large pebbles of 5-10 cm diameter or even amongst large rocks. In the last
case, no proper nest is dug (Ricker, 1966). Preferred sites have less than
10% of the gravel larger than 75 mm in diameter, about 501 between 25 and 75
mm, and the remainder smaller than 25 mm (Hoopes, 1962). During preparation
of the nest, the female is attended by a dominant male and often fay several
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subordinate males as well. The female is aggressive towards other females and
sometimes towards the subordinate males. The male is aggressive towards other
males.
Between digging acts, the female often rests over the pit. The dominant
male courts her by approaching her from behind, gently nudging her side with
his snout, then coming to rest beside her and quivering. He may also swim
over her, brushing her dorsal fin with his body and fins.
When the nest is completed, the female drops into it with her vent and
anal fin well down in the deepest part of the pit. The male at once comes
close beside her, both fish arch their bodies, open their mouths, quiver and
extrude the eggs and sperm. One or more subordinate males may come to the
other side of the female and join in the spawning. Three to five days are
normally required to deposit all the eggs.
The eggs, which are bright orange-red in color and 4.5-5 mm in diameter',
fall into crevices in the gravel at the bottom of the nest, where they are
fertilized by the male's milt. A female may deposit 500-1,000 eggs in a
single nest. Having done so, she moves to the upstream edge of the nest and
digs again. The gravel so disturbed is carried into the nest, covering the
eggs, and a new nest is dug just upstream from the previous one. The female
usually produces between 2,500 and about 4,300 eggs, and will occupy 3-5 nests.
She may spawn with several dominant males, and a male may breed with several
females. All adult sockeye die after spawning.
Development of the eggs takes 6-9 weeks in most areas, the time depending
largely on water temperature, but may require as long as 5 months (Hart, 1973).
Hatching usually occurs from mid-winter to early spring and the young emerge
from the gravel in April to June. There is strong positive correlation between
intra-gravel water flow and egg survival. Deposition of silt in the redd, re-
ducing water flow, may result in heavy mortality of the eggs. Oxygen consump-
tion by the alevins in the nest is considerably higher than by eggs, further
emphasizing the need for adequate flow (Cooper, 1965).
Upon emerging from the gravel, the fry at first tend to avoid light,
hiding in the stones and gravel of the stream bottom during the day and
emerging at night. In a few populations, the fry go to sea during their
first summer, but the vast majority spend one or two years in a lake before
migrating, with rather rare cases of three or even four years in freshwater
(Margolis, et al, 1966). The young sockeye in inlet streams go downstream
to the lake, while those hatched in outlets swim upstream. The mechanisms
of these different responses are unknown, but appear to be genetically con-
trolled.
Once in the lake, the young fish spend the first few weeks close to
shore, feeding largely on ostracods, Bostm'na and insect larvae. Subsequently,
the fish become pelagic and move offshore, where they feed on plankton in the
upper 20 m or so. The major food items during the summer are Calarius and
Bosmina, with the latter preferred by the younger fish (Hoag, 1972). The
abundance of the fry is positively correlated with the abundance of adult
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spawners, and their growth through June is correlated with water temperature
and an abundant food supply (Nelson, 1964; Narver, 1966; Rogers, 1973), Ex-
perimental fertilization of lakes to increase the standing crop of plankton
has resulted in up to 30% greater size of under-year!ing sockeyes. However,
the increase in plankton was much greater. Apparently the upper layers of
water are too warm for the young sockeye in July and August, leading to under-
utilization of the food (Le Brasseur and Kennedy, 1972; Barraclough and Robin-
son, 1972).
After a year in the lake (two years in most Bristol Bay areas), the young
sockeye lose their parr marks, turn silvery ("smolt up") and migrate downstream
to the sea. Smoltification is largely dependent on size, rather than age, and
the threshold size seems to be determined by the genetics of the particular
stock. Early-spawning adults tend to produce a greater percentage of age I
smelts than do late spawners (Narver, 1966). In the Wood River system of
Alaska, the first smolts to migrate are age II, which are probably late mi-
grants from the upper lakes which encountered a temperature block and were
forced to spend a second year in the lake (Nelson, 1964). In other systems,
however, age II smolts seem to be the usual thing. Peak outmigration in the
Bristol Bay region occurs in June, beginning when water temperatures reach
about 4°C (38-39°F). Most of the migrants move at night, especially between
10 and 11 p.m. (Nelson, 1964), but schools of smolt leave the Agulowak River
for Lake Aleknagik all day long in mid-July (personal observations), and in
Babine Lake, British Columbia, the main movements are at dawn and dusk (Groat,
1965), The migrating smolts show well-directed orientation towards the lake
outlet and travel 5-8 km/day in the lake, farther on bright days than on cloudy
ones. During the course of the migration, the orientation shifts according to
the orientation of the bodies of water traversed. Various mechanisms, in-
cluding responses to celestial phenomena, polarized light and the immediate
surroundings seem to be involved (Groot, 1965).
Once in the sea, the young sockeye stay fairly close to shore at first,
feeding mainly on various zooplankters, but also on small fishes (Manzer, 1969).
As the fish get bigger and stronger, they head out to sea. Fish from areas
south of the Alaska Peninsula head out into the Alaska Gyre in the Gulf of
Alaska. Those from Bristol Bay and northward go through the eastern and central
Aleutian passes into the Gulf. By late winter, the sockeyes are spread in a ,
band across the North Pacific south of 50°N. In late spring, the young fish
move north to between 50°N and the Aleutians, from about 160°W to 170° or
165°E. This appears to be an important feeding area, the major foods of the
sockeye being amphipods, copepods and squid (Le Brasseur, 1966a). The follow-
ing winter, the fish separate into those which will mature the following spring
and those which will spend another year or more in the sea. The early-maturing
fish stay north of 50°N, while the others repeat the previous year's journey.
In the following spring, the mature fish head back to their natal streams,
while the immatures continue to repeat the oceanic circuit (Thorsteinson and
Merrell, 1964; Manzer et al, 1965; Margolis et al, 1966; Royce et al, 1968;
French and Bakkala, 1974).
Most sockeye from British Columbia areas spend one year in fresh water
and two in the sea, returning to spawn in their fourth year of age. However,
farther north, two years in fresh water and two or three in the sea are common.
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Hence, many Alaskan sockeye return in their fifth or sixth years. However,
the four year cycle is generally prevalent.
Growth of the sockeye in the ocean is rapid, especially in the final
year. Because of this growth pattern, the inshore fishing practiced along
the North American coast is the most efficient harvesting of the resource.
Catching the fish at sea in the year before they mature results in loss of
yield of 50 to 65% (Ricker, 1962).
The landlocked form of the sockeye is called kokanee. It differs from
the anadromous sockeye chiefly in spending its entire life in fresh water and
in reaching a much smaller maximum size.
Kokanee, wherever they are native, have been derived from anadromous
sockeye populations and each kokanee population apparently has evolved in-
dependently from a particular sockeye run (Ricker, 1940; Nelson, 1968a). Off-
spring of kokanee occasionally become anadromous, and sockeye offspring some-
times will not go to sea. The life span of the kokanee varies from two to
seven years in different stocks. The fish is confined to lake-stream systems
and spends most of its life in the lake, analagous to the sea life of the
sockeye. They are much smaller than sockeye, averaging about 300 mm long in
most places, although fish of over 600 mm have been recorded (Kimsey, 1951).
Spawning time and behavior df the kokanee are like those of the sockeye
(Schultz and students, 1935; Schultz, 1938). However, related to their small
size, fecundity is much lower, on the order of 300-500 eggs per female.
Kokanee are either weak swimmers, or, more likely, lack the strong urge
to swim upstream so characteristic of the salmons. At any rate, small barriers
effectively stop their migrations (Seeley and McCammon, 1966). Spawning
usually takes place in inlet or outlet streams, though sometimes in lakes near
the shore (Lorz and Northcote, 1965). All kokanee die after spawning.
Kokanee are mainly plankton feeders throughout their lives. At least in
some lakes, there is seasonal variation in the type and quantity of food eaten.
They feed most heavily in June and October, less through the summer, in Nicola
Lake, B.C. Diaptomids dominated the food in the spring, chironomids in the
summer and Daphnia in the fall (Northcote and Lorz, 1966).
IMPORTANCE TO MAN
The sockeye salmon is one of the most important of the Pacific salmons.
Its flesh is of the highest quality, and although it is not as abundant as the
pink and chum, the higher price it commands makes it the real "money fish" in
the salmon industry. The kokanee is primarily a sport fish, generally well
accepted by anglers, despite its small size. Like the sockeye, the kokanee is
excellent as food. It has been fished commercially in Lake Pend Oreille,
Idaho. It is also well regarded in some areas as a forage fish for large
trout, but in other places is looked upon as a competitor with the trout.
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ARCTIC GRAYLING
Thymallus arcticus (Pallas)
DISTINCTIVE CHARACTERS
Adult grayling are distinguished by the greatly enlarged dorsal fin and
the small mouth with fine teeth in both jaws. Juveniles may be separated from
coregonids of similar size by the presence of narrow, vertically elongate parr
marks. These are round or absent in coregonids.
DESCRIPTION
Body elongate, somewhat compressed. Greatest depth at origin of dorsal
fin, depth about 20% FL. Head short, about 17% of FL. Snout about equal to
or a little less than eye diameter, 21-25% of head length. Eye large, round,
about equal to snout. Mouth terminal, small, maxilla reach about to middle
of eye. Teeth small, present on both jaws, tongue (may be missing in large
adults), palatines and head of vomer. Gill rakers 16-23 on first arch.
Branchiostegals 8-9 on each side. Lateral line straight. Pyloric caeca
18-21.
Fins
D. 17-25, greatly enlarged in adults, especially in males. Adipose
dorsal fin present. Dorsal fin of adult males reaches adipose fin when de-
pressed, shorter in females. A. 11-15. P]. 14-16. ?2- 10-11, rather long,
reaching anal in adult males, shorter in females. A small axillary process
present. Caudal fin forked, lower lobe often longer than upper.
Scales
Cycloid, fairly large, 77-103 pored scales in lateral line.
Color
Back dark purply blue
times with a pinkish wash,
numerous in young. A dark
pectoral and pelvic fins.
to blue gray, sides silvery gray to blue, some-
with scattered dark spots on sides, these more
longitudinal stripe along lower sides between
Dorsal fin dark with narrow purple edge, rows of
on body of fin.
Adipose, dorsal
Pel vies dark
anal, caudal
reddish to orange and/or purple to green spots
with irregular diagonal orange-yellow stripes.
and pectoral dusky to dark.
Size
The largest grayling known is one of 759 mm and 2.7 kg (29 7/8 in, 5 Ib,
15 oz) from the Katseyedie River, Northwest Territories. However, in most
areas the average angler-caught fish would be on the order of 300-350 mm
(10-14 in) long and weigh 450-700 grams (1-1 1/2 Ib).
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RANGE AND ABUNDANCE
The grayling is common throughout Alaska and northern Canada, from the
west side of Hudson Bay to the western shores of Alaska. It is present in
the headwaters of the Missouri River above Great Falls, Montana, and ranges
northward from central Alberta. To the west, it is present on Saint Lawrence
Island and on the Asian mainland west to the Kara and Ob rivers of Siberia,
south to the upper Yalu River.
The grayling has been introduced into a number of lakes in southeastern
Alaska, as well as into mountainous areas of Colorado, Utah and Vermont.
HABITS
Grayling spawn early in the spring, immediately after break-up. In
interior Alaska, they begin to congregate at the mouths of clearwater tribu-
taries in April, and may start upstream through channels cut in the ice by
surface runoff (Reed, 1964). The fish run upstream to the spawning grounds,
sometimes a matter of more than 160 km, as soon as the streams open. Spawning
takes place from mid-May to June, with no particular preference for substrate,
although sandy gravel seems to be used most often, perhaps because of prevalence,
The males establish a territory which they defend against intruding males
by erecting the dorsal fin, opening the mouth, and assuming a rigid posture.
Persistent intruders may be rushed and driven off. Rarely, females may be
attacked. At spawning, the male follows a female, courting her with displays
of his dorsal fin. He then drifts over beside her and folds his dorsal fin
over her back. Both fish arch, vibrate and release eggs and milt. They may
gape, also. No redd is constructed, but the vibrations of the tails during
the spawning act stir up the substrate and produce a slight depression (Laird,
1928; Wojcik, 1955; Kruse, 1959; Reed, 1964; Bishop, 1971).
The eggs are orange in color and about 2,5 mm in diameter but quickly
swell to about 2.7 mm on hardening (Reed, 1964). During the next few days,
they continue to swell and after three or four days are about 3.5 mm in
diameter (Ward, 1951). Egg number varies with size of the individual and
the stock, with counts as low as 416 and as high as 12,946 recorded (Brown,
1938b). Development to hatching requires 11 days to three weeks, depending
on temperature (Lord, 1932; Brown, 1938b; Nelson, 1954; Bishop, 1971). The
young, which have been described as resembling "two eye-balls on a thread"
(Schallock, 1966), begin feeding the 3rd or 4th day after hatching, and all
are feeding actively by the 8th day (Lord, 1932; Brown and Buck, 1939). Growth
is rapid during the ensuing summer. By September, the young of the year are
about 100 mm (4 in) long. In general, growth is somewhat faster in the
southern part of the range than in the northern, but rates may vary widely
from tributary to tributary within a single drainage (Miller, 1946; Kruse,
1959; Reed, 1964). At age I, grayling of interior Alaska average about 148 mm
(5.8 in) in fork length; at age II, 192 mm (7.6 in); age III, 228 mm (9 in);
age IV, 265 mm (10.4 in); age V, 304 mm (1? in). A 300 mm grayling will weigh
between 200 and 250 grams (about 1/2 lb).
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Following the spawning migration, the adult grayling move away from the
spawning areas, generally but not always going farther upstream, and take up
more or less permanent summer residence in pools (Schallock,, 1966; Vascotto,
1970). Territoriality and social hierarchy develop quickly in each pool, with
the largest and strongest fish occupying the most advantageous positions near
the head of the pool, the smaller subordinate fish farther downstream, and
the smallest of all occupying the foot of the pool without any territories.
Territories are established and maintained through a series of ritualistic
"challenge displays," involving seven distinct steps. First, an "invader" moves
up parallel with a "defender." 2. Both fish then lie still, with dorsal fins
folded. 3. The invader drifts sideways toward the defender until they are
less than 1 cm apart. 4. One fish, usually the invader, moves forward about
15 cm and bends the body so that the inside of the curve is presented to the
head of the defender. 5. The invader drifts toward and beneath the defender.
6. The invader rises towards the defender, who drifts back. 7. Either (a)
the defender keeps on retreating and leaves the territory, or (b) the defender
moves around the invader and the pattern is repeated, with the roles reversed.
Once territories and hierarchies have become firmly established in a pool, the
ritual rarely goes beyond the third stage (Vascotto, 1970; Vascotto and Morrow,
1973).
The establishment and use of the territories is related to the feeding
habits of the grayling. Virtually the entire diet is composed of insects,
larvae, pupae and adults, which are taken drifting. The majority of these
are aquatic forms which have been disturbed from the bottom by one means or
another and thus become part of the benthic drift. The grayling is primarily
a surface and mid-depth feeder and does not generally feed on the bottom ex-
cept in the fall, when benthic drift is much reduced. (Brown, 1938a; Reed,
1964; Vascotto, 1970; Vascotto and Morrow, 1973).
A downstream migration occurs in mid-September. The fish leave the
smaller tributaries and the vast majority move downstream to over-winter in
deep water. A few stay in the major clear-water streams and apparently over-
winter in the deeper pools (Schallock, 1966).
IMPORTANCE TO MAN
The grayling is one of the most important sport fishes of Alaska and
northern Canada. It also makes a significant contribution to subsistence
fishing in some remote areas.
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SECTION 6
SMELTS—FAMILY OSMERIDAE
The Osmeridae is a group of small fishes which rarely, if ever, exceed
about 30 cm (1 ft) in length. They are related to the salmons, but differ
morphologically from that group chiefly in having no axillary process at the
base of the pelvic fins; the lower jaw is longer than the upper; the head is,
in general, smaller; the body is more slender, and there are few or no pyloric
caeca.
The smelts are slender-bodied, silvery or dull, and have forked tails.
Sexual dimorphism, usually in the form of nuptial tubercles, appears at spawn-
ing time.
Smelts may be entirely freshwater, anadromous or marine in their life
history. Spawning usually takes place in fresh water or on beaches, although
a few species spawn in the sea.
The smelts are generally quite abundant, at least during the spawning
season, and are of considerable economic value.
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LONGFIN SMELT
S|?in'nchus thalelchthys (Ayres)
DISTINCTIVE CHARACTERS
The longfin smelt is distinguished by the long upper jaw,' reaching at
least to below the middle of- the eye in adults; fine teeth in a single row on
vomer and palatine bones; and 38-47 gill rakers on the first arch.
DESCRIPTION
Body elongate, slender, compressed, its depth 14-20% of fork length.
Head short, 20-251 of FL. Snout 22-28% of head. Eye fairly large, its di-
ameter about equal to snout length. Mouth large, oblique, lower jaw project-
ing beyond upper when mouth is closed, upper jaw reaching backwards at least
to below middle of eye in adults, shorter in young. Teeth small, fine, present
on both jaws, tongue, vomer and palatines. Gill rakers long, 38-47 on first
arch. Branchiostegal rays 7-8. Lateral line incomplete, 14-21 pored scales,
reaching not quite to below dorsal fin. Pyloric caeca 4-6.
Fins
D. 8-10, moderately high, origin near middle of body length. Adipose
dorsal fin present. A. 15-19. PI- 10-12, reaching to origin of pelvic fins.
?2- 8, origin under anterior edge of dorsal fin. Paired fins with tubercles
on. upper sides in breeding males. Caudal fin forked.
Scales
Moderately large, cycloid, 55-61 (rarely 62 or 63) in a midlateral row.
Tubercles present in breeding males.
Color
Dusky to olive or olive brown above, sides and belly silvery white. Fin
rays usually dusky, membranes clear.
Size
Up to as much as 200 mm (8 in), but average adults are about 150 mm
(6 in) or a bit less.
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RANGE AND ABUNDANCE
The long-fin smelt is found along the coast of North America from San
Francisco Bay (Rutter, 1908) north to Prince William Sound, Alaska, where it
has been found near Hinchinbrook Island at 60°12'N, 146°15'W (Dryfoos, 1961).
The record from the Nushagak River in the Bristol Bay area of Alaska (Gilbert,
1895) is an error which has been uncritically accepted even by some modern
authors, even though Hubbs (1925) long ago pointed out that the specimens in
question were pond smelt, Hypomesus olidus. Within this range, the species
is locally and seasonally abundant, appearing chiefly during the spawning runs.
HABITS
Extraordinarily little is known of the biology of the longfin smelt. The
fish are anadromous, although a few landlocked populations are known. Spawning
runs occur between October and December in most areas, with a few populations
breeding as late as February. Details of spawning have not been recorded, but
it is known to take place in streams not far from the sea. Fecundity varies
from 535 eggs in small females of a landlocked stock to more than 23,600 for
large anadromous fish. Average fecundity for anadromous females is said to be
about 18,000 eggs. The eggs, about 1.2 mm in diameter, hatch in 40 days at
6.9°C. The newly hatched larvae are 7-8 mm long. They probably spend con-
siderable time in fresh water, as young fish up to 72 mm long have been taken
in the Fraser River (Hart and McHugh, 1944). Year old fish average about
75 mm, while two-year-old males are about 106 mm, females about 110 mm long.
Sexual maturity is reached at the end of the second year of life. Most indi-
viduals die after spawning, although a few females apparently survive to three
years. It is not known whether or not these three-year-olds have already
spawned at the age of two. Members of landlocked populations apparently do
not reach as great a size as do sea-run fish.
In the sea, the fish apparently stay fairly close to shore. They are
taken in shrimp trawls at depths to 41 meters in the winter time. Food of
the smelt in the sea consists of small crustaceans, especially euphausiids,
copepods, and cumaceans, while in fresh water the young feed on a small,
shrimp-like crustacean, Neoroysis mergedis (Hart and McHugh, 1944), small,
bottom-dwelling crustaceans and insect larvae (Hart, 1973).
IMPORTANCE TO MAN
This smelt is but little utilized. Fish in the sea are reported to have
a good flavor, but the supply is limited and the species is lumped with other
smelts in the market (Hart, 1973). The longfin smelt is often abundant during
the spawning run, but at this time the flesh is soft and oily and spoils
quickly (Scott and Grossman,,1973),
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POND SMELT
Hypomesus oildug (Pallas)
DISTINCTIVE CHARACTERS
The small mouth, teeth in two rows on vomer and palatines, and 51-62
mid-lateral scales distinguish the pond smelt.
DESCRIPTION
Body elongate, somewhat compressed. Depth 16-242= of standard length.
Head short, 20-252 of standard length. Snout short and blunt, its length
usually a little less than eye diameter. Eye round, moderate, diameter equal
to 20-30% of head length. Mouth small and oblique, lower jaw protruding
beyond upper when mouth is closed; maxillary reaching to below pupil. Small,
pointed teeth present on both jaws and on tongue, in two rows on vomer and
palatines. Gill rakers long and slender, 26-34 on first arch. Branchio-
steqal rays 6-8 on each side. Lateral line incomplete, 7-16 pored scales,
reaching scarcely beyond tip of pectoral fin. Pyloric caeca 0-3, rarely 4,
usually 2.
Fins
D. 7-9, its anterior edge slightly before middle of standard length.
fin present.
8, arising
A. 12-18, usually 12-16. P1. short,
below or ahead of origin of dorsal fin.
10-12,
Caudal
Adipose dorsal
rarely 13.
fin forked.
Seales
Large, cycloid, 51-62 in a mid-lateral series. Males develop breeding
tubercles on scales, as well as on head and fin rays.
Color
Light brown to olive green above, with a metallic silvery band along
middle of sides. Ventral regions silvery white. Fins pale.
Size
Reported to reach 185 mm (7 1/2 in) in the Anadyr River (Andriyashev,
1954). Most North American adults are 100-150 mm (4-6 in) in total length.
RANGE AND ABUNDANCE
In Asia, the pond smelt is found from Wonsan, Korea, north to the Chuk-
hotsk Peninsula, westward to the Alazaya River in Siberia. It is also
present on Hokkaido Island, Japan, and on Sakkalin. An isolated population
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is present in Lake Krugloe in the Kara Sea drainage (Ivanova, 1952). In
North America, the pond smelt is known from the Copper River north along
coastal regions to the Kobuk River in Alaska, and from the Mackenzie drainage
as far upstream as Great Bear Lake in northern Canada.
The pond smelt is often extremely abundant in suitable ponds and streams
(Turner, 1886).
This species (or possibly the closely related H_. transp_acjf i cus Japoni_-
cu_s) was introduced into a number of reservoirs in California, but only one
of the introductions, that in Freshwater Lagoon, Humboldt County, seems to
have been at all successful (Wales, 1962).
HABITS
Spawning occurs in shallow water in streams or ponds, over pebbly
bottoms in streams but often in littoral areas of ponds where the bottom is
covered by organic debris (Jordan and Evermann, 1896; Nikolskii, 1956; Nar-
ver, 1966). Time of spawning is late April to May in Asia, but not until
June in the Copper River and certain lakes in Alaska. On the spawning
grounds, females outnumber males by about 3 to 1. Each female produces 1,200
to nearly 4,000 eggs. The eggs are about 0.75 nun in diameter at fertiliza-
tion, but quickly swell to about 0.95 mm on contact with the water. They are
demersal and sticky, adhering to whatever they touch. Like other smelt egqs,
those of the pond smelt form a short pedicle from the inverted outer coat.
However, about two days after fertilization the egg breaks off the pedicle
and thereafter develops pelagically (Nikolskii, 1956). This pelagic develop-
ment has not been reported by other authors. Hatching occurs on the llth
day at temperatures between 11 and 15°C, (Nikolskii, 1956) but not until the
18th - 24th day at 10°C (Narver, 1966).
The larvae are 4-5 mm long, with a well developed yolk sac. They begin
to feed almost immediately after hatching, even before the yolk sac is ab-
sorbed, feeding on minute crustaceans and rotifers (Sato, 1952). When they
are about 10 mm long, the young school up and move close to shore. They
tend to stay in this region for most of the summer, moving into deeper water
with the approach of fall. After a month of planktonic life, the young are
about 20 mm long, and reach 30 to 40 mm by the end of the summer (Nikolskii,
1956). During this time, the larval population may decline drastically, but
the cause is unknown (DeGraaf, 1974).
Pond smelt are about 60 mm (2.4 in) long at the end of the first year
of life, about 80-100 mm (3-4 in) at the end of the second year. Apparently
only a few survive beyond three years of age. Sexual maturity is reached at
the end of the second year of life in most populations, although in some
Japanese populations, spawning occurs at age 1 + , with virtually no survivors.
Most pond smelt are 50-70 mm long by the end of the first year, 70-90 mm at
,age 2, 90-110 mm at 3, and about 120 mm at 4 years of age. The last two age
groups are extremely scarce.
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'A unique population of pond smelt is reported from an unnamed lake on
the arctic coastal plain of Yukon Territory at 69° 18' N, 138° 57' W
(DeGraaf, 1974). If the age determinations are correct, a few of these fish
may survive to 9 years. The growth rates given are extra ordinarily slow,
with 1+ fish smaller than young-of-the-year from other places; five-year-
olds no longer than one-year fish from Black Lake, Alaska; and the supposed
eight and nine year old fish about the same length as Black Lake 2+ fish.
Pond smelt do not migrate as a rule. However, within a lake or pond, -
the adult fish generally stay on the inshore spawning areas in spring and
early summer, later moving offshore. Young-of-the-year follow the same
patterns, while immature fish stay in the pelagial regions all summer.
There may be outmigrations to other parts of the drainage system if popu-
lation are high in relation to the abundance of food (Narver, 1966).
Food of the pond smelt is virtually exclusively zooplankton. Very young
fish feed largely on rotifers, while adults take larger zooplankton such as
copepods and cladocera, as well as insects and, occasionally, algae (Sato,
1952; Nikolskii, 1956).
IMPORTANCE TO MAN
The pond smelt is of no direct importance to man, due chiefly to its
limited distribution. Hhere it is present, it is of value in subsistence
fisheries (Turner, 1886). They are doubtless used locally in Asia as well
as in Alaska.
It has been suggested (HcPhail and Lindsey, 1970) that, since the adults
feed mainly on zooplankton, they may sometimes be important competitors with
young sockeye salmon.
SURF SMELT
Hypomesus pretiosus (Girard)
DISTINCTIVE CHARACTERS
The surf smelt is distinguishable from other species of Hyppmesus by
its mid-lateral scale count of 66-73, and the presence of 4-7 pyloriccaeca.
DESCRIPTION
Body elongate, slender, slightly compressed. Depth 14-22% of fork
length. Head rather short, 19-211 of fork length. Snout moderate, about
30% of head length and about equal to interorbital width. Eye round, its
diameter about equal to snout length. Mouth rather small, upper jaw
(maxilla) reaching to below front half of eye. Teeth small, pointed,
present on both jaws-. Gill rakers 31-36 on first arch. Branchiostegal rays
7-8. Lateral line short and incomplete, reaching about to tip of pectoral
fin. Pyloric caeca 4-7.
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Fins
D. 8-10, its origin above or slightly behind middle of body lenqth. A
small, sickle-shaped adipose fin present. A. 12-16. P-j. 14-17, slightly
longer in males than in females. ?2- 8, origin below or behind front of
dorsal fin. Caudal fin forked.
Scales
Cycloid, 66-73 in a mid-lateral series. Scales are deciduous and easily
removed.
Color
In males, the back is dull olive to brownish, the belly yellowish-
silver. Females are brighter, more metallic green above and silver on the
belly. Both sexes have a bright silver band along the middle of each side,
which becomes dark in preserved specimens.
Size
The surf smelt reaches a lenqth of about 254 mm (10 in) in California
(Roedel, 1953) but the maximum size of northern fish seems to be about 200-
230 nm (8-9 in).
RANGE AND ABUNDANCE
Along the coast of North America from Long Beach, California in the
south (Miller and Lea, 1972) to Chignik Lagoon in southwestern Alaska (Phin-
ney and Dahlberg, 1968). A single specimen was taken in Herendeen Bay, on
the north side of the Alaska Peninsula, in the summer of 1976 (I. Warner,
Alaska Dept. Fish and Game, Kodiak, pers. comm.). On the western side of the
Pacific, another subspecies ranges from southern Kamchatka to Korea.
The surf smelt is most abundant in the central part of its range, espe-
cially in Puget Sound and adjacent regions. Good runs occur in Alaska as
far north as Prince William Sound. In the Columbia River to Puget Sound
area, the surf smelt is fished not only for sport but also commercially.
Landings of smelt contribute significant amounts to the fisheries of Oregon,
Washington and British Columbia, However, abundance is far from uniform.
In Puget Sound, for example, 57.3 linear km (35.8 mi) of beach are used by
spawning smelt, but only 9.8 km produce more fish than the remaining 47.5 km.
HABITS
The surf smelt, as its name suggests, spawns alontj beaches in light to
moderate surf. The spawning period for all populations stretches from May to
March, according to locality and population (Yap-cMonggo, 1941). At spawn-
ing time, schools of smelt approach the beach, the females appearing a few
107
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days before the males. Spawning is most frequent on a falling afternoon or
evening high tide, and takes place from extreme low tide line to more than
half-way up the beach (Thompson et al, 1936; Yap-Chionggo, 1941). Other ob-
servers have found surf smelt spawning most actively within one to one and
one half hours before or after the flood (Loosanoff, 1937). At spawning,
large numbers of males mill about near the edge of the surf. When a female
is seen, numbers of males pursue her and try to get into spawning position.
If she swims into deeper water, the males desert her. A ripe female, how-
ever, moves up the beach, accompanied by 1-5 males who swim parallel to and
slightly behind her. The males press against the female, sometimes tilting
their bodies to bring male and female vents close together. As they reach
shallow water, 25-50 mm (1-2 in) deep, the fish bend and vibrate, releasing
eggs and sperm, then retreat to deeper water. Spawning may take place on
both incoming and outgoing waves (Schaefer, 1936). The spawning act lasts
only 5-10 seconds and may be repeated on several successive waves. A female
usually requires several days to deposit all her ripe eggs, which number
from about 1,500 to over 36,000 (Schaefer, 1936), depending on the race and
size of the female. Females normally contain several size-groups of eqgs at
any given time and apparently spawn more than once in a single season. Al-
though the female releases only a few eggs at a time, the males emit rather
large quantities of milt, so that the water may become slightly milky on a
heavily used beach. Preferred beaches are those not exposed to stronq sun
and composed of coarse sand and fine gravel 2.5-4 mm (0.1-0.16 In) in dia-
meter. Suitable-sized substrate particles appear to be a most important
factor in the selection of a spawning beach.
The eggs, about 1.0-1.2 mm in diameter after water-hardening and pale
yellow in color, are sticky and adhere to the particles of gravel. Shortly
after fertilization, the outer membrane ruptures and turns inside out, ad-
hering to the egg at the point where it sticks to the substrate and forming
a short pedicel. Wave action buries the eggs, usually to a depth of 25-100
mm (1-4 in) but sometimes to as deep as 300 mm (1 ft).
Incubation requires 8-11 days at 12.2-15.5°C, but three,or four times
as long at 7.5-8.5°C. Once development is complete, hatching requires the
stimulus of wetting, which is accomplished by the incoming tide. The younq
are 3-6 mm (0.1-0.24 in) long at hatching, transparent, with yellow eyes and
a small yolk sac. They are positively phototropic and swim actively towards
the light. Thirteen days after hatching they are about 7 mm long.
Very little is known of the subsequent life history. Young smelt 33 mm
(1.4 in) closely resemble the adults and may go up rivers, where they feed
on Diptera and Ephemeroptera. Presumably those fish which do ascend rivers
return to the sea very shortly, for only a few specimens have been taken in
fresh water. One was recorded from the Sandy River, 16 km (10 mi) east of
Portland, Oregon (Me Allister, 1963).
The surf smelt matures at one to two years of age, at a minimum length
of 90 mm (3.5 in), with males maturing earlier than females. Within a single
age group, larger fish mature and spawn earlier in the season than do smaller
ones (Schaefer, 1936; Loosanoff, 1937). Maximum life is about two years for
males, three for females.
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Nothing is known of the movements of the surf smelt. They disappear,
presumably offshore, as young and re-appear inshore as mature adults.
IMPORTANCE TO MAN
The surf smelt is one of the minor constituents of the total commercial
fisheries of the west coast of North America. Total landings run to several
hundred thousand kg per year and it is considered the smelt in British Colum-
bia. As food, it is considered a delicacy, being of fine flavor and texture.
The size of the sport catch is unknown, but it is probably quite large.
EULACHON
Thaleichthys pacifiaus (Richardson)
DISTINCTIVE CHARACTERS
Large canine teeth on .the vomer and 18-23 rays in the anal fin distin-
guish the eulachon.
DESCRIPTION
Body elongate, slender, slightly compressed in females; thicker dorsally
than ventrally (cross-section more or less pear-shaped) in adult males.
Depth 15-20% of standard length. Head short, 20-261 of standard length,
rather prominent concentric striae present on gill covers. Tubercles present
on heads of breeding males, poorly developed or absent in females. Snout
fairly long, 25-28% of head length. Eye round, small, its diameter 50-66% of
snout length. Mouth oblique, large, maxilla reaching to or behind posterior
margin of eye in adults. Small, pointed teeth present on both jaws, tongue
and palatines. A pair of moderately large canine teeth on vomer. All teeth
tend to be lost at spawning. Gill rakers long, about 66% of eye diameter,
17-23 on first arch. Branchiostegals 6-8. Lateral line complete, with 70-
78 pored scales. Pyloric caeca 8-12, quite long.
Fins
D. 10-13. Dorsal adipose fin present, sickle shaped. A. 18-23.
P]. 10-12. Py. 8. Caudal fin forked. Pectoral and pelvic fins longer in
males than in females, pelvic fins of males may reach anus but always much
shorter than this in females. All fins with well developed breeding tuber-
cles in ripe males, these poorly developed or absent in females.
Sc ales
Cycloid, rather small. Well-developed breeding tubercles present on
adult males, but tubercles only poorly developed on females.
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Brown to dark bluish dorsally, fading to silvery white or while ventral-
ly. Fins immaculate, although pectorals and caudal often dusky.
Si ze
Maximum length about 225 mm (9 in) fork length (Taranetz, 1933), but
most not over 200 mm (8 in).
RANGE AND ABUNDANCE
The eulachon ranges from Bodega Head, California (Qdemar, 1964), north
along the coast of North America to Bristol Bay, Alaska and westward in the
Bering Sea to the Pribilof Islands. It is seasonally abundant in all the
streams utilized for spawning.
HABITS
The eulachon is anadromous, spending most of its life in the sea but re-
turning to freshwater streams to spawn. There is evidence, based on signifi-
cant differences in men's tic characters such as numbers of vertebrae, indi-
cating that eulachon return to their natal streams (Hart and McHugh, 1944),
The spawning run occurs in the spring, beginning in mid-March in the southern
part of the range and extending well into May at the northern end. Hales
predominate in the early part of the run, but may be equaled or exceeded by
females later. The fish apparently do not feed in fresh water (McHugh, 1939;
Hart and McHugh, 1944).
Spawning takes place over coarse sand and pea-sized gravel in water up
to about 7.6 m deep. Water temperature at spawning time is usually between
4.4 and 7.8°C (McHugh, 1940; Smith and Saalfeld, 1955). The eggs are about
0.8 - 1.0 mm in diameter and a female produces from about 17,000 to as many
as 60,000 eggs. Fecundity increases with the size and age of the individual.
Shortly after the egg is extruded, its outer membrane splits, separates from
the inner membrane and turns inside out, remaining attached to the inner
membrane at one point and thus forming a short stalk. The free edges of the
broken membrane are sticky and easily become attached to the bottom sub-
strate. Hatching occurs after 30-40 days of incubation at 4.4 - 7.2°C, but
in less than three weeks at 9.4 - 12.8°C. The larvae are transparent, only
4-5 mm long. They are weak swimmers, stay near the bottom of the stream and
are soon carried downstream to salt water. In the sea, the young fish become
distributed in the scattering layer of coastal waters, where food is
abundant. Food of the youngest fish seems to be mainly copepod larvae.
Later young, 25-50 mm long, have a more varied diet, including phytoplankton,
copepod eggs and adults, mysids, ostracods, barnacle larvae, cladocerans,
worm larvae and the smaller larvae of the eulachon itself (Hart, 1973, citing
unpublished studies by Barraclough). Juvenile eulachon feed heavily on
euphausiids, which are abundant in these waters. By mid-winter (December-
February) the young fish average about 66 mm (2.6 in)long and by the time
they are-a full year old they are about 80 mm (3.1 in) long. Subsequent
no
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growth is much slower. Two year old fish average not quite 90 mm, 3+ about
117 mm and 4+ about 154 mm in April. Adult eulachon in the sea feed on
plankton, mostly euphausiids and cumaceans (Hart and McHugh, 1944; Smith and
Saalfeld, 1955; McAllister, 1963; Barraclouqh, 1964).
Little else is known of the marine life of the eulachon. Occasional
commercial catches of this smelt in near-shore waters suggest that the fish
do not travel far offshore, but probably move about at no great distance from
the natal river.
Sexual maturity may be attained at the end of the second year of life
in some populations, not until the end of the third year in others. Most
smelt die after spawning, but some survive and may spawn a second time. The
spawning run from the sea begins whin river water temperatures rise to about
4.4°C, but the fish stop running if the temperature exceeds 7.8°C. Most of
the runs occur in the larger rivers, such as the Columbia and Fraser, al-
though smaller streams reaching the sea may also have eulachon population.
The spawning grounds may be in the major rivers themselves or in tributaries.
Some of the spawning areas in the Columbia River are above Vancouver, Wash-
ington, more than 160 km from the ocean.
IMPORTANCE TO MAN
In Washington and Oregon, commercial fisheries exist which take more
than 454,000 kg (1,000,000 lb) per year. A fishery of similar size formerly
existed in British Columbia, but landings at present are much smaller. Eula-
chon are important in the subsistence fisheries, and have been for many
years (Swan, 1881). Today, the native fishery in British Columbia probably
takes more fish than does the commercial fishery in that province (Scott and
Grossman, 1973).
The flesh of the eulachon contains a high percentage of oil which can be
rendered down to a fatty material with color and consistency-much like lard.
This fat was highly prized by the Indians of the Pacific Northwest. The fat
content is so high that, when dried, the fish can be burned directly or with
a wick threaded through it, hence the name "candle fish".
Today the eulachon is utilized principally as food, either for humans
or for commercially reared fur animals. The flavor is good and the eulachon
has a high place among gourmet food fishes. Indeed, it has been called
"unsurpassed by any fish whatsoever in delicacy of the flesh..." (Jordan and
Evermann, 1908).
The eulachon is also important indirectly. When eulachon are congregat-
ed off river mouths preparatory to beginning their spawning runs, they are
fed upon heavily by all sorts of predatory fishes, including halibut, cod
and salnon, as well as by such marine mammals as finback whales, porpoises,
sea lions and seals. The eulachon thus forms an important link in the food
chain between the small animals of the zooplankton and the large carnivores.
Ill
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RAINBOW SMELT
Osmerus mordax (Mitchill)
DISTINCTIVE CHARACTERS
One or more large canine teeth on the vomer (often missing in spawning
fish), 11-16 anal rays, and the front of the dorsal fin above or ahead of the
bases of the pelvic fins distinguish the rainbow smelt from other Alaskan
osmerids.
DESCRIPTION
Body elongate, compressed, slender, its depth about 12-191 of total
length. Head moderate, its length 16.5-23% of total length. Snout pointed,
elongate, longer than eye diameter. Eye large, round, its diameter 17.5-241
of head length. Mouth large, lower jaw longer than upper; maxilla reaches to
middle of eye or farther back. Teeth well developed, canine-like, present
on both jaws, tongue, vomer and palatines, the anterior teeth on tongue and
vomer enlarged. Gill rakers long, slender, 25-36 on first arch. Branchio-
stegals usually 7 on each side, rarely 5, 6 or 8 on one or both sides. Lat-
eral line incomplete, with 14-28 pored scales. Pyloric caeca 4-8.
Fins
dorsal
Caudal
Scales
D. 8-11, its origin ahead of base of pelvic fins. A small adipose
fin present.
fin forked.
A. 12-16. P-j. 11-14, shorter than head. P2.
8.
Cycloid, fairly large, thin and deciduous, 62-72 in a mid-lateral
series. Prominent nuptial tubercles present on scales of spawning males,
absent in females.
Color
Pale green to olive dorsally, silvery below, a bright, metallic silvery
band along sides. Sides often with purple, blue or pink iridescence. Fins
colorless, sometimes with faint speckling.
Size
Reported up to 340 mm TL in the Anadyr River of Siberia (Berg, 1948),
but North American smelt apparently only rarely exceed about 250 mm. The
average rainbow snelt in commercial catches is about 150 mm long with a
weight of about 30 grans.
RANGE AND ABUNDANCE
The range of the rainbow smelt along the Pacific coast of North America
is from Barkley Sound, Vancouver Island, north along the coast of British
112
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Columbia and Alaska through Bristol Bay, thence north to the Arctic coast and
east to Cape Bathurst. The species is also known from St. Lawrence Island.
On the Asian side, the rainbow smelt may be found from Korea and northern
Japan to the Anadyr Peninsula and westward to the White Sea. Rainbow smelt
are also native to the east coast of North America, from northern New Jersey
north to Hamilton Inlet on the northern coast of Labrador and westward in a
number of lakes and streams in Quebec and Ontario (Dymond, 1937; Bigelow and
Schroeder, 1963). References to rainbow smelt as far south as Virginia
appear to be in error (Low, 1896; Backus, 1957; Bigelow and Schroeder, 1963).
Smelt were introduced into Michigan waters from Maine, beginning in
1906, Early plantings were unsuccessful, but a plant of 16,400,000 eggs in
Crystal Lake, Benzie County, made in 1912, took hold and appears to be the
ancestors of most or all smelt in the western Great Lakes (Hankinson and
Hubbs, 1922; Creaser, 1926; Savage, 1935). Introductions into New York
waters were probably responsible for smelt in Lake Ontario, for they appeared
there some years before they were taken in Lake Erie (Mason, 1933).
Wherever it is present, excepting perhaps the most extreme ends of its
range, the rainbow smelt is abundant, although, because of its migratory
habits, the abundance is local and seasonal.
HABITS
Little, if anything, is known from direct studies of the rainbow smelt
of the Pacific coast. Most of the following account is derived from studies
of east coast and Great Lakes populations. Presumably the life history of
Pacific coast smelts does not differ greatly from this.
Like other osmerids, the rainbow smelt is a spring-time spawner. The
adult fish begin to congregate near stream mouths early in the spring, often
long before the ice goes out (Rupp, 1959). In most areas, the majority of
the fish are first-time spawners, usually two years old. These constitute
half to two thirds of the run, with three-year-olds another 25-30% and the
rest older fish (McKenzie, 1964). Movement into the streams begins when
water temperatures reach 2-4°C or higher. Warm days and cool nights seem to
enhance the spawning migrations (Hoover, 1936). As a rule, the upstream run
is short, a few miles to sometimes only a few hundred yards above the head of
the tide and some may even spawn in brackish water behind barrier beaches or
in the tidal zone of estuaries (Bigelow and Schroeder, 1963; McKenzie, 1964).
In the Yenisei River of Siberia, however, upstream migrations of more than
1,000 km have been observed (Berg, 1948), and to at least 320 km in the St.
Lawrence River in Canada (Magnin and Beaulieu, 1965).
The movement from lake or river to the spawning grounds is usually made
at night, although daytime spawning has been observed (Rupp, 1959). In most
populations, there appears to be an influx of fish just after dusk, followed
by a second group running around 2 a.m., (Creaser, 1926; Hoover, 1936; Baldwin,
1950). As a rule, the males appear'earliest, the females following an hour
or so later. Spawning seems to be initiated, at least in part, by the pres-
ence of the proper sex ratio in the group, which has been indicated as being
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no more than four males to one female (Hoover, 1936; Rupp, 1965). In the
spawning act, which takes place over sandy gravel, pebbles and rocks, the
fish crowd together, all headed upstream. They move laterally, in somewhat
exaggerated swimming movements. Contact between males either brings about no
reaction or a separation of the two fish, but when a male and female contact
each other, eggs and milt are released. It has also been reported that the
male pushes the female to the bottom or into shallow water after contact has
been made (Hoover, 1936), but -this action does not seem to be general.
Apparently the prominent nuptial tubercles on the sides of the males provide
a tactile stimulus which serves the individual in discriminating between the
sexes. Females also may have nuptial tubercles on the top of the head and
along the base of the dorsal fin, but they are poorly developed and are loca-
ted in positions where they could not function in the manner described
(Hoover, 1936; Richardson, 1942; Rupp, 1965).
In the spawning act, a female deposits only about 50 eggs at a time
(Hoover, 1936). Since the eggs are quite small (0.8-1.0 mm), a single female
produces a large number, from 1,700 to 69,600 (Langlois, 1935; Hoover, 1936;
Bert, 1948; Bailey, 1964; McKenzie, 1964). It thus requires many spawning
acts, spread over several hours each night for several nights, to deposit all
eggs.
Following each evening's activity, most of the fish drift, tailfirst,
downstream to the larger body of water whence they came to the spawning
grounds. Some, mostly males, may remain in the spawning stream during the
day, but they avoid light as much as possible.
The larger nembers of an age group, ripen earlier in .the season than do
smaller ones, and older fish ripen earlier than younger. Hence the spawning
season may extend over several weeks, or even months, with newly ripe fish
coming in as the spawned out fish leave. Because of this correlation of
spawning time with age and size, the average size of the fish tends to de-
crease as the run progresses.
Many of the spawned-out fish, especially males, die after spawning, but
those which survive will spawn again the following year. Sexual maturity is
achieved at the end of the second or third year of life. -Since a few fish
may live to six years, some obviously spawn several times during their lives
(Creaser, 1926; Bailey, 1964).
Spawning is not always confined-to streams, for rainbow smelt have been
observed breeding in shallows along lake shores. Here the fish come inshore
in small schools and swim about over rather restricted areas, apparently
without any distinct pairing. The composition of the schools changes con-
stantly, for individual fish apparently engage in spawning activities for
only 15-30 minutes at a time, then leave the school (Lievense, 1954; Rupp,
1965).
Whenever spawning takes place, the eggs settle quickly to the bottom.
Within 15-30 seconds of exposure to water, they become sticky and adhere to
whatever they touch. As with the eggs of other smelt, the outer cover
ruptures and turns inside out, adhering to the egg at one point, so that the
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egg is held above the substrate on a tiny pedestal. Hatching occurs in about
29 days at 6-7°C, 25 days at 7-8°C, 19 days at 9-10°C, 11 days at 12.2°C and
10 days at 15DC (Hoover, 1936; HcKenzie, 1964). Density of eggs on the
spawning grounds varies tremendously, according to the number of adults using
the area. Greatest production of larvae results when the density of eggs is
on the order of 100,000 - 130,000 per square meter. Under these conditions,
average survival to hatching is on the order of 0.5-2%. Major causes of mor-
tality are mechanical crushing, abrading, dislodging by waves and currents,
and fungus infections. However, year-class abundance is not related to the
number of larvae produced. Factors affecting the post-hatching survival of
the young apparently are more important than the number of fish hatched
(McKenzie, 1947, 1964; Rothschild, 1961; Rupp, 1965).
The young are about 5-6 mm long at hatching and are transparent. Being
weak swimmers, they are soon carried downstream to the lake or estuary. They
reach a length of about 40 mm in 3-4 months (Gordon, 1961; Bigelow and
Schroeder, 1963). Growth rates vary tremendously from one population to an-
other. However, averages, compiled from a number of sources, yield the
following approximate average lengths at various ages: 1+, 111 mm; 2+, 167
mm; 3+, 190 mm; 4+, 218 mm; 5+, 228 mm; and 6+, 242 nm. Females grow faster,
get bigger and live longer than do males (Greaser, 1926; Beckman, 1942; War-
fel et al, 1943; Baldwin, 1950; McKenzie, 1958; Bailey, 1964).
Except for the spawning runs, the rainbow smelt apparently does not
undertake definite migrations. Fish which go to sea stay within 8-10 km of
shore and probably do not stray far along the coast from the estuary (Bigelow
and Schroeder, 1963).
Although the rainbow smelt apparently returns to the river system in
which it was hatched, return to the spawning streams, both by first-time
spawners and by older fish, is often not precise. The degree of homing seems
to vary from one population to another and may be genetically controlled
(McKenzie, 1964; Rupp and Redmond, 1966).
Food of young-of-the-year smelt is mostly copepods and cladocerans, as
well as rotifers, eggs, and algae. Adults in saltwater feed on decapod and
mysid shrimps, copepods, amphipods and small fishes, as well as small shell-
fish of various sorts, crabs, squid and worms. The diet in fresh water in-
cludes the same major groups; copepods, amphipods, cladocerans, mysids and
small fishes, as well as various insects, especially Ephemeroptera and Dip-
tera nymphs and larvae. Feeding virtually ceases during spawning (Greaser,
1926, 1929; Kendall, 1927; Beckman, 1942; Baldwin, 1950; Gordon, 1961;
Bigelow and Schroeder, 1963).
IMPORTANCE TO MAN
In areas where it is sufficiently abundant, such as the Great Lakes re-
gion of the U. S. and Canada, and the New England states, the rainbow snelt
is an important part of both the sport and commercial fisheries. Since 1960,
the commercial catch by the U. S. and Canada in the Great Lakes and the
international lakes has been between 4,500,000 and 9,000,000 kg annually,
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averaging about 6,800,000 kg per year, with a value of about $0.07 per ka._
These fish are taken mainly by otter trawls, traps and gill nets. In the
New England states, landings of smelt reached 550,000 kg in 1889, but declin-
ed to a yearly average of only 215,250 kg in the period 1951-1954, and to an
average of only 69,700 kg yearly for 1969-1971. This decline is probably
caused in part by lessened abundance caused by obstructions in and pollution
of streams and in part by the high price commanded by New England smelts,
about ten times the price of Great Lakes fish (Bigelow and Schroeder, 1963;
Anonymous, 1972, 1973, 1974).
Great Lakes catches were formerly much greater, but an epidemic, ascrib-
ed to either bacterial or viral agents, decimated the population in 1943-
1944, causing an estimated loss to the commercial catch of nearly 23,000,000
kg for the 1943-1946 fishing seasons (Van Oosten, 1947). Other epidemic die-
offs, particularly in New England, have been ascribed to the microspridian
parasite Glugea hertwiqi (Haley, 1952),
The rainbow smelt is also a popular sport fish. It is taken by dip-net-
ting or with hook and line, the latter being especially popular in winter
through the ice. Sport catches in the U. S. are estimated to equal or exceed
the commercial landings.
As food, the rainbow smelt is highly esteemed, and has been ever since
white men came to North America. Its flesh is firm and tasty.
The rainbow smelt was not always considered an asset in lakes where it
was introduced, as it was thought to be a serious predator on and competitor
with various native fishes, and often fouled the nets of fishermen in the
days before it became marketable. However, there appears to be no evidence
that smelt in the Great Lakes have had any adverse effect on other species
(Gordon, 1961).
The rainbow smelt is too scarce in Alaska to be of any great importance,
although it undoubtedly enters subsistence fisheries from time to time.
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SECTION 7
MUDMINNOWS AND BLACKFISH--FAMILY UMBRIDAE
The five living species in this family, belonging to three genera, were
previously classified in two or three separate families. Current ichthyo-
logical thinking lumps them all together in one group. The family is strict-
ly northern hemisphere in its distribution. One species is found in eastern
Europe, two more are known from eastern and central North America, another
from northwestern Washington, and the fifth, Dal 1i a pectoralis, is found in
Alaska and western Siberia.
All members of the family are small, elongate fishes with rounded tails
and rather broad, flattened heads. They live in small, muddy ponds and quiet
streams with abundant vegetation. They are of no commercial value, but the
Alaska blackfish may be used for food for both dogs and people.
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ALASKA BLACKFISH
Dal 1ia pectoral Is Bean
DISTINCTIVE CHARACTERS
The short, flattened snout; dorsal and anal fins far back; pectoral fins
of about 33 rays and pelvic fin .of only 3 rays identify the Alaska blackfish.
DESCRIPTION
Body elongate, cylindrical anteriorly but compressed posteriorly. Depth
of body about 16% of total length. Head short, blunt, somewhat flattened,
about 20% of TL. Snout short, near 20% of head. Eye small, round, its dia-
meter usually somewhat less (751) than snout length. Mouth large, broad,
lower jaw protruding, maxillary reaches behind middle of eye. Small, sharp
teeth present on lower jaw, premaxilla, palatines and head of vomer. No
teeth on maxilla or tongue. Gill rakers 9-12, short. Branchiostegal rays
7-8 on each side. Lateral line with minute pores, 76-100 scales in mid-lat-
eral series. No pyloric caeca.
Fins
D. 10-14, located far back on body. A. 12-16, more or less under
dorsal. P-j. 32-36, edge of fin round. Po. usually 3, the fins, very small
and located just before anus. Caudal fin Broad, rounded.
Scales
Cycloid, small, more or less imbedded in skin.
Color
Dark green or brown above and on upper sides, pale below with dark speck-
les, four to six irregular dark bars or blotches on sides. Fins with dark
brownish speckling. Dorsal, anal and caudal with pale margins, these mar-
gins pink to red in spawning males.
Size
In most of its normal habitat, the blackfish rarely exceeds about 200 mm
in length. However, in the Yukon-Kuskokwim delta, fish up to about 255 mm
are not uncommon. In the area around Anchorage, Alaska, fish.up to 304 mm
and 366 g have been recorded (Trent and Kubik, 1974) and one specimen of
about 330 mm is known (Alaska Dept. Fish & Game, Anchorage).
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RANGE AND ABUNDANCE
The Alaska blackfish is found in lowland areas in eastern Siberia, St.
Matthew, St. Lawrence and Nunivak islands in the Bering Sea, and in Alaska.
On the mainland of Alaska, its natural range is from the Colville River delta
on the Arctic coast east and south to the central Alaska Peninsula near Chig-
nik. It is present in the Yukon-Tanana drainage as far upstream as Big Eldo-
rado Creek, near Fairbanks. Wherever it is present, it is usually quite
abundant.
The Alaska blackfish was introduced successfully to St. Paul Island in
the Pribilofs, unsuccessfully to Ontario. It was accidentally introduced
into Hood-Spenard lakes at the Anchorage airport in the early 1950's and has
thrived there. Subsequently, it spread via interconnecting waterways and
illicit transplants to a number of other lakes in that area, where it creates
a serious problem in the management of the sport fisheries for rainbow trout.
HABITS
Little is known of the biology of the Alaska blackfish. Spawning occurs
in the spring and summer, beginning soon after break-up in May and continuing
into July in the interior of Alaska, but apparently taking place only in late
July in the Bristol Bay area (Blackett, 1962- Aspinwall, 1965). Upstream
movement appears to coincide with a rise in water temperature to 10-15°C
(50-59°F) (Blackett, 1962). Spawning has not been observed, but the eggs,
about 1.5-2.3 mm in diameter, are probably deposited in vegetation at the
bottom of shallow ponds and quiet streams (Nelson, 1887; personal observa-
tions). Females deposit from about 40 to about 300 eggs, the number increas-
ing with the size of the fish. Spawning of a particular female probably goes
on over a period of several days, possibly longer, with only a few eggs being
extruded at each spawning act. At any rate, partially spawned females have
been found during most of the spawning season. Nearly ripe ovarian eggs are
yellow and opaque, but are not capable of being fertilized. Fully ripe eggs
are nearly transparent. The fertilized eggs are demersal and extremely
sticky. Females normally contain two sets of eggs, the large, mature ones
about 2 mm in diameter, and a larger number of small "recruitment" egqs,
which average less than 1 mm in diameter. These are presumably the eggs
which will be spawned the following year.
Development to hatching requires about 10 days at 12-13°C (ca 54°F) under
experimental conditions. The young are about 5.7 mm long at hatching and
have a large yolk sac. By the tenth day post-hatching, the young are about
9 mm long and the yolk sac has virtually disappeared. By the 22nd day, the
little fish are about 12 mm long and are beginning to take on the character-
istics of the adults. At 20-21 mm, reached in about 44 days, metamorphosis
is virtaully complete (Aspinwall, 1965). Subsequent growth in the first
year of life is fairly rapid, but later slows. In interior Alaska and the
Anchorage area, the blackfish reach about 64 (50-95) mm by age 1, 108 (76-
146) mm at age 2, 138 (115-160) mm at age 3 and 178 (155-200) mm at 4. By
contrast, fish from Lake Aleknagik in the Bristol Bay area grow very slowly.
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A two year fish was only 36 mm long. A three was 50 mm, a four 63 Tim, and
one of 8 years was only 135 mm long. Sexual maturity is reached at 2 or 3
years of age, so some females probably spawn several times during their
lives (Blackett, 1962; Aspinwall, 1965; Chlupach, 1975).
The blackfish does not undertake extensive movements, as far as is known.
Its migrations appear to be limited to inshore or upstream movements to
spawning grounds in the spring of the year, and (presumably) reverse migra-
tions to deeper water in the fall.
Blackfish feed almost exclusively on small invertebrates. The smaller
individuals subsist mostly on copepods and Cladocera, shifting, as they grow
larger, to insects and small fishes. Hemiptera are especially important, but
the diet also includes Diptera larvae, Trichoptera larvae, Ephemeroptera,
Odonata, ostracods, molluscs, annelids and even algae (Ostdiek and Nardone,
1959; Chlupach, 1975). They also eat smaller members of their own kind.
Because of their hardiness* their ability to tolerate crowding and low oxy-
gen, and their diet, Alaska blackfish make tough competitors for rainbow
trout wherever the two co-exist.
Blackfish are renowned for their cold tolerance, but despite the old
story of frozen fish being eaten by dogs, thawed by the heat of the dogs'
stomach and then vomited up alive (Turner, 1886), they cannot withstand
complete freezing. Fish have survived exposure to -20°C for up to 40 min-
utes, and can survive for a few days after conplete freezing of parts of the
body, even the head. However, complete freezing results in death (Borodin,
1934; Scholander et al, 1953). Blackfish can also withstand complete anoxia
for up to 24 hours if the temperature is 0°C (Bonnet, unpublished research).'
IMPORTANCE TO MAN
The Alaska blackfish was formerly a most important source of food for the
native peoples and their dogs. In the early 1880's, it was estimated that
not less than 103.5 tons were taken annually in the Yukon-Kuskokwim delta
alone, with 69 tons taken from October through December. Total harvest all
along the coast was estimated at not less than 155 tons, and it was felt that
double this amount might be a more realistic estimate (Nelson, 1884). Nowa-
days, however, such use has decreased greatly, although unknown quantities
of blackfish are still utilized in some of the more remote villages.
The Alaska blackfish makes an interesting aquarium fish, but it shoud be
noted that possession or export is prohibited by law unless a permit is
secured.
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SECTION 8
PIKES—FAMILY ESOCIDAE
The pikes are elongate, somewhat compressed fishes, with elongate, flat-
tened snouts and large, sharp teeth. Five species in one genus are distribu-
ted in the northern hemisphere. One species is confined to Siberia, three
are endemic to eastern North America, and the fifth, the northern pike,
Esox luclus, is of circumpolar distribution.
Pikes are carnivores, their diets consisting mainly of other fishes.
They are classified as qame fishes in North America, although the smaller
members of the family might not be so considered by some people. They are
quite edible when properly prepared, and commercial fisheries for northern
pike are important in the larger lakes of Canada and the United States. The
muskellunge reaches the largest size of any of the six species, with an an-
gling record of 31.8 kg (69 Ib 15 oz). Next largest is the northern pike.
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NORTHERN PIKE
DISTINCTIVE CHARACTERS
Esox lucius Linnaeus
The pike is easily recognized by its long, flat, "duck-bill" snout;
large mouth with many large, sharp teeth; and the rearward position of the
dorsal and anal fins.
DESCRIPTION
Body elongate, slender, moderately compressed, its depth 10-17% of total
length. Head long, 25% or more of total length. Snout elongate, flattened,
40-47% of head length. Eye round, large, near middle of head length and
close to dorsal surface. Mouth large, maxilla reaches to eye, lower jaw
often projects beyond upper. Usually five large sensory pores on each side
of lower surface of lower jaw. Large sharp canine teeth on lower jaw, head
of vomer and inner edge of palatines. Smaller sharp, curved teeth on pre-
maxilla, tongue, vomer and palatines, as well as on basi-/-and pharynqo-
branchials behind tongue. Gill rakers present only as patches of sharp teeth
on gill arches. Branchiostegals 14 or 15 on each side. Lateral line with
55-65 pored scales, these scales notched posteriorly. Scales rows along
middle of sides 105-148. No pyloric caeca.
Fins
D. 17-25 soft rays, fin located far posterior. A. 14-22, located under
and arising a little behind dorsal fin. P] . 14-17, low on
under opercle. ?2- 10-11 s low on body, about at middle of
Both PI and Pg rounded, paddle-shaped. Caudal fin slightly
Scales
body, base arisinq
total length.
forked.
body.
Color
Cycloid, moderately small, with numerous pitted scales scattered over
In adults, back and sides dark grayish-green to green or dark brownish,
sides with numerous yellow spots arranged in irregular longitudinal rows.
Scales usually with a tiny gold spot at edge. Belly and ventral surface of
head creamy white. Dorsal, anal and caudal fins green to yellowish, some-
times more or less orange or reddish, marked with dark blotches. Pectoral
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and pelvic fins dusky. Head dark green above, pale below, an inconsmcuous
dark line running below eye.
Color of young pike is similar to that of adults except that the sides
are marked with irregular pale vertical bars instead of spots, and the eye-
bar is usually more conspicuous.
A color variant is the so-called "silver pike". Here the color is gray-
ish green or deep blue on the back through silvery gray on the sides to
creamy white below. There are no spots or bars on the sides, although pale
marks may be present on the caudal peduncle. The fins are marked with small
dark spots concentrated along the rays. This color variant is a genetic
mutant that breeds true. It was first reported from the area near Sharbot
Lake, Ontario (Prince, 1898), and subsequently has been found in north-cent-
ral United States; in Ontario, Manitoba and Mackenzie in Canada; Sweden; and
Alaska. It is said to be much hardier than the ordinary form (Eddy and Sur-
ber, 1947; Lawler, I960, 1965b; Bartholomew et al, 1962).
Si ze
The northern pike in North America reaches at least 133 cm (4 ft) in
length and 22.3 kq (49 Ib) in weight. The angling record is a specimen of
21 kg (46 Ib 2 oz) and 133 cm (52.5 in) length from Sacandaga Reservoir, N.Y.
A pike said to have weighed "about 45 pounds" (20.5 kg) was taken near
Circle, Alaska, in the early 1960's. European pike apparently get biqger
than those of North America. There are authentic records of pike up to 26 kg
(57 Ib 3 oz) from Ireland and Scandinavia (Scott,and Grossman, 1973) and an
apparently authentic record of a 34 kg (74 Ib 13 oz) fish from Lake II'men'
in Russia (Berg, 1948).
In most areas, the average pike caught by anglers will run between 1 and
3 kg (2-6.6 Ib) weight. A fish of over 7 kg (15 Ib) is a very good one and
one of more than 11 kg (24 Ib) would be considered rather remarkable.
RANGE AND ABUNDANCE
The northern pike is circumpolar in fresh water. It is present widely
in Europe as far south as Spain and northern Italy, with an east-west distri-
bution from the British Isles to the Pacific shores of Siberia. In North
America, the pike ranges from western Alaska to eastern Canada (though absent
from the Keewatin Peninsula) and from the Arctic coast south to Nebraska,
Missouri, southern New England. Except in the Ahrnklin River drainage in
Alaska and the headwaters of the Alsek and Taku rivers in Yukon Territory
and British Columbia, pike do not occur naturally west of the continental
divide. Pike have been introduced into a number of places in the United
States, and the introductions are a nuisance or a blessing, according to
one's point of view. Recent illegal transplants by private individuals have
placed pike in the Susitna River drainage in Alaska.
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HABITS
Pike spend the winter in relatively deep water in lakes and rivers. With
the approach of spring, they begin to move inshore or uostream to the marsh
areas where they spawn. This movement usually occurs soon after the ice
goes out (Clark, 1950), but may start even before that in far northern areas
(Cheney, 1972). Most of this movement to the spawning grounds takes place at
night (Franklin and Smith, 1963). Spawning does not begin until water temp-
eratures in the shallows reach about 6-9°C (43-48°F) (Bennett, 1948; Clark',
1950, Cheney, 1971).
The spawning grounds are marshy areas, with shallow water, emergent vege-
tation and a mud bottom covered with a vegetation mat. Suitable vegetation
and quiet water seem to be the most important factors in the choice of spawn-
ing areas. Actual spawning occurs in water less than 51 cm (20 in) deep,
with most activity in water of less than half that depth (Clark, 1950). A
male (sometimes two or three males) courts a female by pushing with his snout
against her head and pectoral region. If the female is not yet ripe or is
spawned out she repels the male by turning her head to one side and straight-
ening with a jerk. She may also adopt the threat posture, with branchio-
stegals lowered, back arched, paired fins spread downward. Although pike are
not territorial or monogamous, a male attending a female may threaten other
males with this same posture.
If the female is receptive, the two fish swim about side by side, appar-
ently oriented eye to eye. At mating, both fish lower the branchiostega'ls
and swimming speed increases. The male flaps his caudal fin towards the fe-
male, then swings the caudal sharply away from her and brakes with his paired
fins. This thrusts his vent close to that of the female, and he ejects milt.
At the .same time, the female erects her pelvic fins, thrusts her caudal fin
towards the male, and, by powerful contractions of her abdominal muscles,
ejects some eggs. The final return of the caudal fins to normal position
mixes eggs and sperm, and scatters the eggs. Only a few eggs, 5 to 60, are
released at a time. The spawning act is repeated every few minutes for up
to several .hours, after which the fish rest for some time before resuming
spawning. During this resting period, both male and female may take new
mates, or they may continue together for several days until the female's eggs
are all extruded. Spawning occurs only during daylight, and reduced light,
as from cloud .cover or ripples on the water reduces spawning activity.
Excessively cold nights have the same effect (Svardson, 1949; Clark, 1950;
Fabricius and Gustafson, 1958). The spawned out adults may stay on the
spawning grounds for as long as 3 1/2 months, but most leave within 6 weeks.
The eggs are 2.5-3.0 mm in diameter, tan to yellow in color. Egg number
increases with the size of the fish, from as few as 2,000 in fish of about
33 cm (14 in) to nearly 600,000 in a fish weighing 14.5 kg. Ripe females
contain both large, ripe eggs and minute, immature eggs which will ripen the
following year. Thus,'new eggs continue to develop during the life of the
individual. Some of the immature eggs are resorbed between spawning periods,
as are all of the residual mature eggs (Carbine, 1944). Fertilization is
usually highly efficient. After fertilization, the eggs settle to rest on
124
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the weeds or the bottom. Development time to hatching varies inversely with
temperature. At 6°C, 23-29 days are required, but at 18°C, only 4-5 days.
Mortality rates of the eggs tend to increase with increasing temperatures
(Swift, 1965). The newly hatched larvae are 6.5-9.3 mm long and have a yolk
sac but no mouth. The mouth develops at 10.5-12 mm, by which time the yolk
sac is absorbed. Active feeding begins when the young fish are 11-13 mm
long about 10 days after hatching (Franklin and Smith, 1960). Early foods
are various entomostracan zooplankters, especially copepods and cladocerans.
As the little fish grow, they shift to insect larvae and nymphs, mainly
Tendepedidae, then to fish and other small vertebrates. By the tine they
are about 50 mm (2 in) long, the little pike are feeding almost exclusively
on fish (Hunt Carbine, 1951; Mateeva, 1955; Franklin and Smith, 1970, 1963).
When the young have reached a length of about 20 mm, they begin to move
out of the marshes. Movement is in the daytime and is positively correlated
with light intensity. Mortality from the fertilized egg to migrant fry is
heavy, as much as 99.91. Competition for food, predation and cannibalism
are the most important factors here. Water quality also may be important,
as the fry are rather sensitive to extremes of pH and the concentrations of
carbonate and bicarbonate (Hunt and Carbine, 1951; McCarraher, 1962; Franklin
and Smith, 1963).
Growth of the young is rapid, about 0.5 mm per day for the first 20 days,
about 2 mm per day for the next 30 days, and about 1 mm per day for the next
40 days. The fastest growers are those which first make the shift to a fish
diet. These young are especially likely to become cannibals. One fish of
only 23 mm had eaten another young pike of 16 mm (Hunt and Carbine, 1951).
The fastest growing fish may achieve a length of 446 mm (17.5 in) and a
weight of 460 grams (1 Ib) by mid-October. This represents growth in length
at the rate of 2.6 mm per day from hatching, and in weight of 2.69 grams per
day (Carbine, 1945).
Growth remains rapid in the first year, although it depends on tempera-
ture and the availability of food. Subsequently, as with most fishes, the
growth rate slows (Rawson, 1932; Van Engel, 1940; Lagler, 1956; McCarraher,
1959; Cheney, 1972). Pike appear to require less food for maintenance than
do most fishes and are therefore able to convert a relatively large portion
of their ration into growth. After maintenance requirements have been met,
1 gram of food will produce 0.437 grams of pike, a ratio of 2.29:1. Peak
maintenance requirements occur in late spring, then decline and remain low
through the winter, rising again in the spring (Johnson, 1966). This cycle
appears to be related to the energy requirements of reproduction.
Growth rates tend to be faster in the warmer, southern parts of the
range than they are in the far north, but northern fish live longer. How-
ever, even within a relatively small area, growth is highly variable from
one body of water to another. The average size of fish of a given age may
vary by a factor of 2 or even more, and the difference between the largest
and smallest may represent almost a full order of magnitude. In the Minto
Flats area of interior Alaska, pike reach average lengths of 140 mm (5.5 in)
by September of their first year of life, 186 mm (7.3 in) as 1 year olds,
287 mm (11.3 in) at 2 years, 555 mm (21.9 in) at 5 years, 801 mm (31.5 in)
125
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at 10 years, and 990 mm (39 in) at 21 years. These growth rates are rather
faster than those found in Great Bear, Great Slave, Lesser Slave, and Atha-
basca lakes in northern Canada, or in a number of small lakes in the upper
Tanana River drainage in Alaska (Miller and Kennedy, 1948b; Cheney, 1972).
Adult pike feed almost exclusively on fish, the kinds eaten depending
largely on what is available. The variety of species eaten is tremendous
and often includes smaller pike. Other organisms included in the diet in-
clude water fowl, frogs, small mammals such as mice and shrews, crayfish and
insects. In Alaska, coregonids appear to be the major food item, followed by
by small pike, blackfish, burbot and suckers, as well as insects, especially
Odonata naiads. In general, pike seem to eat whatever they can catch and
swallow (Lagler, 1956; Lawler, 1965b; Cheney, 1971). Digestion of fish food
is fairly rapid at usual summer temperature, with digestion 50% complete in
20 hours. Birds, on the other hand, require a much longer time, almost 130
hours for 50% digestion, due, no doubt, to the protection provided by the
feathers {Solman, 1945; Seaburg and Moyle, 1964). Pike are believed to be
serious predators of young waterfowl. This is no doubt true in some places,
but not in others. In the deltas of the Saskatchewan and Athabasca rivers
in Canada, pike were estimated to eat nearly 101 of the young ducklings, but
in the Seney National Wildlife Refuge in Michigan, only 0.21 of the pike
stomachs examined contained waterfowl (Solman, 1945; Lagler, 1956).
Pike do not,' as a rule, undertake long migrations, although occasional
individuals may move considerable distances. In the Minto Flats, in interior
Alaska, 36% of the fish tagged moved more than 10 miles during the summers.
The Minto Flats lakes become anoxic in winter and the pike must travel con-
siderable distances to find suitable wintering areas. One fish moved 288 km
(180 mi) downstream in 10 months (Cheney, 1971).
Different populations of pike often show statistically significant
differences in such things as the numbers of fin rays, lateral line pores
and vertebrae. These differences may indicate isolation in separate glacial
refuges. However, since such differences have been found in lakes only 32
kin (20 miles} apart and draining into the same river, even interpretation
is open to question. There is certainly no strong evidence to indicate sub-
speciation (Morrow, 1964; McPhail and Lindsey, 1970; Cheney, 1972).
IMPORTANCE TO MAN
The pike enjoys a most varied reputation. In some areas it is consider-
ed a nuisance, scarcely worthy of the name fish, but in other regions it is
highly regarded and attracts large numbers of anglers and dollars. Pike
will take a large variety of lures and live baits. They put up a strong
fight at first, but it is usually of fairly short duration. They seldom
jump, but will often rear out of the water, shaking their heads and all too
frequently throwing the hook. Much of the reason behind the pike's poor
reputation as a sport fish is too heavy gear. Taken on light tackle, pike
give very good accounts of themselves. A short wire leader-is a necessity,
as the pike's sharp teeth are all too liable to cut a line.
126'
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On the commercial side, the pike is unimportant in the United States.
Landings from 1960 through 1970, almost entirely from the Great Lakes, aver-
aged only a bit more than 79,500 kg (175,000 1b) per year. On the other
hand, the Canadian catch runs on the order of 2,273,000 kg (5,000,000 Ib) per
year, much of which is exported to the United States. Pike are usually sold
fresh, generally in the round or gutted and beheaded.
For eating purposes, pike are delicious. The meat is white, flakey and
flavorful. The so-called "muddy" taste of which some people complain is
confined to the skin. If the fish be thoroughly scaled, scrubbed and washed,
the muddy taste is eliminated. But it is better and easier to skin the fish.
Another complaint against pike is its boniness. The dorsal ribs or Y-
bones are only lightly fastened to the backbone and tend to come loose in the
meat. However, with a very sharp knife and a little practice, it is easy to
cut boneless filets of pike and thus eliminate the problem.
Pike are often used as farm-pond fish, particularly in combination with
blue gills. They seem able to crop the forage fish at such, a rate that the
latter cannot overpopulate and become stunted.
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SECTION 9
MINNOWS—FAMILY CYPRINIDAE
This is the largest of all families of fishes, including some 275 genera
and more than 1500 species. They are virtually world-wide in their distri-
bution, being absent only from Australia, South America and Greenland. They
are of variable shape, but generally are moderately elongate. They have no
spines in the fins, although some forms, such as carp, may have the first
one or two rays of the dorsal and/or anal fin stiffened and spine-like. Sex-
ual dimorphism is often marked in the spawning season, when the males may
develop breeding tubercles and/or bright colors. The larger species, such as
carp, are used for human food in some parts of the world. However, most mem-
bers of the family are small and find their chief value as forage for larger
fishes.
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LAKE CHUB
Co ties jus plumbeus (Agassiz)
DISTINCTIVE CHARACTERS
The spineless fins, normal jaws, and tiny barbet at the corner of the
mouth distinguish the lake chub from other Alaskan fishes.
DESCRIPTION
Body elongate, slender. Depth 14-2111 of total length. Head short, about
19-22% of total length, relatively longest in young fish. Snout moderate,
rounded, a little shorter than eye diameter. Eye round, about 25% (20-33%)
of head. Mouth fairly small, slightly oblique, upper jaw usually extending
slightly beyond lower anteriorly and reaching posteriorly about to anterior
edge of eye. A small but well developed barbel near posterior end of maxil-
la. No teeth in jaws, but pharyngeal (throat) teeth well developed, usually
2, 4-4, 2, but variable. Gill rakers short, 4-9 on first arch. Branchio-
stegals 3 on each side. Lateral line with 53-79 pores, usually 56-69.
Scales
Cycloid, small, those on back often smaller than those on sides. Breed-
ing adults develop nuptial tubercles on the head and along the back of the
dorsal fin as well as on the \ipper parts of the pectoral fins and on the
breast. These tubercles are better developed in males than in females.
Color
Brown to greenish above, silvery below, A rather indistinct dark or
lead-colored band along sides, often extending forward on to head in small
specimens. Lower sides and belly often with fine dots of dark pigment. In
some populations, breeding males develop bright orange-red patches on sides
of head and at bases of pectoral fins (Richardson, 1944), but the presence of
this color varies from place to place. So far, orange patches have not been
found in Alaskan lake chubs.
Size
Known to reach 227 mm (8.94 in) total length in northern Quebec (Scott
and Grossman, 1973), but most are 50-100 mm (2-4 in) long.
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RANGE AND ABUNDANCE
The lake chub is strictly North American. It ranges from interior Alaska
east to Nova Scotia and from the Arctic coast at the McKenzie River delta and
Ungava Bay south to New England and the Great Lakes, except Lake Erie. Scat-
tered, isolated populations are present in Iowa, Nebraska, Wyoming, Colorado,
Montana and the Dakotas. In Alaska, the lake chub seems to be confined to
the Yukon-Koyokuk-Tanana-Porcupine drainage, from about Nulato on upstream.
Wherever it is present at all, the lake chub is usually one of the more abun-
dant fishes.
HABITS
Despite the fact that the lake chub is wide-spread and common, very
little is known about it. Spawning apparently occurs in the spring and sum-
mer, as early as April in the southern part of its range and as late as Aug-
ust in the far north (McPhail and Lindsey, 1970; Scott and Grossman, 1973).
Spawning occurs in shallow water over rocky or gravelly bottoms in rivers and
smaller streams (Allin, 1953; McPhail and Lindsey, 1970), the fish moving out
of lakes or deeper parts of streams in large schools for this purpose. No
nest or redd is built, nor do the adults guard the eggs (McPhail and Lindsey,
1970). In Lake Superior, the lake chub occasionally hybridizes with the
longnose dace, Rh i n ichthys cataractae (Hubbs and Lagler, 1949). The eggs are
yellowish in color. A female of 70 mm length was estimated to contain about
500 eggs (Richardson, 1935). Nothing has been reported of spawning behavior
or rate of development of eggs and young. In Pyramid Lake, Alberta, one
year old chubs averaged about 28 mm long, 2-year-olds 48 mm, 3-year-olds 71
mm, and 4-year-olds 114 mm (Rawson and Elsey, 1950). Lake chubs are thought
to mature at 3 or 4 years of age, seldom surviving beyond 5 years, in central
British Columbia. Females apparently grow faster and live longer than do
males (McPhail and Lindsey, 1970).
Young lake chub feed primarily on zooplankton, chiefly cladocerans and
copepods, while older fish eat mostly insect larvae and algae. Large chubs
may also take small fishes (Simon, 1946; Rawson and Elsey, 1950; McPhail and
Lindsey, 1970; Scott and Crossman, 1973).
, The lake chub inhabits all sorts of waters - lakes, clear streams, heavi-
ly silted rivers. In general, it seems to be nost common in fairly shallow
water, but may move into deeper parts of lakes during hot weather. In some
areas, as Great Bear Lake, NWT, it is found only in streams, in others it
seems to utilize stream habitats only for spawning, while in still other
places it inhabits streams almost exclusively (Hubfas and Lagler, 1949; Allin,
1953; Personius and Eddy, 1955; Johnson, 1975). In interior Alaska, in the
Tanana and upper Yukon drainages, it is much more common in the heavily-
silted main rivers than in the clear tributaries (personal observation).
The lake chub has been classified as a sight feeder, since it has large
optic lobes and relatively few taste buds (Davis and Miller, 1967). However,
130
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since it is so common
nil, other mechanisms
IMPORTANCE TO MAN
in heavily-silted rivers, where visibility is virtually
for finding food must also be present.
The lake chub is of little direct importance. In some regions, anglers
said to be a nuisance in some trout
readily and "it is almost impossible to
these minnows" (Allin, 1953). Lake chub,
and abundance, are probably an important
use it extensively for bait. It is
fisheries, for it takes bait or fly
make a cast without catching one of
in view of their diet, distribution
forage species, but confirmatory data are lacking.
In Ontario streams, the lake chub is sometimes taken and eaten by smelt
fishermen who do not recognize it (Scott and Grossman, 1973). It has been
cold-smoked, with satisfactory results (Lantz, 1962).
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SECTION 10
SUCKERS—FAMILY CATOSTOMIDAE
These fishes are elongate, mostly subcylindrical, with a ventrally-
placed, protrusible mouth and no teeth in the jaws. They feed by swimming
slowly along the bottom, sucking up the bottom debris, including therein the
small insect larvae and other invertebrates that live on or in the bottom.
Some of the suckers may reach considerable size, up to several kg.
There are about 65 species of suckers, all but two of them found only in
North America. One of the remaining species is endemic to southern China,
while the other, the longnose sucker of North America, is also present in
eastern Siberia.
Suckers are generally considered more of a nuisance than anything else,
especially by trout fishermen. They are popularly supposed to do great dam-
age to the eggs of other fishes, but the supposed extent of this damage is
probably greatly exaggerated. The larger species,.especially the longnose
sucker and white sucker, are taken commercially in the Great Lakes and the
large Canadian lakes and are marketed as "mullet". Suckers make excellent
bait for large game fish such as pike and bass.
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LONGNOSE SUCKER
Catostomus cjttostotnus (Forster)
DISTINCTIVE CHARACTERS
The ventrally-placed sucking mouth with thick, papillose lips distin-
guishes the longnose sucker from all other Alaskan fishes.
DISCRETION
Body elongate, nearly cylindrical. Depth of body greatest somewhat
forward of dorsal fin, 14-19% of TL. Head long, about 20% of TL, top of
head rounded and seal el ess. Snout long, 38-481 of head length, ending ant.eri-
orly in a rounded point. Eye round, small, its diameter about 22-31% of
snout length. Mouth ventrally located, behind tip of snout, protrusible.
Lips, especially upper, thick and papillose. No teeth in jaws, Pharyngeal
teeth numerous, comb-like, in single rows. Gill rakers short, 23-30 on
first arch. Branchiostegals 3 on each side. No proper, well-differentiated
stomach, no pyloric caeca. Lateral line complete, inconspicuous, with 90-120
pored scales.
Fins
D. 9-11. A. 7, Pr 16-18. P2. 10-11. Caudal moderate-ly forked, the
tips slightly rounded.
Scales
Cycloid, small.
Color
Adults may be, reddish brown, dark brassy green or gray to black above,
paler on the lower sides, the ventral parts white. Young fish are usually
dark gray with mottling of paler color on the back. Young of the year are
gray with small black spots. Breeding males are usually dark above with a
brilliant reddish stripe along each side, while females are greenish gold to
copper, with a less brilliant red stripe. The breeding males show prominent
tubercles on the rays of the anal and caudal fins, and also on the head.
Size
The longnose sucker is reported to reach a length of 643 mm (25 in) and
a weight of 3.3 kg (7.26 Ibs) (Keleher, 1961), but the usual run of fish are
much smaller.
, - 133
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RANGE AND ABUNDANCE
The longnose sucker ranges from New England to Labrador in the east,
westward through the Great Lakes, the northern part of the Mississippi-Mis-
souri system and the Columbia to the west coast. It reaches the Arctic Ocean
in northeastern Labrador, but is absent from the arctic coast of Canada from
Ungava to the Horton River. Present throughout the rest of mainland Canada
(except Nova Scotia and eastern Labrador) and Alaska, but absent from the
islands along the Pacific and Bering coasts and from the arctic islands.
Found also in eastern Siberia from the Yana to the Anadyr rivers.
In North America, the longnose sucker is abundant throughout the north-
ern part of its range, especially in the northwest.
HABITS
Breeding occurs in the spring, as early as May in the southern part of
the range, as late as July in the far north. Spawning runs begin after water
temperatures have reached 5°C, with greatest intensity occurring at tempera-
tures above 10°C (Rawson and Elsey, 1950; Brown and Graham, 1954; Geen, et al,
1966). The fish move from lakes into inlet streams or from slow, deep pools
into shallow, gravel-bottomed portions of streams. Spawning occurs only dur-
ing daylight Hours most commonly over gravel of 0.5-10 cm diameter, in shal-
low water 10-60 cm deep with a current of 30-45 cm/sec (Geen et al, 1966).
At spawning time, the males lie close to the bottom in the current of the
spawning area and show no aggressive behavior, while the females stay along
the banks and in still water. In the breeding act, a female swims out to
the males, 2-4 of whom escort and crowd around her. Dorsal fins erect, the
males clasp the female with their pelvic fins and vibrate their anal fins
against her. There is considerable thrashing about for a few seconds during
which eggs and sperm are released. No nest is built. After a spawning, the
female returns to quiet water. She may spawn many times in an hour, with the
same or different males. A single female may produce up to 60,300 eggs
(Harris, 1962; Geen et al, 1966). Fish which have moved out of a lake to
spawn generally return to the lake a few days after spawning is completed.
However, river-resident fish may stay on or near the spawning area for much
of the summer.
Post-spawning mortality of adults is on the order of 10-30%. Many fish
spawn in two or even three consecutive years, while others may skip one or
two years between spawnings (Geen et al, 1966).
The eggs are about 3 mm in diameter, yellow in color, adhesive and demer-
sal, sinking to the bottom and lodging in crevices in the gravel. Develop-
ment to hatching takes one to two weeks, according to temperature. The young
are about 8 mm long at hatching and remain in the gravel for one to two
weeks. In some areas, a nocturnal, downstream movement of fry begins as soon
as the fish emerge from the gravel. However, in interior Alaska at least
some of the young stay in the streams all summer.
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Growth rates vary tremendously from place to place, apparently correlated
more with food supply than with temperature. In Great Slave Lake, young of
the year may reach 80 mm FL by the end of Augusts but in Yellowstone Lake
they are less than half that length (Brown and Graham, 1954; Harris, 1962).
Similar variations in growth rates are continued through later years of life.
Even in a single population, the largest fish of a given age may be twice as
long as the smallest. Age determinations by means of scales, which often
underestimate the true age (Geen et al, 1966), have given the following age-
length relationship for suckers from the south end of Great Slave Lake, one
of the faster growing populations recorded (Harris, 1962): 2 yrs, 179 mm;
5 yrs, 306 mm; 10 yrs, 496 mm; 15 yrs, 557 mm.
Age at maturity increases with latitude. Sexual maturity is achieved at
2 years by males, 3 years by females in Colorado, 4 and 5 years in Yellow-
stone Lake, but not until 9 or 10 years of age in Great Slave Lake. The old-
est fish on record is one of 19, years from the northern part of Great Slave
Lake (Brown and Graham, 1954; Hayes, 1956; Harris, 1962).
Except for movement to and from the spawning areas, the longnose sucker
apparently does not undertake any definite migrations. During the summer,
fish in interior Alaskan streams appear to wander more or less at random,
some going upstream, some down. The general trend of movement is down
stream, however, so that by October no suckers remain the the upper reaches
or on the spawning areas.
The longnose sucker feeds almost entirely on material found on the bottom.
The fish swim slowly along, their protrusible lips touching the bottom in a
way which suggests that the papillae have a sensory function. Food is suck-
ed into the mouth and swallowed. In streams, major food items of the adults
include algae and other plants, Diptera, Ephemeroptera, Trichoptera, Cole-
optera, spiders, molluscs. Lake-dwelling adults feed mainly on various
crustaceans, especially cladocerans and amphipods, as well as insect larvae
and nymphs. Diptera and Ephemeroptera are the most important of these.
Occasionally, large adult suckers will feed upside down on terrestrial
insects floating on the surface of eddies. The very young fish seem to feed
chiefly on cladocera and insects.
The longnose sucker is reputed to destroy the spawn of other fishes,
especially trout. Undoubtedly, fish eggs are eaten when available, but, as
far as trout eggs are concerned, these are undoubtedly "floaters", which
were washed out of the redd or were dug up by subsequent spawners (Stenton,
1951).
IMPORTANCE TO MAN
Utilization by man of the longnose sucker ranges from zero to extensive.
In many areas, it is a major source of dog food. In the Great Lakes and the
Canadian lakes, considerable amounts are landed and marketed, together with
other suckers, as "mullet". The flesh is said to be firm, white, flaky and
sweet, although bony. This may be true of fish taken in the winter, but the
flesh of spawning fish is soft and glutinous and has an unpleasant taste.
135
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Because of the similarity in food habits, longnose suckers are important
competitors with trout and other desirable sport fish. An intensive gill
netting program to remove longnose suckers from Pyramid Lake, Alberta, re-
sulted only in improved growth of young suckers and did not improve angling
success (Rawson and Elsey, 1950).
136
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SECTION 11
TROUT-PERCHES—FAMILY PERCOPSIDAE
This family, which includes only two species, gets its name because
the fish have some of the characteristics of the trouts and some of the
characteristics of the perches. They have, for example, a dorsal adipose
fin and abdominally placed pelvic fins, like the trouts; but there are one or
two spines in the dorsal, anal and pelvic fins and the scales are ctenoid,
like the perches.
The trout-perches are known only from North America. One species, the
sand roller, is found in Washington, Oregon and Idaho. The other, the trout-
perch is wide-spread over much of the northern U. S. and Canada.
Trout-perches are important forage species for larger fishes.
137
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TROUT-PERCH
Percopsis om1_s_cgmaycus (Walbaum)
DISTINCTIVE CHARACTERS
The combination of a dorsal adipose fin; small, weak spines in the dorsal
and anal fins; rough, ctenoid scales; and pectoral fins reaching well behind
the bases of the pelvic fins distinguishes the trout-perch from all other
Alaskan freshwater fishes.
DESCRIPTION
Body elongate, terete, noticeably heavier anteriorly. Greatest depth
just behind head, about 181 of TL. Head large, more or less conical, its
length 22-27% of TL. Snout long, rounded anteriorly, 30-38% of head length.
Eye round, large, 22-27% of head. Mouth small, subterminal, maxillary not
reaching eye. Teeth small, bristle-like, in bands on lower jaw and premax-
illa. No teeth elsewhere in mouth. Gill rakers short, stubby mounds with
small teeth, 8-13 on first gill arch. Branchiostegals 5-7, usually 6, on
each side. Lateral line nearly straight with 41-60 pored scales. Pyloric
caeca 7-14.
Fins
D. I-III (usually II), 9-11, adipose dorsal fin present. A. I, 5-8.
12-15. P2. 8-9. Caudal forked.
P1
Scales
Rather large, ctenoid, noticeably rough to the touch when brushed from
tail to head. No scales on head or,nape.
Color
Pale yellowish to silvery, often almost transparent. A row of about 10
dark spots along midline of back, 10 or 11 along lateral line, another row of
spots high on sides above lateral line. Fins transparent.
Size
The maximum size attained in the southern part of the range is said to
be about 200 mm (8 in). In most areas, however, the largest are only about
half this size. Alaskan specimens have run up to about 80 mm.
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RANGE AND ABUNDANCE
Trout-perch are found in North America. They range from the Del-Mar-Va
peninsula north to the shore of Hudson's Bay, west through West Virginia,
Tennessee and Missouri, northwesterly in the Mississippi drainage to the
prairie provinces and north eastern British Columbia, northward in the Macken-
zie drainage to the Arctic coast and down the Porcupine and Yukon Rivers
almost to the mouth. In Alaska, it has been found at the mouths of the Taton-
duk, Kandik and Charlie rivers, at Circle and Nulato, and at the mouth of the
Andreafsky River {UAFC 1327; Morrow, 1965; McPhail and Lindsey, 1970).
The trout-perch appears to be rare in Alaska (Morrow, 1965), but in other
areas it is an abundant species. In Heming Lake, Manitoba, the population of
adult fish was estimated at 2929-3636 fish per hectare (Lawler and fitz-Earle,
1968).
HABITS
Spawning takes place in late spring and summer. In Red Lake, Minnesota,
spawning fish were found from early June to mid-August (Magnuson and Smith,
1963), while in Heming Lake, Manitoba, most spawning occurred in the latter
half of May (Lawler, 1954). In Lake Erie, ripe individuals have been taken
from early May to mid-August (Scott and Grossman, 1973). Ripe fish were taken
at Circle, Alaska, on June 28 (McPhail and Lindsey, 1970).
At spawning time, adults move inshore to shallow water, or into shallow
tributaries of lakes (Magnuson and Smith, 1963; Langlois, 1954). Males gener-
ally dominate the spawning population, sometimes by as much as 10:1. Some
populations breed almost exclusively at night (Magnuson and Smith, 1963), but
others show no diurnal variation (Lawler, 1954). In the spawning act, two or
more males cluster with a single female within four or five inches of the
surface. They press close to the female, often breaking the surface of the
water, and eggs and milt are released. The eggs are yellowish in color,
about 1.5 mm in diameter before fertilization, demersal and sticky. They
sink to the bottom and adhere to whatever they settle upon. Water hardened
eggs are about 1.9 mm in diameter. Females 60-78 mm FL produce 210-728 eggs,
larger fish having more eggs (Lawler, 1954). Spawning has been observed in
water with temperatures between 4.5° and 17°C (Lawler, 1954). Hatching
occurs in about one week at 20°-23°C (Lawler, 1954; Magnuson and Smith, 1963).
Because Alaskan streams are so much colder than this, it is probable that
trout-perch eggs here require at least twice as long to hatch.
The young are about 5.5 mm long at hatching. They have a prominent yolk
sac, a large head with a prominent, pointed snout and a small, inferior mouth,
an oval eye, and are unpigmented save for a number of large, black, stellate
chromatophores on the yolk sac. By 7 mm in length, the yolk sac has been
absorbed and only a few large pigment cells are present on the right side of
the stomach region. All fins except the pel vies are well-developed by the
time the larvae are 9.5 mm long; the pel vies are just beginning to appear at
this stage. The young trout-perch leave the shallows and move out into deep
water about three weeks after hatching (Magnuson and Smith, 1963). The young
139
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fish at 50 mm is completely transformed and is fully scaled (Fish, 1932). By
the end of the first growing season, they may achieve lengths of 50-84 mm,
although Alaskan fish appear to grow much slower than this. Average length
at one year of age is about 81 mm; at two years, 94 mm; at three, 102 mm; and
at four, 115 mm. Females grow faster and live longer than males. Males
apparently do not survive beyond the third year (Trautman, 1957; Magnuson and
Smith, 1963).
Males begin to mature in their second summer (age 1+), but most females
do not mature until a year later. This difference in age at first maturity
apparently accounts, at least in part, for the predominance of males in the
spawning populations. Spawning groups are composed of 25-31% one-year-olds,
61-71% two-year-olds, 4-8% three-year-olds and 0-}% four-year-olds (Lawler,
1954; Magnuson and Smith, 1963). There is often (usually?) heavy post-spawn-
ing mortality. Amongst 2-year-olds, 96% of the males and 67% of the females
die after spawning; for 3-year-olds, 100% of males a'nd 96% of females; and
all four year old females (Magnuson and Smith, 1963).
Trout-perch are typically found in fairly deep water, 10-61 m, in lakes,
or in long, deep pools in streams. They move into shallow water for spawning.
Diurnal movements are rather noticeable, for the trout-perch spends the day
either in deep water (in lakes) or under cut-banks, roots, debris, etc. (in
streams), moving into the shallows to feed at night. Major foods are algae
(Microcystis), Cladocera, amphipods and the larvae of Chironomids and may-
flies. Small fishes are sometimes eaten in the winter (Nurnberger, 1930;
Langlois, 1954; Trautman, 1957).
IMPORTANCE TO MAN
The trout-perch is of no direct importance to man. However, it is an
important forage fish for larger carnivores wherever it is abundant. It is
known to be a major food for such fishes as pike, burbot, wall-eye, lake trout
and others. Its habit of feeding in the shallows, then moving to deep water
where it, in turn, is fed upon by larger fishes, may make it an important
factor in nutrient transfer, especially in stratified lakes (Lawler, 1954;
Langlois, 1954; McPhail and Lindsey, 1970),
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SECTION 12
CODFISHES--FAMILY GADIDAE
The codfishes are a large family, most marine. They have large heads,
wide gill openings, two or three dorsal fins and one or two anal fins, with
no spines in any fins, and usually have a small barbel at the tip of the
chin.
Cods are mostly northern hemisphere marine fishes, some of which enter
fresh water. One species, the burbot, is strictly freshwater and has a
circumpolar distribution in the northern hemisphere. A few species are
known from-the southern oceans,
The larger forms are very important food fishes and are the objects of
wide-spread and intensive commercial fisheries.
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BURBOT
Lota lota (Linnaeus)
DISTINCTIVE CHARACTERS
The long second dorsal fin, at least six times
ingle barbel on the chin distinguish the burbot.
rnTDTTnM
a single
DESCRIPTION
times as long as the first, and
rbot.
Body enlongate, robust and nearly round anteriorly, strongly compressed
behind anus. Depth 13-15% of total length. Head flattened, broad, its
length 19-20% of total length. Snout long, 27-33% of head length. Each
nostril in a prominent tube. Eye small, its diameter about 30% of snout
length. Mouth terminal, large, upper jaw reaching to below eye. A single,
prominent barbel present at tip of lower jaw. Teeth small and numerous, in
bands on jaws and head of vomer. Gill rakers short, 7-12 on first arch.
Branchiostegals 6-8 on each side.
Fins
60-80
short, 8-16 rays. Dp.
ays, joined to caudal. A.
fin short and rounded.
dal fin rounded.
2*
long, at least six times length of D-,
58-79, joined to caudal. P,. 17-21
mei
5-8, second ray elongate and filamentous.
with
the
Cau-
Scales
Minute, cycloid, embedded in the skin.
Color
Yellowish through brown to dark olive green above and on sides, color
generally darker in northern regions but also depending to some degree on
the color of the local environment. Sides blotched and mottled with pale
and dark shades, ventral parts usually pale yellow or white. Pelvic fins
pale, others dark and mottled.
Size
Reported up to 34 kg (75 Ib) and 1,524 mm (5 ft) long in Alaska (Turner,
1886; Dall, 1898), but the angling record for the state is only 10.2 kg (22.5
Ib). The average angler-caught fish probably weigh in the neighborhood of
0.5-1 kg (1-2 Ibs).
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RANGE AND ABUNDANCE
The burbot is circumpolar in the northern hemisphere. It is present,
in one or another of several subspecies, all across northern North America
from about 40°N to the Arctic Ocean, excluding only coastal Quebec and
Labrador and the northeastern part of Keewatin. In Europe and Asia it is to
be found from the Pacific (though not on Kamschatka) to the Atlantic, but is
missing from northern Scandinavia, Scotland and Ireland. Throughout its
range it is moderately to extremely abundant. In some of the lakes of north-
central North America, burbot are so numerous as to be a nuisance to commer-
cial gill-netters (Hewson, 1955).
HABITS
Burbots are winter spawners. The gonads begin to enlarge in August, but
spawning does not begin until well into winter. Depending on geographical
location, the actual spawning period may be from as early as mid-December to
as late as early April, with most populations breeding in January or February
(Cahn, 1936; Bjorn, 1940; Hewson, 1955; Mac Crimmon, 1959; Lawler, 1963;
Chen, 1969). In Alaska, as in most other areas, spawning apparently occurs
in late January and February (Chen, 1969). In the spawning act, which is
difficult to observe, since it goes on at night under the ice and the fish
show strong negative phototrophism, the burbot congregate in moderately
shallow water, 0.3 to more than 1.5 m (1-5 ft) deep but sometimes in 18-20
meters (Clemens, 1951b)s over bottom composed of clean sand, gravel and
stones. The males reach the spawning area first. At breeding, males and
females form a great globular mass of fish, individuals pushing toward the "
center and releasing eggs or sperm (Cahn, 1936), or at least milling around
close together (Mac Crimmon, 1959). There is no pairing, although Fabricius
(1954) observed definite pairing of European burbot in an aquarium.
Burbot mature at 2-4 years of age in the southern part of the range, but
not until six or seven years in interior Alaska (Chen, 1969). An average
adult female will produce between 500,000 and 750,000 eggs, with numbers up
to nearly 1,500,000 having been recorded (Bjorn, 1940; Lawler, 1963; Chen,
1969). Egg size seems to be quite variable, from a diameter of 1.7 mm in
Lake Erie to as little as 0.5 mm in Heming Lake, Manitoba. The eggs of
interior Alaskan fish range between 0.71 and 0.87 mm (Fish, 1932; Lawler,
1963; Chen, 1969). The eggs are not sticky, but are demersal and contain an
oil globule. After being spawned, they settle to the bottom and develop
without any care from the parents. Development time varies with temperature,
possibly also with the particular population. At 6.1°C, hatching occurs in
about 30 days, but 71 days are needed at temperatures between about 0 and
3.6°C (Bjorn, 1940; Mac Crimmon, 1959).
The newly hatched larvae are only 3-4 mm long, colorless, transparent
and without a yolk sac. Fin rays begin to appear in the dorsal, anal and
caudal fins at about 10 mm. By the time the young are about 19 mm long, fin
rays are present in all fins, the chin barbel is present and the little fish
is readily recognized as a burbot (Fish, 1932). Young burbot at this stage
of development have been found in the Chena River, interior Alaska, in June
(Chen, 1969).
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Growth rates are quite variable from place to place. As a generality,
it may be said that fish in the northern part of the range tend to grow more
slowly and to live longer than fish in more southern regions, although there
are exceptions. Males tend to be shorter lived than females and to grow more
slowly after age 10. Typical lengths at various ages for burbot in interior
Alaska are: 1 + , 136 mm; 2+, 185 mm; 3+, 238 mm; 4+, 283 mm; 5+, 355 mm, 10+,
595 mm; 15+, 715 mm; 20+, 871 mm; 24+, 972 mm (Chen, 1969). Fish from lakes
Erie, Simcoe and Winnipeg, for example, grew much faster than Alaskan fish,
but those from Heming Lake, Manitoba, grew more slowly, at least after the
first year (Clemens, 1951b; Hewson, 1955; Mac Crimmon, 1959; Lawler, 1963).
Chen's 24 year old fish seems to be the oldest on record. In general, it
appears that relatively few burbot reach an age much beyond 15 years.
The burbot is usually a resident of fairly deep water, whether in lakes
or in rivers. In lakes, burbot have been taken as deep as 213 m (700 ft),
and seem to be confined to the hypolimnion in the summertime. River fish
tend to congregate in deep holes throughout the year, except at spawning.
Optimum temperature for burbot is reported as 15.6-18.3°C (Scott and Cross-
man, 1973).
Migratory patterns of burbot are inadequately known. The fish are gen-
erally rather sedentary and do not move about very much. However, there do
appear to be definite movements toward spawning areas, although as a rule
these are individual movements rather than migrations of a whole school to-
gether. They also may move into shallow water to feed at night in the sum-
mer. Burbot have also been observed to make post-spawning runs up river,
apparently for feeding (Mac Crimmon, 1959).
In its food habits, the burbot is an omnivorous carnivore, although
displaying strong preference for a fish diet. Young Alaskan burbot in their
first and second years feed mostly on insect larvae, especially Plectoptera
and Ephemeroptera, and on small sculpins, Cottus cognatus. Beginning in the
third year, at a length of about 180 mm, there is a steady shift towards
fishes of all sorts and away from invertebrates. From about age 5 on (355
mm TL), fishes of various sorts compose 67-90% of the food, with inverte-
brates making up less than 12% (Chen, 1969). In more southerly climes, the
dependence on invertebrates seems to change at about the same age (5 years)
but at a much larger size (500 mm).
The list of food items is a long one, depending at least in part on
what is available. In addition to those items already mentioned, it includes
molluscs, asellids, Mysis, Pontoporela, Gammarus, Trichoptera, and crayfish,
as well as Cisco eggs, mice,shrews and at least 20 species of fishes. Young
burbot are an important food of large burbot (Nurnberger, 1930; Van Oosten
and Deason, 1938; Clemens, 1951a; Mac Crimmon and Devitt, 1954; Hewson, 1955;
Beeton, 1956; Bonde and Maloney, 1960; Lawler, 1963; Chen, 1969). Two fish
taken by the author in the Tanana River near Tetlin Junction, Alaska, in the
early 1960's had adult bank swallows in their stomachs.
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IMPORTANCE TO MAN
The burbot makes up a relatively small part of the freshwater fish catch
of the United States. Maximum landings (over 277,000 kg per year) occurred
in the late 1940's and early 1950's, since declining slightly to an average
of 173,600 kg (382,000 Ib) per year for the six years 1968-1973. Most of
this catch comes from the Great Lakes and the international lakes, although
since 1968 the upper Mississippi basin has supplied about 25% of the total.
The price has fluctuated widely during this time, from less than $0.02 to
more than $0.11 per pound. Fish from the Mississippi drainage generally
coimiands a higher price than fish from the Great Lakes. Almost all U. S.-
caught burbot is used for food, either for men or for animals (Anonymous,
1971-1976). Attempts to popularize the burbot in Canada have thus far met
with but little success (Scott and Grossman, 1973). Burbot are an important
commercial product in Siberia (Nikolskii, 1961).
As a food fish, burbot has enjoyed a mixed reputation. Early explorers
in the Canadian Arctic ate it "...only in times of great scarcity" (Richard-
son, 1836), but the same author also noted that "When well bruised and mixed
with a little flour, the roe can be baked into very good biscuits...". The
liver is also considered a delicacy, at least in Europe, and is used as a
pate for canapes, as well as fried or smoked and eaten for itself. Burbot
liver oil contains as much vitamins A and D as does cod liver oil (Branin,
1930). Actually, burbot is an excellent food fish, for its flesh is white,
flaky, of good flavor and almost boneless. The repulsive appearance of the
fish apparently militates against its popularity.
In Alaska, small numbers of burbot are taken for subsistence. Sport
fishermen probably take more than do the subsistence fisheries. Icefishing
for burbot^in winter is reasonably popular and more and more anglers are
learning that a burbot on light tackle is a worthy opponent at any time of
year.
ARCTIC COD
Boreogadus saida (Lepechin)
DISTINCTIVE CHARACTERS
The distinctive characters of the Arctic cod are the three dorsal and
two anal fins; the lower jaw as long as or slightly longer than the upper;
the minute chin barbel; and the forked caudal fin.
DESCRIPTION
Body slender, elongate. Depth about 15% of fork length, greatest depth
just behind head. Head moderate, 24-27% of fork length. Snout rather long,
about 30% of head length. Eye round, large, its diameter 25-33% of head.
Mouth fairly large, terminal; rather small, sharp teeth in two rows in front
of upper jaw, in one row on posterior part as well as on lower jaw and vomer,
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the teeth wide set. A very small barbel usually present at anterior tip of
lower jaw, sometimes scarcely visible. Gill rakers long and slender, 37-46.
Lateral line discontinuous. Pyloric caeca 20-37.
Fins
D]. 10-14, rarely to 16. D2. 14-17. D3. 18-24. A]. 15-20. A2.
18-24. P-j. about 19. P£. 6, trie second ray notably elongate. Caudal fin
forked, the tips of the lobes slightly rounded.
Scales
Cycloid, minute.
Color
Brownish or grayish brown above, sides above lateral line somewhat
lighter, often with a yellow or purple tinge. Sides below lateral line, and
ventral surface, silvery. Fine black dots scattered over body, most numerous
on back. Fins dusky; dorsal and caudal with a narrow white edge; anal pale
at base.
Size
One of the smallest of the cods, most adults about 130-160 mm (ca 5-6
in) long, the largest specimen known said to have been only 321 mm (Svetovi-
dov, 1948). Weight to about 75 g (about 1 oz).
RANGE AND ABUNDANCE
The Arctic cod enjoys a circum-polar distribution in most of the Arctic
Ocean, being absent only from the region just north of Scandinavia. On the
Atlantic side of North America it has been found as far south as the St.
Lawrence River (Vladykov, 1945) and even to the estuary of the Miramichi
River, New Brunswick (HcKenzie, 1953). It ranges throughout the Arctic
Ocean across the arctic coast of Canada and Alaska, south in the Bering Sea
to Norton Sound on the east side, to the Gulf of Anadyr on the west; westward
along the Siberian coast to about 37-38°E; and north probably to the North
Pole (Andriyashev, 1954). Although primarily a marine species, the Arctic
cod seems to prefer the less saline waters, 15-30 parts per thousand salini-
ty, and not infrequently runs far upstream in rivers. The Arctic cod is one
of the most abundant fishes present in arctic seas (Bean, 1887a, b, c; Alver-
son and Wilimovsky, 1966). The density of juveniles in the eastern Chukchi
Sea in September and October, 1970, was estimated at 28 fish per 1,000 n3 of
water, or about 700 kg of young fish per km2 (Quast, 1974).
HABITS
The Arctic cod is a demersal species, the adults associated with some
kind of substrate, either the sea bottom or the underside of ice packs. Its
distribution is closely correlated with the presence of ice and/or lowered
salinity. Sexual maturity is reached at 3-4 years of age, and the fish are
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reported to spawn only once in their lives (Nikolskii, 1961), The eggs are
the largest of any gadid, 1.5-1.9 mm in diameter. The large eggs and the
small size of the fish combine to produce low fecundity, 9,000-21,000 eggs
per female. Spawning takes place from October to March, usually in January
and February. At this time, the fish move close inshore, often in tremendous
numbers. They may enter rivers, and spawning in fresh water has been record-
ed in Siberia (Svetovidov, 1948). The eggs are pelagic, floating in the
surface waters.
Development to hatching requires several months. The first larvae,
about 5 mm long, appear in May to July. The larval stage lasts about 2
months. Small individuals 5-9 mm long have been taken in Siberian waters in
Hay, Larger larvae, 9-8 mm long, are known from near the west coast of
Greenland north of 68°N in late June. Transition to the juvenile stage
occurs at the end of summer at lengths of 30-50 mm (Jensen, 1926; Andriya-
shev, 1954; Rass, 1968; Quast, 1974).
The young-of-the year appear to be negatively phototrophic, avoiding
light. They are much more numerous below 20 m depth than above. This may
be a mechanism whereby the fish reduce predation by sea birds (Quast, 1974).
Growth of Arctic cod in the northern Bering Sea has been given as
follows: 0+, 31 mm; 1 + , 75-100 mm; 2+, 144-158 mm; 3+_, 190-200 mmg; 4+,
220-230 mm. This is rather faster growth than is found in more southerly
regions along the east coast of Siberia (Andriyashev, 1954).
Arctic cod apparently undertake onshore-offshore migrations, in part
associated with spawning and in part with movements of ice. They are
common along the edge of pack ice, where they are said to swim rapidly be-
tween the blocks and to hide in cracks in the ice. The Arctic cod shows
strong preference for low temperatures. Eggs do well at 0°-2°C, larvae at
2°-5°C, and fry at 5°-7°C (Rass, 1968). Older fish prefer lower tempera-
tures. Adults seem to be most abundant in water with temperatures around
-1° to -1.85°C (McKenzie, 1953; Backus, 1957; Leim and Scott, 1966).
Arctic cod feed mostly on plankton. Larvae and young fry eat copepod
eggs and larval stages. Adults have a more varied diet, with euphausiids
and calanoid copepods (especially Calanus) as the most important items but
also including amphipods, shrimps, fish eggs and small fishes (Svetovidov,
1948; Andriyashev, 1954; Lei in and Scott, 1966; Ponomarenko, 1967;
Hognestad, 1968).
IMPORTANCE TO MAN
The Arctic cod's chief importance to man is indirect. It is one of
the major foods of many arctic marine animals, including fishes, seals,
belugas, narwhals and birds. At Cape Thompson, Alaska, Swartz (1966) esti-
mated that the sea bird colony consumed about 13,100,000 kg of fish per
breeding season, of which the majority was Arctic cod.
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Direct importance to man is relatively low. The Arctic cod is reported
to be of low palatability. Hence, although it is fished commercially and in
subsistence fisheries, especially by the Russians, most of the catch is used
for animal food. The liver, however, contains up to 50% valuable oil.
PACIFIC COD
Gadus macrocephalus Tilesius
DISTINCTIVE CHARACTERS
Three dorsal and two anal fins, a long chin barbel (about 3/4 as long
as eye diameter in young, longer than eye diameter in adults) and the space
between the second and third dorsal fins shorter than the eye diameter distin-
guish the Pacific cod.
DESCRIPTION
Body elongate, nearly cylindrical anteriorly, more or less compressed
posteriorly. Depth about 2Q% of total length, deepest below first dorsal fin.
Head large and broad, its length 26-29% of total length. Snout blunt, long,
about 33-37% of head length. Eye round, its diameter 50-65% of snout length.
Mouth large, subterminal, lower jaw shorter than upper, maxilla reaches to
below eye. Small, sharp teeth present on both jaws and on head of vomer, the
outer row of jaw teeth larger than the others. Gill rakers 18-23 on first
arch. Lateral line with a prominent arch under first and second dorsal fins,
straight posteriorly, ends under third dorsal.
Fins
D]. 13-15; D2. 16-21; D3. 15-21. A]. 17-22; A2. 16-20. P-j. 19-22;
6-7, second ray longest. Caudal fin truncate.
Scales
Cycloid, small.
Color
Quite variable, usually brownish or olive gray above, paler below, with
numerous brown or grayish spots on sides and back. Yellow color phases are
known. Fins dusky; dorsal, anal and caudal fins with white edges, wider on
anals and caudal than on dorsal.
Size
Reaches a
caught fish is
(10 lb).
length of 1,200 mm (nearly 4 ft).
around 700-750 mm (27-29 in) with
Average size of trawl-
a weight of about 4.5 kg
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RANGE AND ABUNDANCE
Pacific cod are found along the North American coast from southern
California, off Santa Monica at 33° 59.5'S, north to Norton Sound and St.
Lawrence Island in the Bering Sea. On the Asian side the species is present
from the southeastern tip of the Chukhotsk Peninsula through the Anadyr Gulf
and southward to the Yellow Sea (Popov, 1933; Andriyashev, 1937, 1954;
Svetovidov, 1948; Pinkas, 1967). It is rare south of Oregon, however,
becoming most numerous, as well as larger, in the northern part of the range.
From Puget Sound on northward and well into the Bering Sea, Pacific cod are
abundant and support commercial fisheries of considerable size. Cod appear
to prefer temperatures between 6° and 9°C (Ketchen, 1961).
HABITS
Like other cod fishes, Pacific cod spawn in the winter, usually in
January and February. Spawning grounds are in deep water, to as deep as
250 tn (137 fath). The eggs are small, average diameter about 1 mm, and
fecundity is high. Egg numbers tend to increase with the size of the fish.
A female 400 mm long contains about 228,000 eggs while one of 800 mm may
produce about 3,350,000. Egg counts as high as 5,700,000 have been recorded.
The eggs are demersal, with specific gravity of about 1.050, and are adhes-
ive at first. They lose their adhesiveness about 30 hours after fertiliza-
tion. Development to hatching requires about 8-9 days at 11°C but 29 days
at 2°C. Experiments have shown that the maximum numbers of larvae are
produced at salinities of about 19 o/oo and temperatures of about 5°C
{Andriyashev, 1954; Thomson, 1962, 1963; Forrester, 1964, 1969b; Forrester
and Alderdice, 1966).
The larvae are 3.5-4.8 mm! long at hatching and have a fairly large
yolk sac which is absorbed in 6-10 days, the time depending on temperature.
Early growth is rapid and by the end of the first year of life the young
fish are 230-260 mm long. Subsequent growth is, of course, much slower, and
quite variable with different locations and populations. Typical lengths
for various ages are: 2+, 500 mm; 3+, 614 mm; 4+, 673 mm; 5+, 726 mm; 6+,
762 mm. Use of otoliths for aging Bering Sea cod gave much shorter lengths
for corresponding ages. In contrast to most fishes, Pacific cod appear to
grow just as fast in winter as they do in summer. Cod on the- Asian side of
the Bering Sea do not grow as fast as these eastern Pacific fish. Sexual
maturity is reached at 2-3 years of age in the southern parts of the range,
but about two years later in the northern areas. Maximum age attained is
about 13 years, but in most areas few live beyond about 7 years (Svetovidov,
1948; Masher, 1953; Andriyashev, 1954; Forrester and Ketchen, 1955; Ketchen,
1961, 1964; Forrester, 1969b).
Pacific cod do not, as a rule, undertake extensive migrations. They
move inshore after spawning, to depths of 30-60 m; then, from late summer
to mid-winter, move out into the deeper water (100-250 m) of the spawning
areas. In at least some North American stocks, there appears to be a good
deal of movement from one bank to another. Most of these migrations are
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relatively short, but at least one tagged fish was recovered about 125 km
(200 mi) from the point of release (Andriyashev, 1954; Forrester and Ketchen,
1955).
Young cod, like so many young fishes, probably feed largely on copepods
and similar organisms. Adults feed heavily on shrimps, herring, sand lance
and crabs, in that order, but also take almost any kind of small fish avail-
able, as well as worms, euphausiids, amphipods, isopods, echinoids, octopus
and even ducks (Cobb, 1927; Hart, 1949; Andriyashev, 1954; Forrester and
Ketchen, 1955).
IMPORTANCE TO MAN
The Pacific cod is one of the more important secondary commercial fishes
of the Pacific coast. Total landings in the U. S. and Canada run on the
order of 5-10 million kg (11-22 million lb) annually. The fishery began in
1863, when Captain Turner, on the brig Timandra, brought a fare of salt cod
to San Francisco from Sakhalin. A small fleet was organized and fished the
same region two years later. Fishing off the Shumagin Islands, Alaska,
began in 1867. Subsequently, nearly all the cod fishing was done in Alaskan
waters and the fishery prospered. Catches rose, more or less steadily, to a
peak level of 2-4 million fish (probably about 10-20 million kg or 20-40
million Ib) in the 1920's. Nearly all the fish were landed at ports in
Washington and California. Economic factors resulted in the decline of the
Alaskan fisheries and the concommitant rise of the industry around British
Columbia and Washington. By the late 1950's, the Alaska catch was no longer
listed separately in the fisheries statistics and did not re-appear until
1973 when landings of 72,000 kg (158,600 Ib) were reported (Power, 1962;
Anonymous, 1976).
The entire catch was salted in the early days, and the oil was extracted
from the livers (10,000 gallons in 1866). With the advent of modern freez-
ing techniques, more and more of the catch is frozen and used for fish
sticks and similar products. The liver oil contains large quantities of
Vitamins A and D, and is quite comparable to liver oil of Atlantic cod in
this respect. Oil derived from the viscera is 3-9 times as ootent, but the
average yield is less than 0.1 that of the liver (Pugsley, 1938). The
development of synthetic vitamins has adversely affected the market for
these products.
SAFFRON COD
Eleginus gracilis (Tilesius)
DISTINCTIVE CHARACTERS
The saffron cod is set off by the three dorsal and two anal fins,
lower jaw shorter than upper, chin barbel- not longer than 1/2 eye diameter,
and the space between the second and third dorsal fins equal to or longer
than the eye diameter.
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DESCRIPTION
Body elongate, nearly round in cross-section anteriorly, slightly
compressed posteriorly. Depth about 16% of total length. Head 21-251 of
total length. Snout long, about 331 of head length. Eye round, moderate,
its diameter about 18% of the head length, or 1/2 snout length. Mouth
large, subterminal, lower jaw distinctly shorter than upper, maxilla reach-
ing to below eye. Teeth small, slender, curved backwards, in several rows
in upper jaw; in two rows in anterior part of lower jaw, a single row pos-
teriorly. A patch of small teeth present on head of vomer. Gill rakers
14-24, usually about 20, on first arch. Lateral line curved in its anterior
portion, ending posteriorly under second dorsal fin.
Fins
11-15. D
15-23. D
18-21.
20-24. A
19-23. P-
2. -. 3. -. ]. -. 2. -. -,
20. P£. 6, the second ray produced. Caudal fin truncate or slightly
emarginate.
Scales
Cycloid, small, about 157 in a mid-lateral series.
•Color
Grayish brown to olive above, upper part of sides paler, sometimes with
a silvery-violet shading, often mottled with indistinct darker blotches.
Lower sides and belly yellowish to silver white. Fins dusky, dorsals and
caudal with white edges.
Si ze
Length usually around 250-350 mm (10-14 in), but up to about 500 mm
(20 in) and a weight of 1 kg (2.2 Ib).
RANGE AND ABUNDANCE
Confined to the North Pacific Ocean and the Bering and Chukchi seas,1
from Sitka (Svetovidov, 1948) to at least as far north as Kotzebue on the
Alaskan coast and from the Chukhotsk Peninsula south to Korea and the Yellow
Sea on the Asian side. The saffron cod not infrequently enters rivers and
may go considerable distances up-stream. It has also been found in fresh-
"water lakes on Bering Island and the Kamschatka Peninsula (Jordan and Metz,
1913; Svetovidov, 1948; Andriyashev, 1954).
Throughout most of its range, the saffron cod is quite abundant,
sufficiently so to be the object of commercial fisheries.
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HABITS
Saffron cod spawn in the winter, December to February. The fish move
from deeper (30-60 m) water to shallower, inshore water for this purpose.
In Norton Sound, saffron cod have been reported spawning over pebbly
bottoms (Turner, 1886). Water temperatures on the spawning grounds are
-1.6 to -1.8°C. Fecundity is between 25,000 and 210,000 eggs per female.
The eggs are said to be adhesive (Svetovidov, 1948). The young hatch in the
spring, April to June, and for the first few months of their lives are plank-
tonic. They are frequently encountered under the bells of large jellyfish
(Cyania ferruginea) (Andriyashev, 1954).
Growth is rather slow. Three year old fish average about 300 mm long
and weigh about 200 g (6 oz); at five years, 375 mm and 375 g; and at 7
years, 460 mm and 800-900 g. Sexual maturity is reached at 2 or 3 years,
after which the fish spawn annually (Svetovidov, 1948; Andriyashev, 1954).
Saffron cod do not migrate long distances. They move inshore in fall
and winter for spawning and offshore in spring and summer to feed in deeper
water. They often enter rivers and may go considerable distances upstream.
However, they usually stay within regions of tidal influence. They appar-
ently are restricted to the less frigid parts of the arctic seas, being
replaced by the Arctic cod in the colder regions (Popov, 1933; Svetovidov,
1938; Alverson and Wiliniovsky, 1966).
Food of the saffron cod consists mainly of small crustaceans, such as
shrimps, amphipods and mysids, as well as polychaete worms and small fishes.
IMPORTANCE TO MAN
Along the Alaskan coast the saffron cod is utilized almost exclusively
in subsistence fisheries. The coastal eskimoes catch considerable numbers
with hand lines through holes in the ice. Along the Siberian coast, commer-
cial fisheries of some magnitude are directed at this species. Current
statistics are not available, but landings in the post-war years were on
the order of 18,000,000 kg annually. The flesh of the saffron cod is said
to be very tasty in late fall and winter but not so during the warmer months
(Andriyashev, 1954).
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SECTION 13
STICKLEBACKS—FAMILY GASTEROSTEIDAE
The members of this family are to be found in marine and fresh waters
of Europe, Asia, North America and the western Mediterranean coast of
Africa. Characteristically, they have three or more free spines in front
of the dorsal fin and one before the anal fin. The pelvic fin is reduced
to a single, strong spine with 0-3 soft rays.
They are all small fishes and are often important in the food of larq-
er fishes, birds, etc. In some situations, they are strong competitors for
food with young salmon.
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THREESPJNE STICKLEBACK
Gastergsteus aculeatus Linnaeus
DISTINCTIVE CHARACTERS
The three (rarely 2 or 4) sharp, free spines before the dorsal fin, the
pelvic fin reduced to a sharp spine and one small ray and the series of
plates along the sides of the. body identify the threespine stickleback.
DESCRIPTION
Body compressed, moderately robust, its depth about" 25% of total length.
(Deeper bodied individuals are usually either gravid females or are infected
with a cestode, Schistocephalus, which may fill the body cavity). Head al-
most 251 of total length. Snout 27-31% of head length. Eye round, its dia-
meter generally a little less than snout length. Mouth oblique, small,
maxilla not reaching anterior margin of eye. Teeth snail, fine, on both
jaws. Gill rakers long and slender, 17-25 on first arch of strictly fresh-
water forms, one or two more in anadromous forms. Branchiostegals 3 or 4.
Lateral line more or less parallel to dorsal outline, pores microscopic.
Pyloric caeca 1 or 2, short.
Fins
D. II-IV (usually III), 10-14, the spines separate from each other and
from the soft-rayed fin, each spine with a reduced membrane attached to its
posterior side. A. I, 6-10, the spine free from the rest of the fin. P-j.
9-11, posterior margin nearly truncate. Pg- 1,1. Caudal truncate to
slightly indented.
Seales
No scales, but the sides with vertically enlarged bony plates. In
freshwater populations, these usually number 5-9 on each side, with 7 the
irost common number. Marine and anadromous sticklebacks are completely ar-
mored with 22-37 plates. A few freshwater populations have 0-2 plates. The
anadromous type has a keel along each side of the caudal peduncle.
Col or
Freshwater forms usually mottled brown or greenish, anadromous fish
silvery green to bluish black. A few isolated populations, in the Chehalis
River system of Washington and in Mayer Lake, Queen Charlotte Islands,
British Columbia, are black. Sides usually paler, belly yellow, white or
silvery. Fins pale, pectoral rays often picked out with dark dots. (Green-
bank and Nelson, 1959; McPhail, 1969; Moodie, 1972a, b).
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Breeding males (except of the black forms) become brilliant bluish or
green, with blue or green eyes, and the forward part of the body, especially
the breast region, bright red to orange.
Size
Maximum size reported to about 100 mm (Jordan and Evermann, 1896), but
the size of adults in different populations is highly variable.
RANGE AND ABUNDANCE
Along the Pacific coast of North America from Rio Rosario, Baja Calif-
ornia, north to the Bristol Bay region of Alaska, and St. Lawrence Island.
On the Asian coast, south to Korea. On the Atlantic side of North America,,
this stickleback is found from Chesapeake Bay north along the coast to the
western shore of Hudson Bay, and to Baffin Island and Devon Island in the
Canadian Arctic. It is also present in southern Greenland, Iceland and most
of western Europe, Wherever present, the threespine stickleback is usually
abundant.
HABITS
Description of the habits of the threespine stickleback is not made any
easier by the great variation shown between populations. It is quite proba-
ble that several subspecies or even full species are involved in this group,
but adequate analysis has not yet been possible and authorities are divided
in their opinions. Some suggest that the variations represent the response
of a single, plastic species to different environmental conditions. Others
feel that various degrees of speciation have occurred/still others that the
differences-are due to several lines of evolution (Hagen, 1967; Miller and
Hubbs, 1969; McPhail, 1969; Hagen and McPhail, 1970; Moodie, 1972b; Ross,
1973; Bell, 1976). The discussion below is an attempt to provide a general-
ized summary, based on many published accounts and personal observations.
Breeding occurs in the spring and summer, usually June and July in
Alaska, with a few fish in breeding condition as late as August. Just prior
to the breeding season, males become strongly territorial and take on the
breeding colors. They defend their territories by ritual displays, a head-
down position, and by chasing and biting invading males. The resident male
builds a barrel-shaped nest, using sand-grains, bits of vegetation, etc,
which he cements together with a glue secreted by a special portion of the
kidney. When the nest is completed, with an en-trance and an exit, the male
courts females by means of a zig-zag dance toward the female, then turning
and swimming toward the nest. The intensity of the zig-zag dance is extrem-
ly variable in different populations, ranging from short dashes each way to
little more than head-wagging. Non-receptive females ignore the dance and
may be driven from the territory, but a receptive female will follow the
male to the nest. There he points out the entrance by posing above it with
his head down. The female enters the nest and deposits a clutch of eggs.
At this time, males may butt the side of the nest, which apparently stimu-
lates the female to lay her eggs. Fecundity ranges from 80 to nearly 1,300
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eggs, but only 50-200 are deposited at one time. Females may lay eggs in
several nests on several days, or may be courted again by the same male. In
between spawnings, which usually occur at intervals of 3-4 days, the female
may eat her own weight of food. If adequate food supply is not available,
egg production goes on at the expense of body tissues, or spawning stops
(Wootton and Evans, 1976).
When the female has laid her eggs, she leaves the nest by the exit.
Attempts to back out of the nest are resisted by the male. Following egg
deposition, which takes several minutes, the male may immediately enter the
nest and fertilize the eggs, or, less frequently, he may court another female
before entering the nest himself. In any case, the female is driven away
after the eggs have been deposited. Several females generally are induced
to deposit eggs in each nest, and it is not uncommon for a male to have
several sets of eggs, at different stages of development, in a single nest.
Following courting and fertilizing, the male guards the nest. He also
takes a characteristic position in front of one of the entrances, maintaining
his body at an angle of about 45° from horizontal with head toward the nest,
and fanning with his pectoral fins.
The eggs are sticky and adhere to each other. Diameter of newly laid
eggs is 1.1-1.6 mm, but they quickly swell to 1.5-1.9 mm. Hatching time may
be as short as six days at 17°C in California to as long as 14 days at 9-16°
C in Karluk Lake, Kodiak Island, Alaska. The young are about 4.2-4.5 mm long
at hatching and have a large, spherical yolk-sac. About 8 days after hatch-
ing, the young, now about 7 mm long and with the yolk-sac absorbed, begin to
emerge from the nest and the male is kept busy catching them in his mouth
and spitting them back into the nest. When the young are fully developed
and free-swimming, they school around the male for a few days, then disperse.
The male may now build a new nest and repeat the cycle.
Growth rates are extremely variable. In Karluk Lake, young threesoine
sticklebacks are about 28 mm by early August, but only about 12 mm long in
the Dixon River, Glacier Bay National Monument. It is not known whether this
reflects slower growth or later spawning in the latter area. Karluk Lake
sticklebacks at one year of age are about 47 mm long, 60 mm at two years.
In Cowichan Lake, British Columbia, threespine sticklebacks reach 64 mm in
only 10 months. Normal life span in this population is 2 1/4 years; else-
where it ranges from 1 to 3 years.
Sexual maturity is first attained at one year, though most do not mature
until the second year. It is not known whether the early-maturing fish
survive to spawn a second time. In at least some populations, the fish are
hermaphroditic. Males have apparently non-functional ovaries while females
may have functional testes and perhaps are capable of self-fertilization.
At any rate, Karluk Lake females have been found with eyed eggs in the
ovaries.
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Freshwater threespine sticklebacks generally inhabit shallow water in
association with a modicum of aquatic vegetation. However, they have been
found in large numbers at the surface over deep water in some Alaskan lakes
and as deep as 24 m (80 ft) in Karluk Lake. (Kuntz and Radcliffe, 1917;
Tinbergen and Van lersel, 1947; Vrat, 1949; Jones and Hynes, 1950; Carl,
1953; Greenbank and Nelson, 1959; Narver, 1969).
The food of sticklebacks consists mostly of zooplankton and insects.
Copepods, Cladocera and chironomids are the most important item, but the
list is a long one, including rotifers, ostracods, oligochaetes, small
molluscs, amphipods, beetle larvae, leeches, planarians, mites, and their
own eggs and larvae (Markley, 1940; Hynes, 1950; Carl, 1953; Greenbank and
Nelson, 1959).
Although the number of lateral plates on freshwater threespine stickle-
backs varies between 0 and 9, the modal number in most populations is 7.
This number of plates seems to be associated with behavioral traits which
make those individuals less subject to predation by larger fishes such as
trout and squawfish. The threespine stickleback relies mostly on camouflage
and cryptic coloration for protection. Lying motionless in the water, as
they often do, the fish resemble a floating stick or twig. Individuals with
more or less than seven lateral plates apparently are more active and hence
more likely to attract the attention of predators than are seven-plated
fish. Why this is so in unknown (Greenbank and Nelson, 1959; Hoodie, 1972a;
Moodie et al, 1973; Hagen and Gibertson, 1973).
The anadromous threespine stickleback differs from the freshwater form
in a number of respects. Morphologically, this difference appears in the
lateral plates and gill rakers. The anadromous form is fully plated, with
up to 37 plates on the sides and a rather pronounced keel on each side of
the caudal peduncle. In addition, the number of gill rakers on the first
arch tends to be slightly higher in the anadronomous form.
Life history also differs. Spawning behaviour is essentially like that
of the freshwater form, but breeding may occur in either fresh or salt water
in salinities at least as high as 28.5 o/oo (McPhail, 1969). Sexual matur-
ity is more often reached at one year rather than two, and the spawning
areas seem to be in thicker vegetation, such as eel-grass beds (Narver,
1969). The young leave the streams and estuaries where they were hatched
and move out into salt water. At first they stay close to shore, sheltering
in seaweeds and such, but enter the open sea in the fall. Some remain near
shore through the winter, but others apparently move out for considerable
distances. Large numbers have been taken at the surface 800 km from shore
in the Gulf of Alaska and on Georges Bank in tha Atlantic (Bigelow and
Schroeder, 1953; Andriyashev, 1954; Clemens and Wilby, 1961; McPhail and
Lindsey, 1970). Whether or not these little fish which go so far from shore
ever find their way back is a moot question.
IMPORTANCE TO MAN
The three spine stickleback is of but little direct importance to man.
It has occasionally been taken commercially in Scandinavia and rendered into
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meal and oil, the latter said to have a healing effect on wounds and burns
(Bigelow and Welsh, 1925; Nikolskii, 1961). It is an important forage
species for predaceous fish like Dolly Varden, lake trout and northern pike.
Large populations in some lakes may have adverse effects on the growth of
young sockeye salmon through competition for food, but its old reputation
and name of "salmon killer" (Jordan and Evermann, 1896) seem to be complete-
ly unjustified. The threespine stickleback makes an interesting aquarium
fish.
NINESPINE STICKLEBACK
Pungitius pungitius (Linnaeus)
DISTINCTIVE CHARACTERS
The ninespine stickleback is set off by the presence of 7-12 free spines
before the dorsal fin, and a long caudal keel, usually reaching to beneath
the dorsal fin.
DESCRIPTION
Body elongate, slender, its greatest depth 19-22^ of total length;
caudal peduncle notably long and slender. Head short, 20-23% of total
length. Snout about equal to eye diameter, about 25% of head. Mouth
oblique, small, reaching back about to anterior margin of eye. Small, fine
teeth present on both jaws. Gill rakers rather slender, 10-15 on first arch.
Branchiostegals 3 on each side. Lateral line nearly straight, the pores
microscopic. No pyloric caeca.
Fins
each with a rudementary mem-
spine stout and curved. P-j
D. VI-XII, 9-13, the spines separate and
brane on its posterior side. A. I, 8-11, the
10-11, posterior edge of fin rounded. Pp. 1,1, the ray closely appressed.
The pelvic girdle is occasionally reduced or absent (Nelson, 1971). Caudal
usually truncate, varies from slightly indented to slightly rounded.
Scales
No scales. 0-15 small, bony plates along anterior part of lateral line.
Caudal peduncle with a noticeable keel which may reach forward to below an-
terior edge of dorsal fin.
Color
Dull olive to light brown above, sides mottled or blotched with darker
patches of the same color. Belly yellowish to silvery white. Fins color-
less. Breeding colors may be quite variable, depending on sex, population
and stage of the breeding cycle. Colors of females are always less intense
than those of males. Aggressive females become dark dorsally and paler
below, then sometimes become paler with more conspicuous saddle marks as
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actual breeding approaches (McKenzie and Keenleyside, 1970), Aggressive
males become totally black, except for the colorless fins and the membranes
on the pelvic spines, which are white. At breeding, the males become paler
dorsally and more intensely black ventrally, especially under the chin
(Morris, 1958; McKenzie and Keenleyside, 1970). Breeding males on the east
coast of North America have been reported as reddish under the head and
greenish on the belly (Bigelow and Welsh, 1925).
Size
Up to 90 mm (3.5 in), but the average is about 65 mm (2.5 in) or less.
RANGE AND ABUNDANCE
The ninespine stickleback is circumpolar in its total distribution. In
Asia, it ranges from the Yangtse River of China north to the Arctic Ocean,
westward across northern Siberia to the Atlantic coast of Europe as far
south as the Loire basin in France. Present on the southern tip of Green-
land. In North America, the species ranges from Cook Inlet, Alaska, north
along the coast to the Arctic Ocean, east to the Atlantic. The Arctic North
American range includes Victoria and Baffin islands and most of the arctic
archipelago. From the mouth of the MacKenzie River, the ninespine stickle-
back ranges southeastward across Canada to the Great Lakes (except Lake
Erie) and south along the coast to New Jersey. Isolated populations are
present in the upper Mississippi drainage (Nelson, 1968b, c,). There are
two quite distinct forms in North America, a coastal, Bering form confined
to Alaska and the coastal regions of Canada and eastern United States, and
an inland, Mississippi form found in the rest of the range. The two groups
are distinguished on the basis of lower average numbers of gill rakers and
dorsal spines and higher numbers of lateral plates in the Bering form
(McPhail, 1963). Wherever it is present, the ninespine stickleback is
abundant.
HABITS
The ninespine stickleback is quite strictly a freshwater form. Although
it can survive salinities up to 20 o/oo, fresh water is required for spawning
(Nelson, 1968c). The coastal form, however, may winter in the sea, at least
in Hudson Bay, and has been taken in Norton Sound at Nome (McPhail and
Lindsey, 1970).
Spawning occurs in the spring and summer, generally Hay to July, but
ripe females have been found as early as April and as late as August
(Nelson, 1968b). The first indication of the onset of the breeding season
is a breakdown of schooling and a beginning of aggressive behavior. Subse-
quently, individuals, both males and females, set up territories which are
sex. At this time, males become black,
(see Color). Males construct nests of
are generally located in aquatic vege-
but may be on the bottom or even
defended against intruders of either
females dark above and silvery below
algae and bits of debris. The nests
tation and 10-15 cm above the bottom
slightly sunk into the bottom. In some areas, nests have been observed
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under or between rocks at depths as great as 2 m (McKenzie and Keenleyside,
1970). The material is stuck together with a glue secreted by the male's
kidneys. Nests are more or less tubular or barrel shaped, with an opening
at each end.
As the nest is completed, the males become paler dorsally but more
intensely black below, especially on the throat and anterior part of the
abdomen. They now begin to court the females, instead of attacking them.
Courtship consists of a series of quick, head-down dancing movements toward
the female. This may be accompanied by bending the body in an S shape in
front of the female. If the female is receptive, she lowers her tail, dis-
playing her swollen belly to the male. He then dances back towards his nest,
the female following with her snout between the male's pelvic spines. If the
female is not responsive to courting, she may be attacked and driven from the
male's territory. At the nest, the male takes up a position just above it,
with his snout pointing to the opening, and fans with his pectoral fins.
The female enters the nest, but, as she is longer than the nest, her head
and tail protrude. The male then contacts his throat against the side of the
posterior part of the female's body, and quivers. This stimulates her to lay
her eggs, after which she leaves the nest, while the male enters the nest
and fetilizes the eggs. Total fecundity of a female is 350-960 eggs (Andri-
yashev, 1954), but only 50-80 are laid at a time. If the female is still
near the nest when the male emerges, he attacks her and drives her out of
his territory. The female soon begins feeding voraciously and in a day or
so is ready to be courted again by the same or other males and lay another
clutch of eggs. The male usually collects several clutches of eggs from
the females on adjoining territories.
Egg laying and fertilization accomplished, the male again becomes black
all over. He spends most of his time at one or the other entrances of the
nest, in a nearly horizontal position, facing the nest and fanning with his
pectoral fins. This keeps gentle currents of water moving over the eggs,
bringing oxygen and removing metabolic wastes. He also moves the eggs
around in the nest and removes and eats any dead eggs. Sometimes the nest
begins to fall apart before the eggs have hatched. The male may then build
a new nest nearby and transfer the eggs to it (Morris, 1958).
The eggs, which are sticky and 1-1.5 mm in diameter, hatch in about a
week at 18°C. At hatching, the male often enters the nest and removes the
broken remains of the egg membranes. The newly hatched larvae move to the
top of the nest and settle on it. At this time, the male may construct a
nursery, a loose mass of vegetation in the weeds a few centimeters above the
nest. As the young develop and begin to move about, the male catches than
in his mouth and spits them back into the nest or the nursery. As the young
grow older, and become still more active, the male is less and less success-
ful in this, and finally loses interest in them, except, perhaps, as food.
The male may now build a new nest and start the reproductive cycle all over
again, or the breeding season comes to an end (Morris, 1958).
Nothing is known of growth rates of ninespine sticklebacks in north-
western North America. However, judging by the size they attain, they must
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grow considerably faster than those in England, which averaged only 37 mm
(1.5 in) long at three years old (Jones and Hynes, 1970). Sexual maturity
is reached at one year by about half the males and most of the females.
Nearly all are mature by the age of two. Maximum life span is less than
3 1/2 years (Jones and Hynes, 1970).
Nothing definite is known of the migrations of the ninespine stickle-
back. However, there would appear to be seasonal movements inshore to
shallow water in the spring for spawning, and offshore to deep water, or
even to the less saline parts of the sea, by the young and such adults as
survive spawning, in the fall. Shortly after hatching, the young are common
in and just below the zone of rooted vegetation in lakes. Adults are common
down to about 20 m and also at greater depths, 90-110 m, in Lake Superior.
They appear to be most numerous in water of temperatures between 6° and 12°C
(Dryer, 1966; Nelson, 1968b).
Food of ninespine sticklebacks consists mostly of copepods and chirono-
mids, but they also eat cladocera and other small crustaceans, ostracods
and oligochaete worms, as well as their own eggs and larvae upon occasion
{Hynes, 1950; Morris, 1958). In turn, they are fed upon by various preda-
ceous fishes such as Arctic charr, Dolly Varden, lake trout, walleye, gray-
ling, and fish-eating birds (Rawson, 1957; Dryer et al, 1965; McPhail and
Lindsey, 1970).
.IMPORTANCE TO MAN
The ninespine stickleback is of little direct value, as it is too small
and of too uncertain occurrence. However, they may be so abundant at certain
times of year the the natives of Siberia, the Yukon-Kuskokwim delta, and
similar regions, catch them for dog and human food. They have also been used
as a source of oil in some areas (Bigelow and Welsh, 1925; Nikolskii, 1961;
McPhail and Lindsey, 1970).
Indirect importance is probably greater than the direct, for ninespine
sticklebacks may form a significant part of the food for larger fish.
They also make excellent aquarium fishes.
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SECTION 14
SURFPERCHES--FAMILY EMBIOTOCIDAE
The members of this family, numbering about 20 North American species,
are almost all marine. One species lives in fresh water in California and
another, the shiner perch, is mainly marine but often enters rivers.
The surf perches are live-bearers, bringing forth living young rather
than laying eggs. In appearance, they are typically perch-like, with spiny
and soft dorsal fins and three spines in the anal fin. However, the scales
are cycloid rather than ctenoid. A remarkable feature of this group is the
fact that, in at least some species, the males are sexually mature at birth.
Surf-perches are small fishes, rarely exceeding 150 mm {6 in) in length,
They are utilized to some extent for human food or as bait for larger fishes,
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SHINER PERCH
i
Cymatogaster agg regata Gibbons
DISTINCTIVE CHARACTERS
The rather deep, compressed body, large scales and three spines in the
anal fin distinguish the shiner perch from other Alaskan freshwater fishes.
DESCRIPTION
Body compressed, deep, its depth 33-38% of fork length. Head compress-
ed, moderate, its length 27-34% of fork length. Snout length usually
slightly greater than eye diameter, 26-33% of head. Eye moderate, round or
nearly so, diameter 24-301 of" head. Nostrils double on each side. Mouth
terminal, rather small, directed slightly upward, upper jaw reaching backward
to below anterior margin of eye. Small conical teeth present on both jaws,
but absent on vomer and palatines. Gill rakers short, slender, 28-33 on
first arch. Branchiostegals 5 or 6. Lateral line slightly arched, complete,
36-43 pored scales.
Fins
C. VIII - XI, 19-22, usually IX, 20, the spiny portion higher than the
soft-rayed part. A. Ill, 22-25, usually 24. P-,. 19 or 20. P2. I, 5.
Caudal forked.
Scales
Large cycloid. A scale sheath present along posterior three fourths of
base of dorsal fin, separated from body scales by a furrow.
Color
Generally silvery, back dusky to greenish. Middle of sides anteriorly
with groups of fine, black dots on each scale, forming about eight longitudi-
nal stripes. The stripes often interrupted, especially in females, by three
vertical, pale yellow bands. Paired fins colorless. Dorsal and caudal plain
or dusky. Anal usually colorless, sometimes with a yellow blotch anterior-
ly. Breeding males may be almost solid black (Eigenmann and Ulrey, 1894).
163
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Size
Reported up to 203 mm (8 In) (Evermann and Goldsborough, 1907), but the
vast majority are between about TOO and 127 mm (4-5 in). Males are general-
ly smaller than females.
RANGE AND ABUNDANCE
The shiner perch is found along the west coast of North America from San
Quintin Bay, Baha California (Miler and Lea, 1972), north to Wrangell,
Alaska (Bean, 1884). It appears to be rather scarce at the extreme ends of
the range, but is an abundant species from about San Diego to Ketchikan.
Occasionally found in brackish or fresh water (Bailey et al, 1970).
HABITS
The surf perch is viviparous, bearing fully developed young which
develop in the ovarian cavity of the female. Mating is preceded by court-
ship. The male moves slowly toward the female, who retreats. Her flight
stimulates pursuit by up to ten males. During the pursuit, males attempt to
bring their anal regions in contact with that of the female. One male heads
off the female, urging her away from the group of males and also driving
them away. The female is conducted to the shelter of a rock or other
object, where the courtship dance begins. In this dance, the male, with
dorsal fin fully erect, swims by means of his pectoral fins in a figure
eight pattern over the female, undulating in both the horizontal and verti-
cal planes. . The male then stops, faces the female head to head, quivers,
moves his jaws and undulates his dorsal fin. He then moves beside the
female, facing in the same direction, and repeats the movements. Finally,
the male tilts on his side, the female tilts slightly away from him, and
the anal regions are brought in contact with each other. Copulation lasts
less than a second (Hubbs, 1917; Wiebe, 1968b).
Ha'les are mature and ripe at birth. Juvenile females are mated soon
after being born. Shortly before breeding, the males develop fleshy lobes
on both sides of the anal fin. One lobe ends in a tubular intromittant
organ which points more or less forward and downward. Mating takes place in
June and early July in most areas, less commonly in May or August. Sperm is
stored in the female's ovary until about December, when fertilization occurs.
Gestation of the young requires 5 to 6 months and they are born in June or
July, about a year after the parents have mated (Eigenmann, 1894-, Turner,
1938).
The eggs are extremely small, only about 0.3 mm in diameter, and shrink
still further during early development to about 0.2 mm (Eigenmann, 1894).
Fertilization occurs while the eggs are still in the follicles, but develop-
ment does not begin until after their release. Eggs which are not ripe at
the time of fertilization degenerate and may contribute to the nourishment
of the developing embryos. During the course of gestation, the epithelial
cells of the ovary become filled with fluid and slough off. The embryos are
nourished by these cells, as well as by the ovarian fluid itself, which
164
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enter the embryonic alimentary canal through the gill opening, later through
the mouth as well. Respiration of later embryos is accomplished by the
development of highly vascularized tissues along the margins of the dorsal
and anal fins. These are in close contact with vascularized portions of the
ovary and most probably function in the exchange of oxygen and carbon
dioxide. In about half the embryos, one gill opening is invaded by a fold
of tissue from the ovary wall, apparently another respiratory device (Turner,
1952). Brood size ranges from 3 to 36 young. Older and larger females
produce more and larger young and generally give birth earlier in the season
than do smaller, younger fish. The development of sex products and second-
ary sex characters of the male is enhanced by warm temperatures and increas-
ing day-length. Development of egg cells of the female, by contrast, is
encouraged by cool temperatures and decreasing light, although development
of the embryos is promoted by warmth (Eigenmann, 1894; Evermann and Golds-
borough, 1907; Wiebe, 1968a, b; Wilson and Millemann, 1969). Breeding
apparently occurs annually during the life of the individual.
The young fish are 34-44 mm long at birth and are ready to mate at this
time. Males are smaller than females, a relation which continues throughout
life. At one year of age, females average about 93 mm long in Yaquina Bay,
Oregon. At 2 years, they are about 111 mm1, at 3, 118 mm; at 4, 130 mm; at
5, 133 mm; and at 6 years, about 135 mm (Eigenmann, 1894; Wilson and Mille-
mann, 1969).
Except for seasonal onshore-offshore movements, the shiner perch appar-
ently does not migrate. However, it does appear in shallow water inshore
in the summer and has been taken in trawls as deep as 128 n in winter
(Clemens and Wilby, 1946; Hart, 1973).
Food of the shiner perch varies with the size of the individual and the
time of year. The young feed mainly on copepods, while the adults eat
various small crustaceans, molluscs and algae. Adults often eat the append-
ages of barnacles on piers and rocks (Clemens and Wilby, 1946; Gordon, 1965).
IMPORTANCE TO MAN
The shiner perch,is of little importance. Some are marketed in British
Columbia, chiefly for the oriental trade in Vancouver (Clemens and Wilby,
1946) and also in California, especially at San Francisco and Los Angeles.
However, the shiner perch is only a minor constituent of the California
"perch" landings {Roedel, 1948). It is used to a slight extent as bait for
larger fishes and provides considerable sport in the summer, when numbers
are found close inshore, for juvenile anglers. It is said to be "a very
fair pan fish" (Evermann and Goldsborough, 1907).
165
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SECTION 15
SCULP INS—FAMILY COTTIDAE
Sculpins are bottom-dwell ing fishes, mostly marine but some freshwater,
with a few of the marine forms entering rivers, sometimes to considerable
distances upstream. There are at least 300 species in the family. They
have large heads, the gill covers usually armed with sharp spines; robust
bodies tapering to a narrow caudal peduncle; both a spiny and a soft dorsal
fin, a single spine in each pelvic fin but none in the anal; and ctenoid-
type scales, but the scalation is usually much reduced and the scales are
often highly modified.
The sculpins are primarily fishes of the northern temperate and arctic
seas. One genus, Cottus, is circumpolar in fresh water in the northern
hemisphere. Most are fairly small fishes, especially the freshwater forms,
but a few of the marine species may reach lengths of 600 mm (2 ft) or more,
Freshwater sculpins occasionally prey upon the young of various salmonids,
but only rarely does this reach significant levels. In turn, they are eaten
by larger fishes such as lake trout and burbot.
A number of old records of freshwater sculpins in Alaska were based on
mis-identifications. Mr. Kevin M. Howe, Department of Fisheries and Wild-
life, Oregon State University, has recently re-examined the specimens which
formed the basis for Evermann and Goldsborough's (1907) record of the riffle
sculpin, Cottus gulosjjs, from Loring and the Boca de Quadra. Mr. Howe finds
these specimens to be a mixture of coastrange sculpins, Cottus aleuticus,
and prickly sculpins, Cottus asper.
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PACIFIC STAGHQRN SCULPIN
Leptocottus armatus Girard
DISTINCTIVE CHARACTERS
The Pacific staghorn sculpin is set off by the large upper preopercular
spine ending in 3 or 4 sharp, upturned, curved spinules; and the large, dark
spot on the posterior part of the spiny dorsal fin.
DESCRIPTION
Body elongate, more or less cylindrical anteriorly, somewhat compressed
posteriorly. Depth about 17-20% of total length. Head rather flat and
broad, its length about 331 of total length. Snout about 25% of head length.
Eye oval, horizontal diameter greater than vertical, length 13-18% of head.
Mouth moderate, terminal, lower jaw slightly shorter than upper, upper jaw
reaches back to below eye. Small, sharp teeth present in bands on both jaws,
vomer and palatines. Gill rakers reduced to flat, bony plates, each bearing
a cluster of small teeth, 8 or 9 on lower portion of first arch, none on
upper portion. Branchiostegals 6. Lateral line nearly straight, with 37-42
pores, each associated with a small, sub-dermal, cartilaginous plate.
Fins
VI-VIII. D2- 15-20, the spinous and soft parts separated by a
short gap. A. 15-20. P-j. 19. P2. I, 4. Caudal slightly rounded.
Scales
No scales. Skin smooth and naked except for the lateral line plates.
Color
Grayish olive to greenish, occasionally with some yellow, dorsally.
Orange-yellow to white or silvery below lateral line. Spiny dorsal dusky,
with a black spot near tips of last 3 spines, a white band below it. Soft
dorsal dusky, with several oblique white to yellowish bands. Pectorals
yellow with 5 or 6 dark greenish bars. Caudal dusky with one or two pale
bands. Anal and pelvic fins pale.
Size
Reaches about 460 mm (18 in) total length.
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RANGE AND ABUNDANCE
Found along the Pacific coast of North America from San Quintin Bay,
Baha California (Roedel, 1948} to Izembek Lagoon in the southeastern Bering
Sea (Tack, 1970; Paulson and Smith, 1974). Throughout most of its range, it
is a common fish of the sub-tidal zone and tide pools. Even in the Bering
Sea, at the northern limit of its range, it is not at all uncommon in favor-
able locations. Although a marine fish, the Pacific staghorn sculpin can
tolerate freshwater and not infrequently penetrates rivers and even fairly
fast small streams. It has been found in fresh water on Admiralty Island,
Alaska, among other places {Gunter, 1942).
HABITS
The Pacific staghorn sculpin spawns from October to March, but the
majority of reproductive activity takes place in January and February.
Spawning areas are probably most common in bays and estuaries with relatively
stable salinities, but must also exist in the ocean, for very young staghorn
sculpins have been found in are'as far from fresh water inputs. Eggs develop
normally in salinities between 10.2 and 34.3 parts per thousand, with great-
est survival in the range of 17.6-26.4 parts per thousand. The higher of
these two figures is probably close to the optimum salinity (Jones, 1962).
Development to hatching under laboratory conditions requires 9-14 days
at 15°C, with most of the eqgs hatching in between 10 and 12 days. The new-
ly hatched larvae are 3.9-4.8 mm long, with a large, Globular yolk sac
(Jones, 1962). Growth and longevity appear to vary tremendously in differ-
ent localities. Fish from Walker Creek, Tomales Bay (north of Point Reyes,
California) were reported to grow exceedingly fast, reaching an average
total length of 127 mm at one year of age, 156 mm at 2 and 197 mm at 3.
The maximum age observed in this population was three years (Jones, 1962).
By contrast, fish from Puget Sound, near Friday Harbor, reached 10 years of
age, with average calculated total lengths of I, 34 mm; II, 55 mm; III, 80
mm; IV, 101 mm; VII, 174 mm; X, 228 mm (Weiss, 1969). Females appear to
reach greater size and age than do males.
Recently metamorphosed young move from the spawning areas in the estu-
arine regions into completely fresh water, where they remain for up to six
weeks; Because of the extended spawning period, the fish farthest upstream
are usually somewhat older and larger than those in the lower estuary. How-
ever, this-migration into fresh water is not an essential part of the life
history, for staghorn sculpins may spend their entire lives in highly saline
water (MacGinitie, 1935). Adults apparently tend to remain in shallow water
in the lower estuary, or farther offshore. The adult staghorn sculpin is
often^one of the dominant species of the sand flats and in some regions, at
least, is common to as deep as 90 m (Jones, 1962; Tack, 1970; Isakson et al,
1971; Pearcy and Myers, 1974; Paulson and Smith, 1974). This broad distri-
bution reflects the considerable degree to which this sculpin can tolerate
168
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different salinities. This tolerance range goes from essentially zero
(fresh water) to a maximum of about 51-53 parts per thousand, considerably
higher than ordinary sea water (Hubbs, 1947; Erkkila et al, 1950; Carpelan,
1961).
The Pacific staghorn sculpin is essentially a sedentary fish, resting on
or moving slowly about just over the bottom. However, it is capable of rapid
movement over short distances when occasion demands. When violently dis-
turbed, it "rushes rapidly to a new place and buries itself, leaving only the
eyes exposed" (MacGinitie, 1935). This mode of life is reflected in the
food habits. Juveniles in the Walker Creek estuary subsisted almost entirely
on amphipods of the genus Corophium, with nereid worms, gammarid amphipods,
insect larvae and nymphs, and 18 other items of minor importance. Adults in
San Francisco Bay had fed chiefly on four species of bottom-dwelling shrimos,
smelt and crabs, with other fishes and invertebrates occurring in small
amounts in the stomach contents (Jones, 1962).
IMPORTANCE TO MAN
The Pacific staghorn sculpin is of little direct importance to man.
Although it is edible, the large head and slender body result in a small
yield of meat per fish. It is sometimes a nuisance to anglers, greedily
taking baited hooks intended for more desirable fishes. It probably competes
strongly for food with other fishes, especially in confined regions such as
bays and estuaries. It is an important food for herons, loons and cormor-
ants (MacGinitie, 1935).
SLIMY SCULPIN
Cottus^ cojnatus^ Richardson
DISTINCTIVE CHARACTERS
The two pores on the chin and the short lateral line ending under the
second dorsal fin distinguish the slimy sculpin.
DESCRIPTION
Body tadpole-shaped, nearly round in cross-section anteriorly, strongly
compressed posteriorly. Depth 17-21% of total length. Head broad, somewhat
flattened, expanded posteriorly, its length 22-28% of total length. Snout
rounded and broad as seen from above, its length 27-28% of head length.
Nostrils in short tubes, one above middle of upper jaw on each side, .the
other near antero-dorsal margin of eye. Eyes more or less on top of head,
diameter a little less than snout length. Mouth broad, terminal, upper jaw
projecting, maxilla reaching anterior margin of eye or farther back. Fine
teeth on jaws and vomer, none on palatines (teeth rarely present on pala-
tines, according to McAllister, 1964). Gill rakers short and stubby, about
six on first gill arch. Branchiostegals 6. Lateral line incomplete, ending
under middle of soft dorsal fin, usually a few isolated pores behind this,
the main portion with 12-26 pores.
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Fins
DI. VII-X. 02- 14-19, A. 10-14. P-|. 12-16, fins large and fan-
shaped, tips of lower rays exserted. Pp. 1,3, sometimes 1,4, but the
fourth ray much reduced. Caudal roundea.
Scaljs$_
No scales, but small prickles sometimes present behind pectoral fins.
t
Col or
Dark brownish or greenish to dark gray on back and sides, whitish below,
with vague, dark matt!ings or bars below soft dorsal. Belly sometimes with
orange tints. Soft dorsal, pectoral and caudal fins usually barred, pelvics
and anal usually immaculate but sometimes barred. Spiny dorsal dark basally,
with a clear margin which may turn orange in breeding males. Breeding males
usually dark, sometimes black, all over.
Size
Known to reach 121 mm total length (Van Vliet, 1964), but most do not
exceed about 75 rrm (3 in).
RANGE AND ABUNDANCE
The slimy sculpin is distributed all across northern North America, from
eastern New Brunswick, Canada, to western Alaska, including Nunivak and St.
Lawrence islands. On the east coast it is found as far south as Virginia
and north to Ungava Bay, though it has not been recorded from the northern-
most parts of Newfoundland and northwestern Labrador, nor the most northerly
parts of Keewatin. It is present in all the Great Lakes, the upper Missis-
sippi drainage, thence northwesterly across most of Saskatchewan to northern
Alberta (absent from most of southern Alberta). Present in the upper Columbia
drainage and the Fraser River, north through British Columbia east of the
Continental Divide, reaching the west coast in southeastern Alaska. Present
throughout the Northwest Territories (except northeastern Keewatin), Yukon
Territory and mainland Alaska. Also known from the eastern part of the
Chukhotsk Peninsula in Siberia.
Throughout its range it is an abundant species. It inhabits lakes to
considerable depths being most common in Lake Superior at depths between 90
and 110 m. In Lake Michigan it has been found all the way from near shore
to 128 M (Deason," 1939). It is common in streams, particularly those with
fairly fast current and rocky bottoms, where it stays on the bottom, darting
from place to place when disturbed. Its quiescent, bottom-dwelling habits
and cryptic coloration make it difficult to observe until it moves.
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HABITS
Spawning takes place in the spring, shortly after break-up, at water
temperatures between 4.5 and 10°C. This may occur as early as late March in
the southern part of the range to late May or even later in the far north.
Before spawning, males select nest sites, usually under rocks, logs, boards,
etc., in shallow water, from about 2 cm to more than 30 cm deep. The floor
of the nest is often sandy and may be excavated to some extent by strong
swimming or fanning movements on the part of the male. He may also remove
gravel from the nest by picking up small particles in his mouth and moving
them outside. The nest is usually 20-40 mm high, both height and width
depending on the size of the fish. At this time, the male assumes a dark
brown to black color and the orange margin of the first dorsal fin is bright
(Koster, 1936; Van Vliet, 1964).
Nesting males are strongly territorial and aggressive towards other
males. They defend the nest by "barking" (quickly opening and closing the
mouth) at intruders, which may be followed by a quick dart with gill covers
spread and head flattened, and by fighting. In contrast to most fishes,
in which agonistic behavior is usually ritual, the loser of a fight between
two male sculpins is often killed (Koster, 1936; Van Vliet, 1964).
Courtship begins when a ripe female approaches the nest. The male
barks, undulates and twitches his body and fins, then rushes to the side of
the female and drives her into the nest. Within the nest, there is more
barking, as well as butting or biting the female's side near the vent. Both
fish turn upside down and the female deposits her eggs on the roof of the
nest. The male presses her against the roof, which may help in extruding the
eggs, and releases milt (Koster, 1936). After spawning, the female leaves
the nest and shows no further interest in the eggs.
Spawning apparently goes on at any time of the day or night. The female
deposits all of her eggs in a single mass. Fecundity ranges from about 42 to
1420 eggs, the number tending to increase with the size of the female. The
average size of a clutch is 150 to over 600 eggs. Males usually mate with
two or three females in the same nest, but nests have been found with as many
as 10 egg masses (Roster, 1936; Van Vliet, 1964; Craig and Wells, 1976).
The eggs are quite large for so small a fish, 2.5-3.0 mm in diameter,
and are yellow to pink in color. Development to hatching requires about 30
days. During this time the male remains in or near the nest, guarding it
against intruders, fanning the eggs with his pectoral fins and removing
debris and dead eggs. The young are about 7.2 mm long at hatching, with a
large, spherical yolk sac. They seek the bottom of the nest and remain there
for about a week. By that time the yolk has been absorbed, the young are
nearly 11 mm long and they leave the nest to take up the solitary life
characteristic of this species.
Growth rates vary from locality to locality, with one-year-old fish as
small as 36 mm total length in the Cree River, Saskatchewan, or as long as
93 mm at Otter Rapids on the Churchill River (Van Vliet, 1964). In general,
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however, growth seems to be rather slow. Most fish mature in their second
year, although in the Chandalar River of Alaska, 4 years was the usual age
at maturity. Maximum age recorded is seven years, in the Chandalar River
fish (Craig and Wells, 1976).
Except for probable movement into shallow water for spawning, slimy
sculpins apparently do not migrate. It is not known whether the sculpins
found at considerable depths in the Great Lakes breed at those depths or
whether they come in to shore in the spring. The slirry sculpin is quite
sedentary, rarely moving more than a few meters, even when violently dis-
turbed. However they have been seen to travel considerable distances through
the semi-consolidated, jelly-like sediments on the bottoms of some deep
lakes on the Canadian shield (Emery, 1973). In some areas, they are common
in brackish water (Dunbar and Hildebrand, 1952). Presumably, these individ-
uals must migrate to and from fresh water, at least for spawning.
The slimy sculpin is almost exclusively insectivorous in its food
habits. In most areas, the major items are the larvae of Diptera, Ephemero-
ptera and Trichoptera, with Odonata and annphipods sometimes important for
larger individuals. The smaller fish feed most heavily on Diptera, espe-
cially chironomids and simuliids, while larger fish tend to make more use of
the larger items (Kendall and Goldsborough, 1908; Rimsky-Korsakoff, 1930;
Sibley and Rimsky-Korsakoff, 1931; Pate, 1933; Koster, 1936, 1937; Van Vliet,
1964; Petrosky and Waters, 1975; Craig and Wells, 1976). Although the slimy
sculpin has often been accused of devouring the eggs and young of trout and
other salmonids, it seems that this happens but seldom. Fish are rarely
eaten, and young sculpins,-as well as sculpin eggs, are taken far more fre-
quently than are other fishes. Sculpins do, on occasion, eat larval trout;
however, salrnonid eggs are too well protected by the redd to be vulnerable
to sculpins large enough to eat them and the eggs are too large to be eaten
by sculpins small enough to burrow into a redd (Koster, 1936, 1937; Kogl,
1965; Clary, 1972; Petrosky and Waters, 1975; Craig and Wells, 1976).
IMPORTANCE TO MAN
The slimy sculpin is of no direct importance to man. In some places it
may be used as bait for larger fishes, but even this seems to be rare. In-
directly, however, it may be more significant. Although this sculpin is
almost exclusively a bottom feeder and the salmonid fishes with which it
frequently co-exists are mostly mid-depth and surface feeders, the two groups
feed on much the same groups of organisms at different developmental stages.
In a Minnesota stream, it has been found that slimy sculpins made up about
1/3 of the total annual fish production, with three trouts (Salvelinus fonti-
nalis, Salmo gairdneri and Sal mo truttaj contributing the remaining 2/3. In
the Chatanika River, Alaska, sculpins and grayling share many of the same
food items. It is conceivable that a large population of sculpins could
compete strongly with the salmonids and limit the production of the latter,
although this matter has not been proven (Schallock, 1966; Vascotto, 1970;
Petrosky and Waters, 1975; Craig and Wells, 1976).
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PRICKLY SCULPIN
Cottus asper Richardson
DISTINCTIVE CHARACTERS
The complete lateral line, a single pore at the tip of the chin, 15-19
anal rays, and well developed palatine teeth distinguish the prickly sculpin.
DESCRIPTION
Body elongate, broad and nearly cylindrical anteriorly, compressed pos-
teriorly. Depth 16-18% of total length. Head large, its length 25-32% of
total length. Snout 21-27% of head length. Eye round, its size variable,
18-33% of head length. Two nostrils on each side, each in a short tube,
anterior nostril near edge of upper jaw, posterior close to anterior margin
of eye. Mouth large, maxilla reaches middle of eye or further back in
adults. Teeth small but well developed, present on both jaws, palatines and
voner. Gill rakers short, 5-6 on first arch. Branchiostegals 5-7 (usually
6) on each side. Lateral line complete, with 32-43 pores. Pyloric caeca
short, 4-6.
Fins
Pa-
scal
D.
I,
es
4
VII to X
(rarely
3
18
or
to
5).
23. A.
Caudal
15-19. P-|. 15-18, large and fan-shaped.
truncate or slightly rounded.
No true scales. Prickles on body, distribution ranging from covering
most of the skin (except on abdomen) to a small patch behind base of pectoral
fins.
Color
Dark brown, olive or gray above and on sides, whitish yellow to white
below. Usually three dark, irregular blotches or bars below soft dorsal fin.
Vague, irregular, dark marks on sides. Fins (except anal) with dark bars,
first dorsal with a dark spot posteriorly. Both sexes show an orange band
on the edge of the first dorsal at spawning time.
Size
The largest species of the genus Cottus, said to reach about 300 mm (1
ft) (Jordan and'Evermann, 1896), but the majority generally not over 150 mm.
However, the original specimen, on which the description was based, was 9 1/2
inches (241 mm) (Richardson, 1836) and Jordan and Gilbert (1882) listed 10
tnches (245 mm) as the maximum length.
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RANGE AND ABUNDANCE
Known from the Ventura River, California, in the south, then north in
coastal streams of the Pacific coast to Seward, Alaska. Ranges well inland
in the San Joaquin and Sacramento rivers of California, the northern part of
the Columbia River drainage (but not the Snake River) as well as in the
Fraser and other major Pacific coastal rivers of British Columbia (Krejsa,
1967a). Isolated populations are present in the Peace River drainage in
British Columbia as far downstream as McLeod Lake (Lindsey, 1956). An abun-
dant form wherever present. The record of £. gulps us from southeastern
Alaska was based on a mixed sample of C_. as per and C_. aleu_t_i_c_iis (Kevin Howe,
Department of Fisheries and Wildlife, Oregon State University, pers, coirm.j-
HABITS
The prickly sculpin exhibits at least two genetically distinct forms.
The inland form is generally more densely prickled over a larger portion of
the body and does not migrate to estuaries to spawn, while the coastal form
shows reduced prickles and spawns in brackish water (Krejsa, 1967b). Except
for the difference in spawning localities, the life histories of the two are
quite similar, as far as is known.
Spawning may occur as early as February in the south and as late as July
in the north. Prior to breeding, the males move downstream and select nest
sites under cobbles, large, flat rocks, etc. The females aggregate some
distance upstream, then move down singly to the spawning area. They display
to the,male,,who courts a female and brings her to his nest. Further court-
ship goes on in the nest. The orange eggs, somewhat less than 1 nm in diam-
eter, are sticky and adhere to the roof of the nest. Fecundity ranges be-
tween 280 and 7,410 eggs per female (Patten, 1971)." Since males may mate
with as many as ten females, there may be 25,000-30,000 eggs in a single
nest (Krejsa, 1967b; Ringstad, 1974). After spawning, the female goes back
upstream to feed, while the male remains at the nest, fanning and guarding
the eggs. He does not feed until the eggs have hatched. Spawning takes
place in water temperatures of 8-13°C. Development to hatching requires
15-16 days at 12°C, but about 3 weeks at 8°C (Krejsa, 1967b).
Within a few hours of hatching, the larvae, about 5 mm long, swim up to
the surface, where they remain in a planktonic existence for the next 30-35
days. They are positively-rheotactic and swim actively, in short bursts,
against currents greater than 1 cm/sec. They are probably swept up and down
the estuary by tidal currents, but any carried out to sea doubtless perish,
as none have been taken in the ocean. However, in late spring and early
summer, they are numerous around and below the spawning grounds (Krejsa,
1967b; Pearcy and Myers, 1974; Mason and Machidori, 1976). Larvae hatched
upstream in freshwater may be carried into a lake or estuary during this
phase of existence (McLarney, 1968).
During the planktonic stage of life, the larvae of the freshwater, non-
anadromous form in lakes show distinct diurnal vertical migrations. They
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are most abundant at the surface during the darkest hours of the night,
between 2100 and 0400 hours (9PM-4AM). During the day and on bright moon-
light nights, they apparently stay deep (Sinclair, 1968).
Metamorphosis has been completed by the end of the planktonic period
and the little fish take up a demersal mode of life. The adults move back
upstream during the late summer, followed by the young in the early fall,
although the young may remain in the estuary for a full year. Upstream move-
ment is usually no more than a few miles and may be blocked by relatively
small obstructions such as falls of 30 cm (1 ft) height (Krejsa, 196?b; Ring-
stad, 1974; Mason and Machidori, 1976).
Yearling fish (and possibly older ones also) have been observed to form
schools in shallow water in lakes (Northcote and Hartman, 1959). Whether
this is related to feeding or represents a remnant of their pelagic life or
has some other explanation is not known.
The prickly sculpin grows rather slowly and shows wide variations in
growth rates within and between year-classes, as well as from place to place.
Some approximate lengths at different ages for fish from Oregon and Vancouver
Island, B. C., are, 1+ 45-46 mm; 2+, 52-65 mm; 3+» 71-89 mm; 4+, 85-98 mm.
The establishment of typical lengths at different ages is difficult because
of the fact that this sculpin appears to make about 50% of the entire year's
growth in only two months, July and August. Hence, the time of collection
makes a great difference in the observed average lengths (Bond, 1963; Ring-
stad, 1974; Mason and Machidori, 1976).
The oldest specimen reported was a 7 year old fish from Oregon (Bond,
1963). If the very large specimens (240-300 mm) reported in the literature
grew at the rates shown here, they must have been at least 13-17 years old.
Food of the prickly sculpin consists mainly of invertebrate animals of
various sorts; as would be expected, these are mostly bottom-dwelling organ-
isms and the developmental stages of insects. The list is a long one, in-
cluding cladocerans, copepods, ostracods, amphipods, molluscs, mites, at
least seven orders of insects, and small fishes. Among the insects, the
Trichoptera, Ephemeroptera and Diptera seem to be the most important. How-
ever, the relative ranking of food items varies with season and locality and
depends largely on availability. In some areas, salmon eggs and fry, sock-
eye, pink and chum, and even year old sockeye are eaten in considerable
numbers. Up to 111 sockeye fry have been found in the stomach of a single
prickly sculpin. This sculpin also eats other fishes, and even its own kind.
One was found which had swallowed another sculpin more than half as long as
itself (Pritchard, 1936b; Ricker, 1941; Robertson, 1949; Northcote, 1954;
Hunter, 1959).
IMPORTANCE TO MAN
The prickly sculpin is of no direct importance to man. It is too small
to be used as food and too difficult to capture in large numbers to be used
for anything else. Indirectly, however, this species may be of considerable
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importance as a predator on the young of more valuable species and as food
for the adults. It is an important item in the food of several fish-eating
birds such as mergansers.
CQASTRANGE SCULPIN
Cottus aleuticus Gilbert
DISTINCTIVE CHARACTERS
The coastrange sculpin is distinguished by having only a single pore on
the tip of the chin, no palatine teeth, and no pronounced gap between the
first and second dorsal fins.
DESCRIPTION
Body elongate, rather round anteriorly, compressed posteriorly; depth
about 18% of total length. Head moderate, 20-23% of total length. Snout
27-33% of head. Eye small, round, its diameter less than snout length.
Nostrils two on each side, well separated, each in a short tube, posterior
nostril at antero-medial margin of eye. Mouth moderate, broad, -naxilla
reaching almost to below anterior edge of eye. Teeth small but well develop-
ed on both jaws and on vomer, absent from palatines. Gill rakers short, 5-7
on first arch. Branchiostegals 6. Lateral line complete, 34-44 pores,
generally with a marked flexion on caudal peduncle. Pyloric caeca 3-6.
Fins
D. VIII-X. 17-20. A. 12-15. P^. 13-16, usually 14. P2. I, 4.
Caudal fin truncate.
Scales
No scales. Skin smooth except for a small patch of prickles behind
each pectoral fin.
f
Color
Dark brown to greenish or grayish on back and sides, with darker
blotches. Sides lighter, ventral areas white. Usually two or three dark,
saddle-like blotches below soft dorsal fin. Dorsal, anal, pectoral and
caudal fins with dark bars. Spiny dorsal of spawning males with orange edge.
Si ze
Up to 172 mm (6 3/4 in) in southern California according to Hubbs
(1921), but generally not over about 115 mm (4 1/2 in) and most are between
50 and 100 mm (2-4 in).
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RANGE AND ABUNDANCE
From Peidras Blancas Point, California, north in coastal streams to
Bristol Bay, Alaska, westward in the Aleutians to Kiska Island. An isolated
population is found in the Kobuk River, Alaska, some 800 km north of the
nearest Bristol Bay fish. Apparently isolated populations also are present
in Lake Washington, Washington, and Cultus Lake, British Columbia, perhaps
also in other lakes of that general area (Ricker, 1960; Ikesumiju, 1975;
Larson and Brown, 1975). The coastrange sculpin is one of the more abundant
small fishes throughout its range.
HABITS
Spawning generally takes place in the lower reaches of the streams and
in the estuaries, but may occur well up stream as well. The eggs are depos-
ited in clumps and clusters in crevices between rocks or (more often) on the
underside of logs, cutbanks, rocks, etc. (Ringstad, 1974; personal observa-
tion). The eggs are yellow to orange, sticky, about 1.24, to 1.91 mm in
diameter, and are usually present in at least two distinct groups of differ-
ent stages of development. Spawning behavior has not been observed, so it is
not known whether the clumps represent several spawnings by a single female
or single spawning by several females, but judging by the numbers of eggs
per nest, it is almost certainly the latter. Over 7,000 eggs have been
found in a single nest, whereas fecundity ranges from 100 to 1,764 eggs per
female (McPhail and Lindsey, 1970; Patten, 1971). The male guards the nest
until all eggs have hatched.
Spawning occurs in the spring, as early as mid-February to as late as
August in various areas, but mostly in May and June, in water temperatures
around 6-8°C (Ikesumiju, 1975). Time required for development to hatching
is 19-20 days at 10-12°C under laboratory conditions (Mason and Machidori,
1976). Larvae are about 5 mm long at hatching and swim to the surface within
a few hours. The young are planktonic, floating near the surface, and are
carried downstream to the inter-tidal portions of estuaries or out into open
waters of large lakes. Feeding begins 6-10 days after hatching, by which
time the yolk is noticeably depleted and most of the larvae have drifted down
to the lake or estuary (Mason and Machidori, 1976). Young coastrange scul-
pins may be 20-30 mm long in the intertidal zone by mid-August (Heard, 1965;
McLarney, 1968). In Carnation Creek, British Columbia, they average 41.5 mm
long at age 1 (in August), 57.0 mm at age 2, 70.0 mm at age 3 and 79.5 mm at
age 4. Sexual maturity is achieved in the third year (Ricker, 1960; Patten,
1971). Maximum age recorded is 8 years, although in Oregon they apparently
do not survive beyond 4 years (Bond, 1963; Mason and Machidori, 1976).
The coastrange sculpin undertakes regular seasonal migrations. The
adults move downstream to estuaries or at least the lower reaches of rivers
in the spring, moving mostly at night. This migration is associated with
spawning, although in some streams spawning takes place in the normal winter
habitat upstream (McLarney, 1968; Mason and Machidori, 1976). A reverse
upstream migration of yearlings and adults occurs in late sunnier to early
winter, August to December. The young fish apparently spend nearly a full
177
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year in the downstream habitat. The winter is spent in the upper parts of
the streams (McLarney, 1968; Ringstad, 1974). Where coastrange sculpins co-
exist with prickly sculpins, in the lower portions of streams, the latter
occupy sheltered areas of low current velocities, while the former are found
more often on the heads and tails of riffles in swifter water. In the
absence of prickly sculpins, especially upstream, the coastrange sculpin
occupies both swift and quiet water (Ringstad, 1974). The coastrange sculpin
is generally solitary, but large aggregations have been seen. It is not un-
common to find two or three individuals in the general vicinity of a nest
(Greenbank, 1957; personal observations).
Food of the coastrange sculpin consists mainly of benthic insect larvae
and nymphs. However, depending on the season of the year and the time of
,day, feeding habits vary. Feeding is most intense at night. During daylight
hours in summer, Trichoptera larvae dominate the food. In late evening and
early morning, Plecoptera and Ephemeroptera nymphs and chironomid larvae are
important. In some streams, this sculpin preys heavily on pink salmon fry.
Other foods include ostracods, mysids, cladocerans, amphipods and isopods.
They can survive without food for 94 days or more, but lose about 33% of
their weight in such circumstances (Hunter, 1959; Sheridan and Meehan, 1962;
Ringstad, 1974; Ikesumiju, 1975).
IMPORTANCE TO MAN
The coastrange sculpin is of no direct importance. However, it may be,
in some situations, a significant predator on salmon fry of several species,
and has been shown to compete for food, sometimes severely, with coho
salmon fry (Ringstad, 1974). In turn, the planktonic larvae of this sculpin
are eaten by juvenile sockeye salmon and the adults may be important food
items for larger predatory fishes such as Dolly Varden.
FOURHORN SCULPIN
Myoxocephalus quadricornis (Linnaeus)
DISTINCTIVE CHARACTERS
The fourhorn sculpin is identified by the four bony protuberances on
the top of the head (smaller in females and young and absent in the fresh-
water form); the chain-like lateral line; the sharp, straight spines on the
preoperculum; and no palatine teeth.
DESCRIPTION
Body tadpole-shaped, tapering posteriorly. Depth about 25% of total
length. Head moderately large, 27-30% of total length, broad and flattened,
its width notably greater than its depth. Four bony or warty protuberances
on top of head, two between the eyes and two on occiput. Four preopercular
spines on each side; upper spine long, straight and sharp, longer in males
than in females; the two lower spines short and pointing downward. Snout
broad, long, 26-30% of head length. Eye small, 60-80% of snout length.
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Mouth large, terminal, lower jaw included, maxilla reaching to below middle
of eye or farther back. Jaws and vomer with bands of small teeth. Gill
rakers reduced to rounded, spiny protuberances, 7-8 on first arch. Branchio-
stegals 6. Lateral line chain-like, with about 45 pores, each with a small
embedded plate.
Pi ns
D. VII to IX - 13 to 16, the spiny and soft fins separated by a space
about equal to eye diameter. Soft dorsal often much enlarged in adult males.
A. 13-17. P-j. 15-18. P2. I, 3-4. Caudal truncate to slightly rounded.
Scales
No true scales. Lateral line with embedded, bony plates. Sides above
and below lateral line with several irregular rows of round, bony, prickly
plates, these best developed and most numerous in adult males, noorly de-
veloped or absent in females and young.
Color
Young are gray above, with three or four dark saddles
and a dark spot on dorsal side of caudal peduncle. Adults
dorsal dusted with black, soft dorsal, anal, pectorals and
bars. Pelvics pale.
below dorsal fins
are darker. Spiny
caudal with dark
Reported to reach a length of 365 mm (Andriyashev, 1954} but most adults
do not exceed about 280 mm and a weight of about 260 grams.
*
There are two distinct forms of M_, quadricornis , the marine, brackish
water and riverine form described here, and the dwarf, strictly lacustrine
form, which is often considered a separate subspecies, M_. quadricornis
thomsoni . In Canada, this lacustrine form is called Deepwater Sculpin. It
differs from the marine type chiefly in lacking the bony protuberances on the
head and the spiny plates on the sides and in its small size. The deepwater
sculpin has not yet been found in Alaska, but may be present in some of the
deep, unexplored lakes of the Arctic.
RANGE AND ABUNDANCE
Circumpolar in cold, brackish and moderately saline water. In Europe,
from the Baltic eastward along the Siberian coast to the Chukchi and Bering
seas, south in the Bering Sea to the Anadyr River. From north-western Green-
land westward along the arctic coast of Canada and the Canadian Arctic Archi-
pelago, the north coast of Alaska and south in the eastern Bering Sea to
Norton Sound and St. Lawrence Island. Ascends rivers for considerable dis-
tances. It has been found in the Meade River, Alaska, as much as 144 km
(90 mi) upstream, and 192 km (120 mi) up the HacKenzie River in Canada. It
is an abundant species, as far as is known.
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HABITS
Most of the knowledge of the biology of the fourhorn sculpin comes from
studies made in the Baltic Sea. Presumably, the life history of the fish in
Alaska is not very different.
Spawning takes place from mid-December through January at depths of 15-
20 m and water temperatures of 1.5-2°C. With the approach of the spawning
season, males become territorial and aggressive towards other males. When
an intruding male appears on the territory, the resident male advances to-
ward him in quick, short darts, stopping 3-5 cm from the intruder with his
dorsal and pectoral fins spread. At the same time, the resident produces a
low-pitched humming sound, with a frequency of 125 cycles per second. The
intruder usually lowers his dorsal fin and turns away. If he does not, the
resident male bites him.
Two or three days before spawning, the females become restless, swimming
about with short periods of resting. When a female swims over a male, he
undulates his body laterally and raises his spiny dorsal fin. If the female
comes to rest near the male, he moves close to her in a series of short
darts. Both sexes fan rhythmically with their pectoral fins, sometimes for
hours, resulting in a hole 20-25 cm in diameter and 10-15 cm deep in the
soft bottom. Finally the male and female lie side-by-side, with the male's
caudal fin twisted under that of the female. The male makes rhythmic
motions toward the female with his tail, which apparently stimulates her to
release her eggs (Westin, 1969, 1970b),
The eggs are adhesive and are extruded in a single clump. They are
quite large, 2.4-2,9 mm in diameter and the color varies from greenish to
bluish to yellow. At one time, the different egg colors were thought to in-
dicate genetic differences (Svardson, 1958; 1961) but subsequent investiga-
tion has shown that the colors of the eggs are controlled by diet (Nyman and
Westin, 1968, 1969; Westin, 1968a, c). As would be expected with such large
eggs, fecundity is low, 792-6,150, but in some populations up to almost .
18,000 eggs (Andriyashev, 1954; Westin, 1968a).
After spawning, the female leaves and shows no further interest in the
eggs. The male, however, remains with the egg mass, guarding it and attack-
ing any fish that comes within about 30 cm. He generally lies close to the
eggs, fanning them with his pectoral fins, and occasionally picking them up
in his mouth and spitting away the dead ones. This cleaning is essential to
successful hatching, as without it the eggs soon become infested with fungi
and die. The male guards and cares for the eggs until they hatch, and does
not feed during this period. Hatching requires 97 days at 1.5°C, 74 days
at 2°C or 55 days at 4.7°C. At 10.5DC, all eggs die before hatching
(Westin, 1969).
Fry hatch in the spring and reach a length of 20-24 mm by August.
Little is known of subsequent growth except that it is slow. Fish one year
old are 40-55 mm long, two-year-olds are about 97 mm, at 5-6 years only
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210-240 mm long, those of 7-8 years are 240-270 mm, and at 10 years the
average length is about 300-310 mm. Sexual maturity is first reached at 4-5
years of age, most fish being mature at 6 years (Andriyashev, 1954; Lukyani-
chov and Tugarina, 1965).
The fourhorn seulpin is most active in the daytime during the winter
(November-April) but is largely nocturnal the rest of the year (Viestin,
1971). It is quite strictly a cold-water fish, although under proper condi-
tions of acclimation it can stand temperatures up to 25.5°C. In its adaption
to very cold temperatures, around 0°C, it develops an organic "antifreeze" in
the blood. At low temperatures, about 4°C, it adapts more readily to slight
rises of temperature than to equal falls (Westin, 1968b; Oikari and Kristof-
fersson, 1973),
The fourhorn sculpin lives permanently near the coast. Barring short
onshore-offshore seasonal movements and mass movements of fry into shallow
water in summer (Andriyashev, 1954; Westin, 1970a) there are no migrations of
large numbers of fish. Movements into fresh water and long distances up
rivers are apparently undertaken by relatively few individuals at a time.
Food of the fourhorn sculpin consists primarily of invertebrates of various
sorts. The list includes priapulids, mysids, isopods, amphipods, molluscs,
small fishes and fish eggs, the last most commonly the eggs of its own
species. In the Baltic, the most important individual items are the isopod,
Mesi dotea entomon, and the amphipod, Ppntopore i a a f f i n1s. It is the latter
which gives the characteristic greenish orbluish color to the eggs. Fish
living on diets which do not include Porvtoj)o_reia have yellow eggs (Andriya-
shev, 1954; Westin, 1968c, 1970a). In other areas, mysids may compose as
much as 90i of the food (Lukyanichov and Tagarina, 1965).
Almost nothing is known of the biology of the deep water, lake-dwelling
form. It lives at such great depths that its presence is often known only
from the examination of stomach contents of deep-feeding carnivores like
burbot and lake trout. It is present in the Great Lakes and in a number of
deep lakes in Canada. It is most abundant at depths below about 75 m in the
Great Lakes, at 25^37 m in Great Slave Lake. Its diet includes mysids, cope-
pods, amphipods and chironomid larvae and pupae. This is a dwarf form. The
three largest on record are a specimen 199 mm long from Lake Ontario and two
of 103 and 109 mm from Lake Michigan. Similar and perhaps identical sculpins
are known from a number of lakes in Europe. The forms in lakes which were
most recently cut off from the sea, such as those on Victoria Island, Canada,
and Lake Ladoga in Russia, are morphologically intermediate between those in
older lakes and those in the sea (Deason, 1939; Rawson, 1951; Martin and
Chapman, 1965; Dryer, 1966; Delisle and Van Vliet, 1968; Me Phail and
Lindsey, 1970; McAllister and Ward, 1972; Dyatlov, 1974).
IMPORTANCE TO MAN
The fourhorn sculpin is of no direct importance. Although it is edible
and is used for food in some regions of the arctic coast, its distribution
far from centers of population and the small yield of meat per fish militate
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against any high degree of utilization. The deep water lacustrine form is,
where available, one of the first fish foods eaten by young lake trout, and
is a minor constituent of the diets of adult lake trout and burbot (Andriya-
shev, 1954; Rawson, 1961).
SHARPNOSE SCULPIN
Cllnocottus acuticeps (Gilbert)
DISTINCTIVE CHARACTERS
The flattened, tri-partite anal papilla and the cirri on eyeballs, head,
lateral line and at the tip of each dorsal spine will distinguish the sharp-
nose sculpin from its relatives which occur in fresh water.
DESCRIPTION
Body elongate, slender, slightly compressed. Depth 15-19% of total
length. Head rather small, about 23% of total length. Snout sharp, short,
its length about 251 of head length. Eye nearly round, diameter about equal
to or a little less than snout length, a slender cirrus on upper part of
eyeball. Mouth terminal, upper jaw reaching to a vertical through anterior
edge of pupil. Small teeth in bands on both jaws. Gill rakers reduced to
low, smooth mounds, about six on lower limb of first arch, none on upper
limb. Branchiostegals 6. Lateral line high anteriorly, recurved downward
to middle of sides, straight posteriorly, 33-36 pores on body, often one or
two more on tail, each of the anterior 15 or so pores with a slender cirrus.
Anus located well forward, about 1/3 the distance from pelvic base to anal
origin, with a broad, flattened, tripartite anal papilla.
Fins
D. VII to
rays before and
anal. PC
IX - 14 to
behind it.
16.
"l
I» 3» short. Caudal rounded.
A. 10-13, next to last ray longer than the
13, reaching to or just beyond front of
Sales
No scales. On each side, one unbranched cirrus at inner side of base of
nasal spine, one on upper part of eyeball, one above eye, two or three behind
this on head, one or two at angle of gill opening, one above each of the
anterior pores of lateral line, one at tip of each dorsal spine.
Color
Varies with the environment. Sometimes nearly uniform bright green,
more often green to light brown above with dark, wedge-shaped saddles, broad-
er below, on upper sides, with lighter color between. Sometimes a dark,
longitudinal stripe along or below lateral line, sometimes interrupted with
182
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light spots. Ventral region creamy to white. Three dark lines radiating
from eye, one to snout, one downward to behind mouth, third posteriorly to,
base of preopercular spine. Spiny dorsal with a dark blotch between first
and third spines. Pelvic fins plain, others usually dusky, mottled or indis-
tinctly barred.
A very small fish, maximum size about 63 rm (2 1/2 in) .
RANGE AND ABUNDANCE
The sharpnose sculpin is known from the Big Sur River, California, north
along the coast to Alaska and westward along the Aleutian chain to Attu
Island. It has been recorded from the Bering Sea side of Unimak and Unalaska
islands (Wilimovsky, 1964;; MUler and Lea, 1972). Within this range it is
a common member of the inshore rock/algae community, especially in sheltered
tide pools in the upper intertidal zone (Green, 1971). At Amchitka Island,
it is most abundant from late spring to early fall (Isakson et al , 1971).
It occasionally ventures into fresh water.
HABITS
Virtually nothing is known of the biology of the sharpnose sculpin. It
spawns in the spring and summer, for breeding has been observed at the San
Juan Islands on 4 July and a ripe -female was taken on 28 April. The eggs
are brown, about 1 mm in diameter (Hart, 1973).
At Amchitka Island in the Aleutian chain, the sharpnose sculpin is most
abundant near shore in warm weather (Isakson et al » 1971). Presumably, it
moves into deeper, more sheltered areas to escape the rigors of winter.
Judging by its size and habitat, the sharpnose sculpin probably feeds
on small invertebrates so common among the sea weeds and tide pools. How-
ever, nothing is known on this subject.
IMPORTANCE TO MAN
The
is known
UANLt iU MAN
The sharpnose sculpin is a small, zoological curiosity, but as far as
lown it is otherwise of no importance.
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SECTION 16
FLOUNDERS—FAMILY PLEURONECTIDAE
The flounders are a remarkable group of fishes in which both eyes are
on the same side of the head. The young are symmetrical at hatching and
swim in an upright position. During larval development, one eye, usually
the left, moves over the top of the head and comes to rest beside the other
eye. The fish settles to the bottom, blind side down, and takes up a demer-
sal mode of life. The eyed side is colored, usually brownish or greenish
brown, and the blind side is white. The dorsal and anal fins are long,
extending almost the entire length of the body, and have no spines.
Flounders are all marine, but a few species enter fresh water. Several
species are highly important food fishes and are the objects of commercial
fisheries. The group is of world-wide distribution in the ocean, but the
greatest numbers of species and individuals are to be found in the northern
hemisphere.
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ARCTIC FLOUNDER
Llopsetta glacial is (Pallas)
DISTINCTIVE CHARACTERS
Both eyes on the same side of the head, the body covered with typical
scales and the generally plain coloration distinguish the Arctic flounder.
DESCRIPTION
Body strongly compressed, broad, its depth 38-46% of total length. Head
short, 22-26% of total length. Snout short, its length 14-18% of head. Eye
a little longer than wide, diameter about equal to snout length, right
(lower) eye a little before left. Nostrils in short tubes. Mouth small,
oblique, asymmetric, jaws of blind side a little longer than on eyed side,
anterior edge of lower eye. Teeth incisor-like,
teeth on eyed side, 12-21 on blind side; lower
Gill rakers 9-15 on first arch. Branchiostegals
5-6. Lateral line with a very low curve above pectoral fin, straight behind,
and a short, accessory branch on head; 73-76 pores in main portion. In
males, the lateral line is complete, but in females as large as about 200 mm
it is an open groove on the posterior part of the body (Walters, 1955).
maxilla reaching about to
small; upper jaw with 3-6
jaw with 4-16 and 15-22.
Fins
D. 50-62, most anterior ray above eye. A. 35-44, an embedded forward-
pointing spine before first ray.
rounded.
Scales
Pj. 9-11 (rarely 12). P2. 6. Caudal
Weakly ctenoid on both sides of the body in males, with cycloid scales
on the abdominal region of the blind side. Females usually with cycloid
scales over most of the body and narrow bands of ctenoid scales along the
bases of the dorsal and anal fins. In some populations the females are
scaled like the males (Norman, 1934; Andriyashev, 1954; Walters, 1955).
Color
Eyed side dark olive green to brown,
dots or indefinite dark blotches. Blind
ping over from the eyed side. Fins pale
ish tinge or traces of dark spots.
sometimes with scattered black
side white, rarely with brown lap-
brownish, sometimes with a yellow-
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Size
One of the smaller flounders, only rarely reaching 350 mm (13 3/4 in).
Most are much smaller than 350 mm (10 in).
RANGE AND ABUNDANCE
The arctic flounder is almost circumpolar in its distribution. It is
found in the Arctic Ocean from Queen Maude Gulf in arctic Canada westward
along the coasts of North America and Siberia to the White Sea and the
Barents Sea. It is present in the Chukchi and Bering seas southward to
Bristol Bay on the American side and to the northern part of the Sea of
Okhotsk on the Siberian side. It is a fish of the coastal waters, not found
far offshore. In some places it is very abundant, at least inshore in
summer. It often enters rivers (Jordan, 1884; Bean, 1887c; Andriyashev,
1954; Walters, 1955; Nikolskii, 1961; Alverson and Wilmovsky, 1966).
HABITS
As with so many arctic fishes, little is known about the arctic flounder.
Spawning usually takes place in coastal areas in January-March, but may be as
late as May in some regions. Spawning goes on in shallow water at depths of
5-10 m in regions of pronounced tidal currents, temperatures below -1°C and
salinities of about 27-28 o/oo. Fecundity is relatively low for flounders,
only 31,000-230,000 eggs per female. The fish usually reach maturity at 4-5
years of age, although maturity as early as 2 years has been reported. Spawn-
ing is said to occur only every second year. Females grow more rapidly than
males, but growth is slow in both sexes. In the Barents Sea, five year old
fish average only 177 mm long for males, 199 mm for females, and nine year old
females are about 249 mm long. Arctic flounders appear to move offshore in
the fall and onshore in spring. They move close inshore in the evenings,
especially on a rising tide. Their food is mainly small molluscs, crustaceans
and fishes (Turner, 1886; Bean, 1887c; Probatov, 1940; Esipov, 1949; Andri-
yashev, 1954; Nikolskii, 1961).
IMPORTANCE TO MAN
The arctic flounder is of little importance. Small quantities are
taken commercially insthe eastern parts of the Barents and White seas. In
Alaska, this species was formerly (and perhaps still is) utilized by coastal
subsistence fisheries, especially eaten raw. Opinions of early workers vary
as to its utility. Jordan (1884) wrote that "...its great abundance and fine
flavor make it an important article of food." By contrast, Turner (1886)
found it "not palatable" and noted that, in sumer, the fish often developed
repulsive tumors along the fins.
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STARRY FLOUNDER
Platichthys stellatus (Pallas)
DISTINCTIVE CHARACTERS
The starry flounder is set off by both eyes on the same side of the
head, the dorsal and anal fins marked with dark and light (white to orange)
bars, and especially by the stellate, bony tubercles scattered over the
body.
DESCRIPTION
Body strongly compressed, asymmetrical, color on one side only. Depth
40-48% of total length. Head moderate, 23-26% of total length. Snout
rather pointed, 15-18% of head length. Nostrils in short tubes, one at
antero-dorsal margin of lower eye, the other just anterior. Eye small,
diameter about equal to snout length or a little longer. Both eyes on same
side of head, either right or left. Mouth terminal, oblique, lower jaw pro-
truding. Teeth incisor-like, about 10 on eyed side of upper jaw, 15 on blind
side, 12 and 16 on lower jaw. Gill rakers 6-11 on first arch. Branchioste-
gels 4. Lateral line with a slight arch over pectoral fin, 63-78 pores.
Pyloric caeca usually 2 in North American specimens, sometimes 3 or 4;
usually 4 in Asian fish.
£i_ns_
D. 52-66, origin over middle of upper eye. A. 38-47, with a sharp,
forward-pointing spine (often buried in skin) before first ray. P-j. about
11, the fin bluntly pointed. Pp. 6. Caudal slightly rounded.
Scales
Stellate, tuberculate, with well developed spines, scattered over head
and body and in a row along bases of dorsal and anal fins. Tubercles more
numerous on eyed side than on blind side, and increasing in number with size
and age. Embedded cycloid scales present between the stellate tubercles, but
their distribution seems to be quite variable (Norman, 1934; Orcutt, 1950;
Batts, 1964).
Color
Eyed side dark brown to nearly black, sometimes with indefinite blotch-
ings. Alaskan specimens sometimes have a greenish tinge. Blind side white
to creamy. Dorsal with 4-7, anal with 4-6 dark bars with white to orange
spaces between. Caudal with 3 or 4 dark longitudinal bars on its posterior
part. Rather rarely, the blind side may be partly or completely colored
like the eyed side; or white may be present on the eyed side, creating a pie-
bald effect (Gudger, 1941; Follett, 1954).
187
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Size
One of the largest, if not the largest, of the true flounders. Report-
ed to reach nearly 1 m in length and 9.1 kg weight.
RANGE AND ABUNDANCE
The starry flounder ranges along the coast of North America from around
Santa Barbara, California, north to the Arctic Ocean, then east along the
Arctic coast of Canada to Bathurst Inlet, possibly even as far east as Queen
Maude Gulf (Walters, 1955). To the west, starry flounders are found along
the Chukchi and Bering coasts of Siberia (but not the Arctic Ocean coast)
and southward to Japan and Korea. In the central parts of its range it is
abundant and from Northern California to the Bering Sea. It is perhaps the
most abundant of the flounders in near-shore areas. It is common in brackish
water and may ascend rivers for some distance, even into fresh water (Gunter,
1942; C. Hubbs, 1947). The starry flounder apparently does not venture far
from shore or into water of high salinity (Alverson and Wilimovsky, 1966;
personal observations in the Chukchi and Bering seas, August, 1974).
HABITS
Spawning occurs in the winter and spring, from November to February off
California, February to April in Puget Sound and British Columbia and still
later farther north, with the height of the spawning season apparently
corresponding with water temperatures near 11°C (Orcutt, 1950; Andriyashev,
1954; Hart, 1973). The eggs are quite small, 0.89-0.94 mm in diameter,
colorless to pale orange, with a thin, vermiculated membrane. They are
slightly less dense than sea water, so tend to rise and float near the
surface. A medium-sized female of 565 mm standard length (about 690 mm total
length) contained an estimated 11,000,000 eggs. Hatching occurs about 68
hours (2.8 days) after fertilization when the eggs are kept at 12.5°C, but
110 hours (4.6 days) at 10.5°C. The newly hatched larvae are about 2 mm
long, with a large yolk sac. For the first few hours, they are largely
quiescent and tend to float with the yolk sac uppermost. By 24 hours, how-
ever, the larvae, now about 3 mm long, have become quite active. The yolk
is completely absorbed in about 10 days. The time of metamorphosis is un-
known, but this phenomenon takes place at a length between 3.4 and 10.5 mm,
probably closer to the latter (Orcutt, 1950).
Scalation with minute cycloid scales is present at 25 mm and these
scales soon begin to develop into the typical stellate plates. Specimens of
100 mm show these plates on the eyed side of the head and along the dorsal
and anal fin bases on both sides of the body. At 200 mm, there are broad
bands of rough scales above and below the lateral line on the eyed side, on
the head and in double lines above and below the lateral line on the blind
side. As the fish continue to grow, the tubercles become more numerous and
cover more and more of the entire body (Orcutt, 1950).
188
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During the first 18 months of life there is a noticeable seasonal dif-
ference in growth rate; growth is rapid through spring and early summer, then
slows through the fall to accelerate again in mid-winter through the follow-
ing spring. The same growth pattern exists in older fish, but is less
obvious. Typical lengths (converted from standard to total length by multi-
plying by a factor of 1.22) for fish from southern California are 1+, 129 mm
cff, 133 mm ?; 2+, 287 d* , 294 $; 3+, 364 $ , 384 $; 5+, 477 (f , 519 $; 7+,
620 S. A fish of this last length will weight about 3,250 g (7.15 lb)
(Orcutt, 1950). Large individuals in more northern waters grow about 25 mm
per year (Manzer, 1952)
Sexual maturity is reached at two years for males, three years for
females. Females grow faster, reach a larger size and live longer than do
males.
The starry flounder makes inshore-offshore migrations with the seasons.
During the summer, they are inshore, often in very shallow water and in
estuaries; they tend to move into deeper water, up to 300 m, in the winter.
They may occur in the deep water at all times of year, for commercial fish-
ermen fishing for other species have taken them at these depths in May *
(Turner, 1886; Bean, 1887c; Orcutt, 1950; Andriyashev, 1954). Other than
these seasonal movements, starry flounders do not migrate much. However,
the young move up rivers, and the adults may also, for as much as 120 km.
Few adults have been taken far up rivers, however. A few fish tagged at the
mouth of the Columbia River and in British Columbia waters had moved 96-200
km, but most were recovered within 5 km of the tagging area (Manzer, 1952;
Westrheim, 1955).
The pelagic young feed on minute plankton. After metamorphosis and the
adoption of a bottom-dwelling life style, demersal copepods and amphipods
are important. As the fish grow, larger and larger items are included in the
diet. From a length of about 100 mm on, food includes clams, snails, star-
fish, amphipods, polychaete worms, crabs, mysids and nemerteans. Fish are
eaten only by the larger flounders, nostly those over 450 mm long. Small
starry flounders graze on the siphon tips of clams, which lie buried in the
bottom with only the ends of the siphons exposed. In Puget Sound, the
starry flounder feeds heavily from July to October, stops feeding in Janu-
ary and apparently does not eat again until June. It is not known whether
this feeding cycle occurs in other areas (Orcutt, 1950; Andriyashev, 1954;
Miller, 1967).
When the fish is at rest, the dorsal and anal fins are curled towards
the blind side, holding the body slightly off the bottom. Movement is most
often a sort of crawling, using the fin rays of the dorsal and anal after the
manner of the legs of a caterpillar. In quick, short movements, the pectoral
fins are used as paddles. When disturbed, the starry flounder flutters its
dorsal, anal and caudal fins so as to wave a covering of sand or mud over
the body. It may bury so deeply that even the eyes are covered, and may be
so confident of its concealment that force must be used to move it.
189
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Adaptive coloration is pronounced. Starry flounders can alter their
color and color pattern to simulate the bottom on which they lie. Fish on
mud bottoms are dark, those on sand tend to be brownish, while on extremely
light colored bottoms they are gray (Orcutt, 1950).
A major point of interest concerning the starry flounder lies in the
fact that, although it belongs to a group which normally has the color and
both eyes on the right side, it may have eyes and color either on the right
side (dextral} or the left side (sinistral). There is some evidence, albeit
incondusives which suggests that sinistral individuals are less viable than
dextral fish, at least in some areas (Hubbs and Hubbs, 1944). Marshalled
against this is the fact that over most of the range, sinistral starry
flounders out-number dextral ones. There appears to be a general trend, with
irregularities, for sinistrality to increase to the north and west. Off
California, sinistral individuals make up 55.2-59.5% of the population; off
the outer coast of Oregon, 49.21; at the mouth of the Columbia River, 60.4%;
off Washington, 66.4%; southeast Alaska, 58.2%; Kodiak Island and the Alaska
Peninsula, 68.01; Japan, nearly 100%. Sinistrality and dextrality are not
correlated with sex, both males and females showing the same degree of
reversal in a given population (Townsend, 1937; Hubbs and Kuronuma, 1942;
Orcutt, 1950; Andriyashev, 1954; Forrester, 1969a).
The starry flounder hybridizes with at least two other species, the
Japanese Karesius bjcoloratus and the North American P_arpjghrys_ vetulus. Both
hybrids were given scientific names before their hybrid status was realized.
The North American hybrid, Inopsetta 1schyra, is not uncommon and has been
studied to some degree. The hybrids have been found from Drake's Bay,
California, north to the Bering Sea. Reproductive level of the hybrid is low
due to abnormal development of eggs and sperm (Schultz and Smith, 1936;
Herald, 1941; Hubbs and Kuronuma, 1942; Aron, 1958; P. Reed, 1965; Ueno and
Abe, 1969).
IMPORTANCE TO MAN
Although the starry flounder is abundant and wide-spread, it is of rela-
tively small importance. It is taken commercially, but mostly as an inci-
dental catch when fishing for other species. Starry flounder is marketed,
along with several other flatfishes, as "sole", so it is difficult to deter-
mine how much is taken. In California, landings have been fairly steady
for many years at between 136,000 and 227,000 kg per year. Total landings
of unclassified flounders on the west coast in 1971 amounted to about
909,000 kg. Probably about 75% of this was starry flounder. Smaller fish
are considered superior to large ones for eating purposes, as the flesh of
the latter is said to be somewhat coarse. Opinions,vary as to the-quality
of the flavor. In Kamschatka it was canned as a luxury item for export
(Andriyashev, 1935; Ripley, 1949; Orcutt, 1950; Anonymous, 1974).
The starry flounder is reasonably popular as a sport fish, for it bites
readily most of the time and may often make the difference between a "dry"
day and a successful one.
190
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SECTION 17
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217
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219
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/
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239
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TECHNICAL REPORT DATA
(Please read Instructions on the ret ene before comptf' —'
1. REPORT NO,
EPA-600/3-79-053
4 TITLE AND SUBTITLE
FRESHWATER FISHES OF ALASKA
Their Biology, Distribution and Value
5 REPORT DATE-
May 1979 issue date
$ PERFORMING ORGANIZATION CODE
7 AUTHORSS)
Janes
8 PERFORMING ORGANIZATION REPORT NO
Morrow
9 PERFORMING ORGANIZATION NAME AND ADDRESS
10 PROGRAM ELEMENT NO.
University of Alaska
Fairbanks, Alaska 99701
BA 608
11. CONTRACT/GRANT NO
12 SPONSORING AGENCY NAME AND ADDRESS ,
Arctic Environmental Research Station - - -
Corvallis Environmental Research Laboratory
'. Office of Research and Development
\ U. S. Environmental Protection Agency
13 TYPE OF REPORT AND PERIOD COVERED
Final .June 1 , 1975-Dec. 31, 976
14 SPONSORING AGENCY CODE
EPA-600/02-
15 SUPPLEMENTARY NOTES
16 ABSTRACT- j.'
^-> '!
A summary of knowledge of the freshwater fishes of Alaska is provided.
Covered are 5& species in 3^ genera and 15 families, including strictly fresh-
water species, anadromous forms and those which normally are marine but which
occasionallv or regularly enter fresh water. For each species, a brief description
is given, followed by discussion of its ranqe and abundance, its aeneral bioloay
and its importance to man, as far as presently known.
This report was submitted in fulfillment of Grant No. R8038^5r01 by the
University of Alaska under the sponsorship of the Environmental Protection Agency.
s report covers the period 1 June 1975 to 31 December 1976 and was completed
as of 31 December 1976.
17
KEY WORDS AND DOCUMENT ANALYSIS
DESCRIPTORS
b IDENTIFIERS/OPEN ENDED TERMS c. COSATI Field/Croup
Fresh water fishes
Fishes
Alaska
18 DISTRIBUTION STATEMENT
Release Unlimited
19 SECURITY CLASS {This Report)
21 NO OF PAGES
20 SECURITY CLASS (This page)
22 PRICE
EPA Foim 2220—1 (Rev 4-77) PREVIOUS eoiT'CN is OBSOLETE
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