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homonym (Suckley 1861, Morton 1970), but there are four other scientific
names available for the bull trout, all proposed by Suckley (1858, 1861 ): Salmo
confluentus, Salmo bairdii, Salmo parkei, and Salmo campbelli. The first bi no-
men, Salvelinus confluentus (Suckley), is selected for use because, 1) it has
precedence over the other three in date of publication (Suckley 1858), 2) fewer
nomenclatural problems, including spelling, are anticipated for the name con-



146 CALIFORNIA FISH AND GAME

fluentus, 3) Suckley's type, a dried head and skin (USNM 1135), is still in
existence, and 4) S. bairdii and S. parkei lack type specimens. Jordan and
Evermann (1896) placed Sa Imo con fluentus Suckley in synonymy with Oncor-
hynchus tshawytscha (Walbaum), probably because Suckley's (1858, 1860)
descriptions read, in part, like that of a Pacific salmon. However, careful com-
parison of the head of the type specimen with Suckley's description has led me
to conclude they are the same.

TAXONOMIC HISTORY

As indicated above, the bull trout 5 was first described as Salmo spectabilis
(Girard 1856). Girard gave the locality of the holotype of spectabilis (USNM
7078) as St. Mary's Mission on the Clark Fork of the Columbia River, Montana,
but George Suckley (1860) who had collected the specimen in 1854, rede-
scribed spectabilis and corrected Girard's locality information, stating that the
holotype came from Ft. Dalles on the lower Columbia River. This specimen is
now a mutilated, half-rotted individual estimated at 200 mm sl.

In another paper, Suckley (1861) realized that the name spectabilis was
preoccupied and substituted Salmo campbelli (after Archibald Campbell, Chief
of the N.W. Boundary Commission). Morton (1970) has given a detailed ac-
count of this name change and the rules that pertain to it. In the same paper,
Suckley also described Salmo bairdii and Salmo parkei, which are both con-
specific with spectabilis. I searched the USNM collections but failed to find the
types of bairdii and parkei. However, the holotype of Suckley's "Salmo con-
fluentus" \n&s found (Figure 2A). This specimen consists of a dried head and
skin now preserved in alcohol. The head is that of a bull trout, but the description
by Suckley (1858) states that the dorsal, adipose, and caudal fins were spotted
profusely with dark brown and black (unlike Salvelinus) . This could not be
confirmed, for on examination of the type, no dark spots were found. Thus, I
believe it possible that the description of Salmo confluentus was based on the
remains or observations of two different individuals, one of which was a bull
trout and the other a Pacific salmon.

The type-locality for Salmo confluentus was given as the Puyallup River near
Ft. Steilacoom, Washington Territory. The type was procured September 27,
1856 by Suckley, apparently from Indians who had captured the fish — a large
(approximately 700 mm) male in spawning condition. It possesses a kype that
fits into a deep notch between the premaxillae. The notch is accentuated be-
cause of the dried condition of the specimen.

Thus, we see that Suckley not only collected the holotype of S. spectabilis and
later redescribed it as Salmo campbelli, but within a few years described the
same species three more times under different names {confluentus, bairdii,
parkei), using specimens he himself had collected.

Jordan (1879), in his key to the species of Salvelinus found within the United
States, included characters of the bull trout — "head large, stout, broad and
flattened above" — in his diagnosis of Salvelinus spectabilis (Girard). He appar-
ently based his key characters on specimens taken from the Clackamas River,
Oregon, by Livingston Stone. He also examined Girard's type of Salmo spectabi-



Although "char" may be a more appropriate term for members of the genus Salvelinus (Morton 1955), "trout"
is used herein as the common name for the bull trout in accordance with the American Fisheries Society's
attempt to stabilize fish nomenclature (Bailey 1970).



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT



147



lis, noting that the holotype was still preserved in the U.S. National Museum,
while that of parkei was lost. He added that parkei was unquestionably the same
as 5. spectabilis. In the same paper, Jordan recognized the bull trout from the
McCloud River as S. bairdii (Suckley), believing it differed from 5. spectabilis
by lacking basibranchial teeth. He did not list S. malma (Walbaum) in his key.
Jordan and Gilbert (1882) were responsible for placing the bull trout in
synonymy with Salvelinus malma (Walbaum). Jordan and Evermann (1896)
followed this precedent, which has been continued to the present. Jordan never





FIGURE 2. (A) Holotype of Salmo confluentus Suckley 1858, USNM 1135, head length 173 mm,
Puyallup R. near Steilacoom, Washington; (B) lateral and ventral view of head of
spawning male Salvelinus confluentus, NMC 66-70, estimated 550 mm standard length,
Deer Cr., British Columbia; (C) lateral and ventral view of head of spawning male
Salvelinus malma, UMMZ 126507, 295 mm, Karluk R., Kodiak I., Alaska. Photograph
by the author.



148 CALIFORNIA FISH AND GAME

correctly distinguished the bull trout from Dolly Varden where they occur to-
gether in Puget Sound. Under his use of the name, "spectabilis", he lumped the
bull trout with the Dolly Varden of coastal areas as far north as Alaska. In later
papers (Jordan 1923, Jordan and McGregor 1925) the name Salvelinus spectabi-
lis was used for a supposed "southern form" which ranged from northern
California northward to Alaska and the name Salvelinus malma applied to a
"northern form" known from Unalaska to Kamchatka.

The earlier descriptions of " Sal mo spectabilis" mentioned a number of distinc-
tive characteristics of the bull trout. Girard (1856, 1858) emphasized the curved
maxilla in the type specimen and its elongate head which entered the standard
length 3.5 times. Under his description of Salmo confluentus, Suckley (1858)
was the first to mention the "projection of the chin anterior to the front teeth";
in fact, he confused this character with the hooked lower jaw of spawning male
Oncorhynchus. He also added that the type of confluentus had 1 3 or 1 4 branchi-
ostegals. Again, in listing characters for Salmo bairdii, Suckley (1861 ) described
the "snout having a deep notch between the extremities of the premaxillaries
receiving a conical fleshy protuberance projecting upwards from the chin." In
the same paper under Salmo parkei, Suckley noted the large head "about four
and a half times in the total length; its top flat; muzzel pointed" and "branchi-
ostegals 13-14." Also in parkei, Suckley mentioned "a disposition toward the
formation of a fleshy 'tit' projecting upwards at the point of lower jaws with a
corresponding notch between the premaxillaries." Despite what Morton (1970)
has published to the contrary, there is little doubt in my mind that in his descrip-
tion of Salmo bairdii and Salmo parkei, Suckley (1861 ) was writing about the
bull trout and not the Dolly Varden.

To conclude this discussion of taxonomic history, it is emphasized that the
name confluentus is to be substituted for spectabilis in accordance with the rules
of zoological nomenclature (Article 53) and the holotype of this species
becomes USNM 1135.

DIAGNOSIS

A large species of North American Salvelinus reaching a greater size (to 18.3
kg, about 40 lb, Hart 1973) than S. malma; distinguished from the latter by its
long, broad head which is flat above and sharply tapered through the snout with
the eye positioned near the dorsal margin; head measures 3.7 in standard length,
averaging 3.6 in juveniles and 3.9 in large adults (5. malma measures about 4.3);
jaws and teeth well-developed, with cleft of mouth terminal; maxilla constantly
curved downward; branchiostegals typically 13 or 14 on the right side, 24-31
both sides combined; mandibular pores usually 7-9 on a side, 16 combined (5.
malma typically has 6 on a side); gill rakers robust with strong teeth projecting
from mesial edges toward the branchial cavity (whereas S. malma has relatively
long, finely tapered flexible gill rakers that are much compressed dorsoventrally
and lack teeth projecting from the mesial edge); gill rakers 14-19, pyloric caeca
21-36, and vertebrae 62-67; basibranchial teeth 0-11, averaging 4, and consist-
ently arranged in a single longitudinal row; in all specimens examined, there is
a pronounced gap on each side between the palatine and vomerine tooth rows;
in anterior view, there is a notch at the lower terminus of the snout (between
the premaxillae) that receives a fleshy protuberance on the symphysis of the
mandibles, although best developed in the adult (both male and female). This



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT 149

character is also seen in larger juveniles and is not to be confused with the kype
of spawning male salmonines, even though the same area is similarly modified
in large, breeding males of S. confluentus.

DESCRIPTION

The characters of S. confluentus described below were found to be consistent
in samples taken from localities throughout its known range. These samples
comprise a total of 332 individuals from eight major river basins draining to the
Pacific and Arctic oceans and Hudson Bay: Sacramento, Klamath, Columbia,
Skeena, Taku, Yukon, MacKenzie, and the Saskatchewan.

Some of the characters which have been employed for many years in Sal-
velinus taxonomy, such as numbers of gill rakers and pyloric caeca, will not
separate 5. ma/ma and 5. confluentus. This is one reason why the bull trout has
not been recognized as a distinct form. Most useful in separating S. confluentus
from S. malma are the shape and size of the head. In addition, characteristics
of the jaws, teeth, and gill rakers, number of mandibular pores, number of
branchiostegal rays, arrangement of basibranchial teeth, neurocranium profile,
and configuration of certain bones of the cranium, such as the supraethmoid,
frontal, preopercle, and opercle, are very useful. Numbers of gill rakers, pyloric
caeca, and vertebrae show considerable overlap between the two taxa.

Head Size and Shape

In dorsal view (Figure 3A) the head appears very broad and flat on top and
is hard to the touch. The frontals slope only slightly in a lateral direction away
from the midline. In S. malma (Figure 3B) the head is more compressed and the
frontals usually peak at the midline. Unlike S. confluentus, the frontals of 5.
malma are usually covered with thick, fatty tissue underlying the skin. This is best
observed in anadromous S. malma.

In lateral view (Figure 3A), the head of S. confluentus is low and sharply
conical (also see Paetzand Nelson 1970). The terminal cleft of the mouth evenly
divides the anterior profile of the head, while in S. malma the snout tends to be
shorter and deeper and often overhangs the tip of the lower jaw, especially in
juveniles (Figure 4B).

The eye of confluentus (Figures 2B, 3A) is more dorsal in position than in 5.
malma (Figures 2C, 3B). The vertical distance from the center of the eye to the
top of the head falls well short of the distance from the center of the eye to the
nares, while in S. malma, this distance reaches the nares or nearly so.

In anterior view, the greater breadth of the head is again noticeable in S.
confluentus (Figure 3). A major characteristic seen in this aspect is the notch
dividing the premaxillae, which receives a fleshy protuberance at the teminus
of the lower jaw ( Figure 3A) . This character is best developed in adults of both
sexes and reaches maximum development in spawning males (Figures 2A, 2B).
Juveniles older than 2 years may also have this feature.

S. malma may have a well-developed kype in spawning males of anadromous
populations, as shown by Morton (1965), but the kype is barely evident in
spawning males of the nonanadromous S. confluentus ( Morton 1965). On speci-
mens I have examined, the kype was best developed on the largest spawning
males (over 500 mm sl). Its existence is probably a function of size. The kype



150



CALIFORNIA FISH AND CAME




FIGURE 3. Head form (lateral, dorsal, ventral, and frontal views) in the adult female of (A)
Salvelinus confluentus, NMC 66-437, 580 mm, Finlay R. trib., Brit. Col.; (B) Salvelinus
malma, UMMZ 126507, 289 mm, Karluk R., Kodiak I., Alaska. Photograph by the
author.

in S. malma is directed dorsally, whereas in S. confluentus it has a slightly more
anterodorsal orientation.

The upper jaw of the bull trout in lateral view (Figure 3A) always exhibits a
pronounced downward curve as seen in the concavity of the toothed margin
and convexity of the dorsal margin, particularly where the supermaxilla is seated.
Typically, the maxilla of S. malma is more slender, with the toothed shaft straight
or only slightly curved downward.

Head length of 5. confluentus (Table 1 ) typically enters the standard length
less than 4.0 times, whereas in S. malma it enters the standard length about 4.25
times. The difference is greater when data from dwarf landlocked populations



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT



151




FIGURE 4. Head form in juveniles of (A) Salvelinus confluentus, OSUM 25212, 235 mm, female,
trib. of S. Fork Flathead R., Montana, (frontal, lateral, dorsal views); (B) Salvelinus
malma, OSUM 25213, 138 mm, female, trib. Karluk R., Kodiak I., Alaska, (frontal,
lateral, dorsal views); (C) Salvelinus confluentus UMMZ 188857, 134 mm, male, trib.
of Clearwater R., Montana (lateral view). Photograph by Cus Spreitzer.

of S. malma are excluded. However, the proportion of head length to standard
length changes with the size of the fish in both S. malma and S. confluentus
(Table 2). While individuals under 100 mm would be difficult to separate using
this character, those over 250 mm show a high degree of separation. I found no
difference in proportionate head size between adult male and female S. con-
fluentus, although I suspect that males over 500 mm will have relatively longer
heads. In S. malma, however, males over 200 mm tended to have larger heads
than females of the same size.

Branchiostegal Rays

S. confluentus had the highest branchiostegal ray count of all Salvelinus I

investigated. It averaged 14 rays on the left side and 13 on the right side in 120

specimens, with a range from 12 to 16 (Table 3). The range for both sides

combined was 24 to 31 with a mean of 27 (Table 4). Among the species of

2 — 77343



152



CALIFORNIA FISH AND GAME



TABLE 1. Head Size as the Proportion of Standard Length to Head Length in Salvelinus

confluentus and Salvelinus malma.

Drainage Sample size Range Mean

Salvelinus confluentus

McCloud River, California 12 3.2-3.9 3.6

Klamath River Basin, Oregon 5 3.5—4.1 3.8

Columbia River Basin, Snake River drainage 11 3.6-4.7' 3.8

Columbia River Basin (excluding Snake drainage)

Puget Sound, Fraser River Basin 23 3.3-4.1 3.7

Skeena River Basin 10 3.5-4.0 3.7

Taku River Basin 5 3.7-4.1 3.9

Yukon River Basin 7 3.6-4.0 3.7

MacKenzie River Basin 25 3.5-4.1 3.7

Saskatchewan River Basin 13 3.5-4.0 3.7

TOTAL 111 3.2-4.1(4.7)' 3.73

Salvelinus malma

McCloud River, California 2 4.5-4.7 4.6

Soleduck River, Washington 5 3.8-3.9 3.9

Puget Sound, Washington 4 4.2-4.6 4.4

Skeena River Basin 3 3.9^.5 4.1

Coastal British Columbia 14 3.5-4.5 4.2

Taku River Basin 7 3.5-4.6 4.1

Gulf of Alaska, Kodiak Island, Coastal Streams and Islands 20 3.6-4.7 4.3

TOTAL 55 3.5-4.7 4.23

1 One individual, Univ. Utah No. 1, with abnormally small head.

TABLE 2. Head Size as the Proportion of Standard Length to Head Length in Salvelinus
confluentus and Salvelinus malma, in Relation to Body Size and Sex.

Standard length (mm) Sample size Range Mean

Salvelinus confluentus

50-100 5 3.3-3.8 3.63

101-150 20 3.4-3.9 3.62

151-200 27 3.2—4.0 3.67

201-250 24 3.5-^.1 3.74

251-300 15 3.6-1.1 3.82

301-350 11 3.5-3.9 3.70

351-568 10 3.6-4.1 3.86

; (200-568) 20 3.5^1.0 3.73

- (200-423) 18 3.5-4.0 3.73

Salvelinus malma

50-100 10 3.3-4.1 3.63

101-150 10 3.8-4.3 4.00

151-200 10 3.6-1.5 4.16

201-250 9 3.9-4.7 4.36

251-300 14 4.0-4.7 4.38

301-445 8 3.8-4.7 4.40

J (200-380) 12 4.1-1.7 4.46

r (200-445) 10 3.8-4.4 4.10



Salvelinus, S. namaycush is closest to S. confluentus in number of branchioste-
gals. Vladykov (1954) reported S. namaycush To average 13 on the left side with
a mean of 25.3 for both sides combined. This character is important in separating
S. confluentus from S. malma. The average for 88 specimens of S. malma was
11.61 on the left side, 11.0 on the right side, with a range of 9 to 12 (Table 3).



TAXONOMY AND DISTRIBUTION OF THE BULL TROUT



153



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TAXONOMY AND DISTRIBUTION OF THE BULL TROUT 155

For both sides combined, S. malma had a range from 19 to 25 with a mean of
22.6. Counting branchiostegal rays on the right side separated 90% of the 5.
confluentus from S. malma. By combining both sides, the separation was not
increased significantly.

Mandibular Pores
Of 118 5. confluentus examined, 63% had 16 or more mandibular pores on
both sides with a range from 12 to 19 (Table 5). In 5. malma there are 10 to 15
total with a mean of 12.1. Elsewhere among the species of Salvelinus, only 5.
namaycush has a higher mandibular pore count (Morton and Miller 1954).

Basibranchial Teeth
Basibranchial teeth of S. confluentus axe characteristically arranged in a single
longitudinal row. They usually number from 3 to 5; however, of 40 individuals,
6 (15%) had no basibranchial teeth. In S. malma basibranchial teeth are usually
more numerous and arranged in one to three rows on the basibranchial plate.
Three of 35 (9%) S. malma that I examined had no teeth, and Morton and Miller
( 1 954 ) found that 4 of 20 ( 20% ) specimens they examined had no basibranchi-
al teeth.

Gill Raker Morphology

After examining large numbers of gill arches from S. confluentus and S. malma,
I conclude that the form of the raker and the characteristics of its dentition are
more important in separating S. confluentus from S. malma than actual gill raker
counts, which have been employed so extensively in salmonid taxonomy. Gill
raker morphology provided a high degree of separation (98%) of S. confluentus
from S. malma. Features of the raker that are most distinctive are the shape and
degree of dorsoventral compression, the relative size of the teeth, and their
presence or absence along the mesial edge of the rakers (the edge that faces
the branchial cavity).

The gill rakers of S. confluentus ate robust and oval in cross section. They have
strong teeth projecting well out from the mesial edge (Figure 5A) as well as
having smaller teeth on the dorsal and ventral surfaces. The raker is ornamented
with prominent ridges along its lateral margin. S. malma, in contrast, has rakers
that are strongly compressed dorsoventrally so that the broad surfaces of the
rakers are flat and weakly ridged (Figure 5B). Usually the rakers possess long
tapered tips that are quite delicate. Although there are usually small teeth on
these surfaces in S. malma, the mesial edge lacks the strong projecting teeth
entirely.

Body Form

The trunk of S. confluentus tends to be slender and rounded with only slight
lateral compression (Figure 6). In S. malma the trunk, like the head, is more
laterally compressed. A proportional difference that helps to separate the two
species is found by dividing the head length into the distance from the vent to
the base of the tail. The head length nearly always equals or exceeds this
distance in S. confluentus, whereas in S. malma it falls short.

Spotting Pattern

The size and distribution of spots are weak characters, although they have
been used for identifying North American Salvelinus, especially by early taxono-



156



CALIFORNIA FISH AND GAME



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Online LibraryCalifornia. Dept. of Fish and GameCalifornia fish and game (Volume 64, no. 3) → online text (page 2 of 10)