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CALIFORNIA
FISH™ GAME

"CONSERVATION OF WILDLIFE THROUGH EDUCATION"




California Fish and Game is a journal devoted to the conservation of wild-
life. If its contents are reproduced elsewhere, the authors and the California
Department of Fish and Game would appreciate being acknowledged.

Subscriptions may be obtained at the rate of $5 per year by placing an
order with the California Department of Fish and Game, 1416 Ninth Street,
Sacramento, California 95814. Money orders and checks should be made out
to California Department of Fish and Game. Inquiries regarding paid sub-
scriptions should be directed to the Editor.

Complimentary subscriptions are granted, on a limited basis, to libraries,
scientific and educational institutions, conservation agencies, and on exchange.
Complimentary subscriptions must be renewed annually by returning the post-
card enclosed with each October issue.



Please direct correspondence to:

Kenneth A. Hashagen, Jr., Editor
California Fish and Game
1416 Ninth Street
Sacramento, California 95814



u









I]



VOLUME 64



JULY 1978



NUMBER 3




Published Quarterly by

STATE OF CALIFORNIA

THE RESOURCES AGENCY

DEPARTMENT OF FISH AND GAME

—LDA—

CALH IRCES AGENCY LIBRARY

Resources Building. Room tl7

1418 9th Street



STATE OF CALIFORNIA
EDMUND G. BROWN JR., Governor



THE RESOURCES AGENCY
HUEY D JOHNSON, Secretary for Resources



FISH AND GAME COMMISSION

BERGER C. BENSON, President

San Mateo

SHERMAN CHICKERING, Vice President ABEL GALLETTI, Member

San Francisco Rancho Palos Verdes

RAYMOND DASMANN, Member ELIZABETH L. VENRICK, Member

Santa Cruz Cardiff-by-the-Sea



DEPARTMENT OF FISH AND GAME
E. C. FULLERTON, Director

1416 9th Street
Sacramento 95814



CALIFORNIA FISH AND GAME

Editorial Staff

KENNETH A. HASHAGEN, JR., Editor-in-Chief Sacramento

DARLENE A. OSBORNE, Editor tor Inland Fisheries Sacramento

RONALD M. JUREK, Editor for Wildlife Sacramento

J. R. RAYMOND ALLY, Editor for Marine Resources Long Beach

DAVID A. HOOPAUGH, Editor for Salmon and Steelhead Sacramento

DONALD E. STEVENS, Editor for Striped Bass, Sturgeon, and Shad Stockton



CONTENTS



137



Page



Taxonomy and Distribution of the Bull Trout, Salvelinus

confluentus (Suckley), from the American

Northwest Ted M. Cavender 139

Reproduction and Spawning of the Northern Anchovy,

Engraulis mordax, in San Pedro Bay, California Gary D. Brewer 175

Hooking Mortality of Juvenile Largemouth Bass, Micropterus

salmoides Ronald J. Pelzman 185

Catch-per-Unit-of-Effort Studies of Northern California

Dungeness Crabs, Cancer magister Daniel W. Gotshall 189

Sex Ratios of the Northern Anchovy, Engraulis mordax, Off
Southern California Richard A. Klingbeil 200

The Origins of Rainbow Trout, Salmo gairdneri Richardson, in

New Zealand. D. Scott, J. Hewitson, and J. C. Fraser 210

NOTES

The First Eastern Pacific Records of Bulleye, Cookeolus
boops, (Bloch and Schneider, 1801) (Pisces,
Priacanthidae) Ronald A. Fritzsche 219

A Hermaphroditic California Halibut, Paralichthys

califomicus Jack W. Schott 221

Spinal Column Deformity in a Pile Surfperch, Damalichthys

vacca Robert N. Tasto 223

A Note on Production Modeling of Populations with Discon-
tinuous Reproduction Alec MacCall 225

Observations of Agonistic Behavior in the Treefish, Sebastes

serriceps (Scorpaenidae) Peter L. Haaker 227

REVIEWS

ERRATUM

Smith, Gary E., 1978. An evaluation of disk-dangler tag shedding by striped bass
(Morone saxatilis) in the Sacramento-San Joaquin Estuary. California Fish and
Game, 64(2):93-97.
Page 94: The equation printed as: log [ A+B N, / ( A ^ B Nj + b-aH)]

should read: log [ A . B N ; / ( A . B N, + 8 _ A N,)] = log (1- A K) + i log (1- A S)

Page 95: The last sentence of Figure 1's caption (Numbers in parentheses
indicate number of fish from which tags were obtained.) should be deleted.



138



CHANGE OF EDITORSHIP

With this issue Kenneth A. Hashagen, Jr. of Inland Fisheries Branch
assumes the duties of Editor-in-Chief of California Fish and Game.

Mr. Hashagen's assumption of the editorship follows the Depart-
ment's policy of rotating the editorship between staff members repre-
senting Marine Resources, Inland Fisheries, and Wildlife Management.

For 7 years Mr. Hashagen, Senior Fishery Biologist, has served as
Inland Fisheries Editor of the Quarterly. Through this service he has
gained a knowledge of editorial policies and procedures of the Journal.

Under his guidance, the Journal will continue its policy of presenting
to the public the results of scientific investigations as they relate to
management programs and the conservation of California fish and wild-
life resources.

Mr. Hashagen will be ably assisted in his duties by five associate
editors: Darlene A. Osborne, Inland Fisheries; Ronald M. Jurek, Wildlife
Management; J. R. Raymond Ally, Marine Resources; Donald E. Stevens,
Anadromous Fisheries; and David A. Hoopaugh, Salmon and Steelhead.

To Mr. Collins, Editor-in-Chief the past 4 years, we wish to express
our appreciation for a job well done. £ C. Fullerton, Director, California
Department of Fish and Game.



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT 139

Calif. Fish and Came 64 (3 ) : 1 39-1 74. 1 978

TAXONOMY AND DISTRIBUTION OF THE

BULL TROUT, SALVELINUS CONFLUENTUS (SUCKLEY),

FROM THE AMERICAN NORTHWEST 1

by

Ted M. Cavender

Museum of Zoology

The Ohio State University

Morphological and distributional evidence is presented favoring the specific dis-
tinction of the bull trout, Salvelinus confluentus (Suckley ), a currently unrecognized
form of Salvelinus native to western North America. This species has been confused
with the Dolly Varden, Salvelinus malma (Walbaum). Separation of the two is based
primarily on characters of the head and cranial skeleton. Diagnosis and description
are given for the bull trout along with a history of its early taxonomy. Past and
present distribution of the bull trout ranges between lat 41° and 60° N. North of the
49th parallel it is found in most of the major drainages originating on both sides of
the Continental Divide. The bull trout is or at one time was sympatric with the Dolly
Varden in at least three major river systems, as well as the waters of Puget Sound.

INTRODUCTION

The salmonid genus Salvelinus has long been recognized as a difficult taxo-
nomic group. Without exaggeration, Vladykov (1954) referred to the taxonomy
of Salvelinus (along with other salmonid genera) as "extremely involved and
time-consuming." This is particularly true for species which are native to areas
bordering the North Pacific Ocean. McPhail (1961) referred to these popula-
tions of Salvelinus as part of the Salvelinus alpinus (Linnaeus) (arctic char)
complex because of their similarity, the unsatisfactory state of their taxonomy,
and the incompleteness of representative material.

The possible existence of more than one species of Salvelinus in the American
northwest has been a controversial subject since the days of the Pacific Railroad
surveys in the 1850's. Morton (1970) was the last to deal with the subject. He
concluded that only one species, Salvelinus malma (Walbaum) was recogniza-
ble, while none of the proposed subspecies was valid. However, in this paper
I will present morphometric, meristic, osteological, and distributional evidence
to show that there are two widely distributed forms of Salvelinus native to the
western United States and Canada; the Dolly Varden, S. malma, and the bull
trout, S. confluentus.

Although primarily an inland species, collection records show that the bull
trout is not strictly an interior, nonanadromous form, but combines coastal and
inland, as well as northern and southern aspects to its distribution. Because of
the taxonomic difficulties with this species, the bull trout has lacked uniform
scientific recognition even though it is a well-known sport fish. The scientific
literature, especially compendia of regional fish faunas, has for so long lumped
information about their ecology, morphology, and life history, that S. malma and
S. confluentus have been largely confused. If there are morphological characters
and aspects of their biology that consistently separate S. malma and S. confluen-
tus over their distributional ranges, why then has there been a taxonomic prob-

1 Accepted for publication September 1977.



140 CALIFORNIA FISH AND GAME

lem? The answer is due primarily to the lack of adequate comparative material
available to any one investigator.

The findings in this paper are part of a more extensive systematic treatment
of the genus Salvelinus I began at the University of Michigan in 1968 and have
continued at Ohio State University. Conclusions on the specific distinction of the
bull trout reached here are essentially those presented in a preliminary report
at the 1969 Annual Meeting of the American Society of Ichthyologists and
Herpetologists in New York City (Cavender 1969).

MATERIALS AND METHODS

Specimens of Salvelinus studied were principally those in the collections of
the University of Michigan Museum of Zoology (UMMZ); the National Museum
of Natural Sciences, Canada (NMC); and the United States National Museum
of Natural History (USNM). Locations given on the distribution map represent
collections housed in these institutions ( Figure 1 ) . Other material was examined
from the California Academy of Sciences (CAS).

Morphometric data were taken only from specimens from which standard
length could be recorded. The specimens of S. malma compared morphometri-
cally with S. confluentus were taken from the Pacific drainages of the United
States, including Alaska, and Canada. Most represent a type which McPhail
(1961) has termed the southern form of the Dolly Varden. The National Mu-
seum of Natural Sciences, Canada, possesses excellent material both of S. con-
fluentus and S. malma. A large part of this material came from the University of
British Columbia, Vancouver (UBC). A number of old and valuable specimens
of Salvelinus axe at the United States National Museum, including holotypes of
Salmo spectabilis and Salmo confluentus; Livingston Stone's collection from the
McCloud River, California, in the 1 870's; and specimens taken in the early 1 880's
from Puget Sound and coastal waters of British Columbia. In 1974 I collected bull
trout from the Flathead River drainage in Montana, which are now housed at
the Ohio State University Museum of Zoology (OSUM).

Osteological data were obtained from both dry-skeletal and cleared-and-
stained material housed at the Ohio State University and the University of
Michigan.

The number of gill rakers was determined by removing the first gill arch on
the right side and counting under a dissecting microscope the individual rakers,
including all rudimentary ones. Branchiostegal rays were counted after locating
the smallest, most anterior ray by dissection. Mandibular pores were counted
by exposing the openings of the mandibular sensory canal with a fine jet of
compressed air. The count includes all pores, but not the opening from which
the canal exists at the rear of the lower jaw. Pyloric caeca were cleaned of
connecting tissue and fat deposits prior to counting. Vertebrae were counted
from radiographs, which have been deposited at the University of Michigan
Museum of Zoology. Counts were repeated until the same number was obtained
twice.

To study gill raker morphology, the anterior right gill arch was removed,
stained in alizarin red-s, and cleared in glycerin. The most posterior gill raker on
the lower limb was then illustrated with the aid of a Wild-M5 Stereoscope with
integral camera lucida attachment.

Osteological abbreviations are as follows: AO (antorbital); ART (articular-



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT



141




• conflunntui

malma

* hybrid



FIGURE 1 . Distribution of Salvelinus confluentus and Salvelinus malma over same latitudinal range
in North America; plotted from localities of specimens examined.

angular); ATL (atlas vertebra); BOC (basioccipital); BR (branchiostegal ray);
BS (basiphenoid); DE (dentary); ECPT (ectopterygoid); ENPT (endop-
terygoid); EOC (exoccipital); EPO (epiotic); FR (frontal); HYO (hyomandibu-
lar); INT (intercalar); IO (infraorbital); IOP (interopercle); LE (lateral
ethmoid); MPT (metapterygoid); MX (maxilla); NA (nasal); ORS (orbito-
sphenoid); OP (opercle); P (parasphenoid); PA (parietal); PMX (premaxilla);
POP (preopercle); PRO (prootic); PTF (posttemporal fossa); PTO (pterotic);



142 CALIFORNIA FISH AND GAME

QU (quadrate); SE (supraethmoid); SO (supraorbital); SOC (supraoccipital);
SOP (subopercle); SPO (sphenotic); SPOP (suprapreopercle); SY (symplec-
tic); V (vomer).

Specimens Examined

Salvelinus confluentus (Suckley)

Sacramento R. Basin, McCloud R. drainage, Calif.: CAS 25691, (1), 110
mm, 2 Shasta Co.; CAS 38787, (1) 136 mm, near Nosoni Cr., Shasta Co.;
CAS 19889, (1 ), 147 mm, near Bollibokka Mt., Shasta Co.; CAS 38788, (1 ),
173 mm, Mt. Shasta State Hatchery near Mt. Shasta, Shasta Co.; USNM
26196, (1), skeleton, USNM 10547, (1), 175 mm; USNM 27820, (3), 211-
284 mm, USNM 15549, (2), 101-163 mm; USNM, (1), 339 mm; USNM
22452, (1), 301 mm.

Klamath R. Basin, Ore.: UMMZ 188849, (5— two cleared and stained),
140-168 mm, Long Cr., Lake Co.; UMMZ 188851, (1 skeleton), 169 mm,
Long Cr., Lake Co.; USNM 16793, (1 skin and skeleton), Linn Creek, Ft.
Klamath.

Columbia R. Basin, Snake R. Drainage: USNM 125309, (2), 142-154 mm,
Meadow Cr. near Sawtooth, Ida.; UMMZ 117879, (1), 216 mm, Stanley
Lake, a tributary to the Salmon R., Custer Co., Ida.; UMMZ 162298, (1), 166
mm, Dave Cr., Elko Co., Nev., a tributary to the Jarbidge R.; Univ. Utah No.
1, (1), 103 mm. Dave Cr., Elko Co., Nev.; CAS 38789, (6— one cleared and
stained 3 ), 90-155 mm, West Fork Jarbidge R., Elko Co., Nev.; UMMZ
127607, (1), 134 mm, Little Lost R. on Custer Co.-Butte Co. line, Ida.

Columbia R. Basin, mainstream and minor tributaries: USNM 7078, (1 ),
type of Salmo spectabilis, Columbia R. at The Dalles, Ore.; USNM 25273-
25276, (4), 279-313 mm, Walla Walla, Wash.; NMC 66-61, (1 ), Columbia
R. at Arrowhead, Brit. Col.; NMC 66-63, (22), Mars Cr., tributary to Co-
lumbia R. at Big Bend Highway, Brit. Col.; NMC 66-64, (5) Kinbasket Lake
at Tsar Cr., tributary to Columbia R., Brit. Col.; NMC 66-70, (3 — includes
one head), Lower Arrow Lake off Deer Cr., tributary to Columbia R., Brit.
Col.; NMC 59-169, (20), Luthead Lake, Banff National Park, Brit. Col.;
UMMZ 164591, (1 ), 252 mm, Emerald Lake, Yoho National Park, Brit. Col.

Columbia R. Basin, Clark Fork Drainage: USNM 38028, (1), 310 mm,
Clark Fork River; USNM 44002, (1 ), 158 mm, Rattlesnake Cr. at Missoula,
Mont.; NMC 59-148, (11), Shepp Cr., tributary to Flathead R., Brit. Col.;
NMC 66-72, (77), Pollock Cr., tributary to Flathead R., Brit. Col.; NMC
66-77, (17), Gumbo Flats Cr., tributary to Flathead R., Brit. Col.; UMMZ
161871, (3), 134-166 mm, Flathead Lake, Lake Co., Mont.; UMMZ 172458,
(7 — including three cranial skeletons), 212-423 mm, Flathead Lake, Lake
Co., Mont.; UMMZ 161866, (1 ), 350 mm, Flathead Lake, Lake Co., Mont.;
UMMZ 188857, (3— one cleared and stained), 130-145 mm, Morrell Cr.
near Seeley Lake, tributary to Clearwater R.; UMMZ 188856, (4), 47-90
mm, Big Creek, tributary to North Fork Flathead R. on western boundary
of Glacier National Park; OSUM 25212, (2— one cleared and stained),
208-235 mm, Hungry Horse Cr., tributary to Hungry Horse Reservoir on
South Fork Flathead R., Mont.; OSUM 25213-5, (1 skeleton), Flathead R.

2 All fish lengths are standard length.

3 The cleared and stained specimen from this lot has since been given a separate number: CAS 38790.



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT 143

at Coram below the confluence of North and Middle Forks, Flathead Co.,
Mont.; UMMZ 102948, (1 ), 257 mm, Pend Oreille Lake, Banner Co., Ida.

Columbia R. Basin, Kootenay Drainage: NMC 66-52, (2), Slocan Lake,
tributary to Kootenay R., Brit. Col.; NMC 66-57, (4) Duncan Lake near
Howser, tributary to Kootenay R., Brit. Col.; NMC 66-59, (1), Kootenay
Lake at the mouth of the Duncan R., Brit. Col.; NMC 66-68, (6), Kootenay
R. at Canal Flats, Brit. Col.; NMC 66-74, (1), mouth of Wolf Creek at
Skookumchuck, tributary to Kootenay R., Brit. Col.; NMC 66-78, (2), mouth
of Gold Cr. near Newgate, tributary to Kootenay R., Brit. Col.; NMC (no
number), (12), Lardeau R., tributary to Duncan R., Brit. Col.

Puget Sound, Washington: USNM 1135, (1), 733 mm, Pullayup R. at
Steilacoom, type of Salmo confluentus; USNM 27264, (2), 295-568 mm, D.
S. Jordan, 1880; USNM 42044, (1 ), 297 mm, Elliot Bay, O. B. Johnson, 21
May 1889.

Fraser R. Basin, Brit. Col.; NMC 55-130, (2), 184-185 mm, Salmon R. at
Hart Highway, north of Prince George.

Skeena R. Basin, Brit. Col.: UMMZ 159357, (3), 223-358 mm, Lakelse
Lake; UMMZ 159345, (2), 181-195 mm, Damshilgwit (Cabin) Lake;
U M MZ 1 59333, ( 1 ) , 230 mm, Morrison Lake; U M MZ 1 59337, ( 2 ) , 1 88-203
mm, Morice Lake; UMMZ 159352, (1 ), 368 mm, Slamgeesh Lake; UMMZ
159351, (1 ), 341 mm, Sustut Lake, USNM 86207, (1 ), 265 mm, Bear Lake.

Taku R. Basin: NMC 68-896, (5), 150-246 mm, Flannigan Slough, Taku
R. at International boundary between British Columbia and Alaska.

Upper Yukon R. Basin: NMC 68-1231, (7 of 15), 97-198 mm, Partridge
Cr., tributary to the Swift R. near Yukon-British Columbia boundary.

MacKenzie R. Basin, Liard Drainage, Brit. Col.; NMC 62-234, (1), 313
mm, Tatsho Cr. near Dense Lake, tributary to Liard R.; NMC 62-235, (2),
340-344 mm, Letain Lake near King Mt.; NMC 68-1230, (4), 129-176 mm,
outlet from Little Lake.

MacKenzie R. Basin, Peace R. Drainage: NMC 66-435, (1), 375 mm,
Chuchi Lake on Nation R., tributary to Parsnip R. N.W.T.; NMC 66-436, (1
of 2), 113 mm, Germansen Lake on Omineca R., Brit. Col.; NMC 66-437,
(1 ), 510 mm, Finlay R., about 4 miles upstream from Ft. Grahame, Brit. Col;
NMC 66-438, (7), 172-333 mm, mouth of Finlay R. to Manson R., Brit. Col.;
NMC 66-440, (1 ), 206 mm, Peace R., 25 miles downstream from Hudson-
Hope, Brit. Col.; NMC 68-802, (1) 171 mm, Peace R., 11 miles west of
Hudson-Hope, Brit. Col.

MacKenzie R. Basin, Athabaska Drainage, Alberta: UMMZ 80837, (1),
253 mm, Jacques Lake, tributary to Rocky R., Jasper National Park; UMMZ
159930, (5), 225-324 mm, Jacques Lake, Jasper National Park; NMC 59-48,
(A — including 2 heads), Jacques Lake, Jasper National Park.

Saskatchewan R. Basin: USNM 64326, (1 ), 715 mm, headwaters of Bra-
zeau R., tributary to North Saskatchewan R., Alberta; UMMZ 164943, (1 ),
164 mm, Banff National Park, Spray R., tributary to Bow R. of South Saskat-
chewan drainage, Alberta; UMMZ 164928, (1), 190 mm, Bow Lake, Banff
National Park, Alberta; UMMZ 164930, (1 ), 215 mm, Bow R., Alberta; NMC
60-343, (1 ), Red Deer R. drainage at Morrin, 65 miles northeast of Calgary,
Alberta; USNM 44444, (1 ), 231 mm, Oldman R., tributary to S. Saskatche-
wan R., Alberta; UMMZ 188900, (8), 190-267 mm, Cracker Lake, tributary



144 CALIFORNIA FISH AND GAME

to St. Mary's R. of S. Saskatchewan R. drainage, Glacier National Park,

Mont.

Salvelinus ma I ma (Walbaum)

Sacramento R. Basin, McCloud River, Calif.: USNM 20819, (2), 235-241
mm, sent by Livingston Stone, catalog entry made Nov. 24, 1877.

Soleduck R., Washington: UMMZ 93829, (14 — three cleared and
stained), 100-134 mm, above Soleduck Falls, Olympic Peninsula.

Puget Sound, Washington: USNM 34301-34305, (5), 252-274 mm, Port
Townsend, James G. Swan, coll. in 1884 or earlier.

Skagit R. Drainage, British Columbia: UMMZ 179422, (2), 50-65 mm,
Skagit R. near Hope.

Skeena R. Basin, British Columbia: UMMZ 159323, (1 ), 200 mm, Alastair
Lake; UMMZ 159344, (2), 140-144 mm, Johanson Lake.

British Columbia: NMC 59-150, (2 of 6), 126-130 mm, East Fork Seltat
Cr. at Haines Rd. and about 1 mile below Snowater Lake, tributary to Klehin
R.; NMC 65-213, (1 head), Nass Harbour, Iceberg Bay near mouth of Nass
R.; NMC 65-212, (3), 146-159 mm, Nass Harbour just north of Jacques
Point, Iceberg Bay; NMC 65-225, (5), 189-242 mm, mouth of stream, cove
on south shore steamer passage about % mile west of Khutzeymaten Inlet;
NMC 65-159, (2), 197-293 mm, cove on west side of Refuge Bay and at
north end of Porcher I., south of Prince Rupert; USNM 31979, (1 ), 285 mm,
Port Simpson, Capt. H. E. Nichols, June 1882; USNM 37610, (1 ), 98 mm,
taken in fresh water at Port Simpson.

Taku R. Basin, Alaska: NMC 58-402, (2), Twin Glacier Lake, tributary
Taku R.; NMC (UBC 58-388), (5— part of large series), Canyon I., Taku R.

Alaska, Aleutian Islands: UMMZ 106266, (13 — two cleared and stained),
52-127 mm, small stream on Atka I.; UMMZ 106529, (3), 383-497 mm,
Unalaska I.

Alaska: UMMZ 128983, (4), 257-310 mm, vicinity of King Cove, Belkof-
ski Bay, Alaska Peninsula; UMMZ 106260, (1 ), 288 mm, freshwater stream
tributary to Three Saints Bay, Kodiak I.; UMMZ 126507, (2), 286-289 mm,
KarlukR., Kodiak I.; UMMZ 126476, (11— two cleared and stained), 71-151
mm, Upper Thumb R., tributary to KarlukL., Kodiak I.; UMMZ 159395, (2),
100-125 mm, North Fork of Upper Thumb R. above falls, Kodiak I.; UMMZ
159393, (1 ), 53 mm, Falls Cr., above falls, tributary to Karluk L., Kodiak I.;
OSUM 25213, (5), 98-138 mm, mouth of tributary entering Karluk L.,
Kodiak I.; UMMZ 106267, (1 ), 174 mm, Lake at north end of Sitkalijak I.;
U M MZ 1 82299, ( 1 ) , 1 34 mm, Baranof I ., stream on north shore of Port Lucy
at west end of Island.

Copper River Drainage, Alaska: UMMZ 162600, (1 ), 173 mm, Chitina R.;
N MC ( BC 58-227 ) , ( 5 ) , 205-327 mm, South L. of Chenan Lakes to Copper
R.

Yukon River Basin: UMMZ 144581 4 , (1 ), 85 mm, Grant Cr. about 30
miles west of Tanana; UMMZ 133553 4 , (2), 186-194 mm, Riley Cr., McKin-
ley National Park; NMC (UBC 58-271), (4), 152-190 mm, Dry Cr. near
Alaska Highway, Tanana R. drainage.

4 Specimens that represent the "northern form" of Salvelinus ma/ma (McPhail 1961).



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT 145

MacKenzie R. Basin, British Columbia: USNM 147661, (1 ), 102 mm, Hot
Springs, 3 miles WNW of junction of Trout R. and Liard R. July 6-8, 1948,
J. R. Alcorn.

U.S.S.R.: UMMZ 145814, (1), 123 mm, Pogarna R., Kamchatka.

Japan and Kuril Islands: NMC 60-154, (6), Shokotsu R., Soya Province,
Hokkaido; UMMZ 186872, (6— one cleared and stained), 130-135 mm,
Pon-mataochi R., Nemuro Province, Hokkaido; UMMZ 188710, (9 — one
cleared and stained), 79-168 mm, Ishikari R. at Sounkei, Ishikari Province,
Hokkaido; UMMZ 188711, (9), 65-180 mm, Kuzira Bay, Paramusiro I.,
Northern Kuril Islands.

Korea: UMMZ 188698, (4), 70-187 mm, Upper Tuman R., Mozan;
UMMZ 188712, (1) 429, Joshin Bay near Seishin.

NOMENCLATURE

Salvelinus confluentus (Suckley)
Salmo spectabilis — Girard, 1856: 218 (orig. descr.)
Salmo spectabilis — Girard, 1858: 307-308 (amended descr.)
Salmo confluentus — Suckley, 1858: 8-9 (orig. descr.)
Salmo spectabilis — Suckley, 1860: 342-343 (correct locality given)
Salmo confluentus— Suckley, 1860: 334-335 (amended descr.)
Salmo bairdii— Suckley, 1861: 309 (orig. descr.)
Salmo parkei—Suck\ey, 1861: 309-310 (orig. descr.)
Salmo campbelli— Suckley, 1861: 313 (subst. name and amended descr.)
Salvelinus spectabilis — Jordan, 1879: 79-81 (amended diag.)
Salvelinus malma — (in part) Jordan and Gilbert, 1882: 319-320 (synonymy
given)

For years the bull trout and the Dolly Varden, here considered specifically
distinct, have been combined under one name, the Dolly Varden, Salvelinus
malma (Walbaum). The latter name is correctly applied to the form which is
generally characterized as anadromous. As now understood, the range of S.
malma spans the entire arc of the North Pacific from the Sea of Japan and Kuril
Islands, across the Aleutian chain to Alaska and south along the North American
Pacific coast to the northwestern U. S. (Figure 1 ). Because of its international
usage, the common name Dolly Varden is better reserved for S. malma although
it may originally have applied to S. confluentus. The latter ranges well inland in
the United States and Canada and is generally nonanadromous. Where S. con-
fluentus reaches an adult size of several kilograms or more, such as in the
Kootenay River of British Columbia and the Flathead River drainage of Montana,
fishermen refer to this species as the bull trout (Dymond 1932; Brown 1971 );
a name inspired by its large, broad head, its large mouth and prominent jaws,
and its highly piscivorous diet.

The original name for the bull trout, Salmo spectabilis Girard, is a secondary


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