ISSN 0704-3716
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CANADIAN TRANSLATION OF FISHERIES AND AQUATIC SCIENCES
No.
4667
Arctic char Salvelinus alpinus (Salmoniformes, Salmonidae)
bodies of water of Bering Island (Commander Islands)
of the
by K.A. Savvaitova, and V.A. Maksimov
Original Title: Golets Salvelinus alpinus (Salmoniformes, Salmonidae)
Vodoemov Ostrova Beringa (Komandorskie Ostrova)
From: Zool. Zh. 54: 1847 1859, 1975
-
Translated by the Translation Bureau (GAD)
Multilingual Services Division
Department of the Secretary of State of Canada
Department of Fisheries and Oceans
Northwest Atlantic Fisheries Center
St. John's, Nfld.
1980
29
pages typescript
ir
-s
DEPARTMENTOFTHESECRETARYOFSTATE
SECRÉTARIAT D'ÉTAT
TRANSLATION BUREAU
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MULTILINGUAL SERVICES
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TFA-5 g6 6 7
TRANSLATED FROM
INTO - EN
TRADUCTION DE
English
Rus sian
AUTHOR - AUTEUR
K.A. Savvaitova and V.A. Maksimov
TITLE IN ENGLISH - TITRE ANGLAIS
ARCTIC CHAR SALVELINUS ALPINUS (SALMONIFORMES, SALMONIDAE) OF THE BODIES OF WATER
OF BERING ISLAND (COMMANDER ISLANDS)
TITLE IN FOREIGN LANGUAGE (TRANSLITERATE FOREIGN CHARACTERS)
TITRE EN LANGUE ÉTRANGÉRE (TRANSCRIRE EN CARACTÉRES ROMAINS)
GOLETS SALVELINUS ALPINUS (SALMONIFORMES, SALMONIDAE) VODOEMOV OSTROVA BERINGA
(KOMANDORSKIE OSTROVA)
REFERENCE IN FOREIGN LANGUAGE (NAME OF BOOK OR PUBLICATION) IN FULL. TRANSLITERATE , FOREIGN CHARACTERS•
RÉFÉRENCE EN LANGUE ÉTRANGÉRE (NOM DU LIVRE OU PUBLICATION), ,AU COMPLET, TRANSCRIRE EN CARACTÉRES ROMAINS.
Zoologicheskii zhurnal
REFERENCE IN ENGLISH - RÉFÉRENCE EN ANGLAIS
Journal of Zoology
PUBLISHER - ÉDITEUR
YEAR
ANNÉE
PLACE OF PUBLICATION
LIEU DE PUBLICATION
USSR
1975
REQUESTING DEPARTMENT
MINISTÉRECLIENT
BRANCH OR DIVISION
•
VOLUME
54
DFO
NW At 1 . Fisheries Center,, St. John's
pp
ISSUE NO.
NUMÉRO
1847-1859
NUMBER OF TYPED PAGES
NOMBRE DE PAGES
DACTYLOGRAPHIÉES
12
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UNEDITED TRANSLATION
For information only
TRADUCTION NON REVISEE
infermation seuloment
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7630-21-029-8333
8 1980
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8 1980
Zoologieheskii zhurnal (Journal of Zoology), 1975, vol. 54, no. 12, pp
1847-1859, USSR.
UDC 597.553.2 (571.683):592/599:001.4-591.5
ARCTIC CHAR SALVELINUS ALPINUS (SALMONIFORMES,
SALMONIDAE) OF THE BODIES OF WATER OF BERING
ISLAND (COMMANDER ISLANDS)
IJNIEDITED TRANSLATION
For informl,Aion only
TRADUCTION NON REVISE
Information seulement
By K.A. Savvaitova and V.A. Maksimov
(Department of Ichthyology of Moscow University)
A description is given for the first time of arctic char of
the Commander Islands (Bering Island). The following were studied:
morphometric indices, qualitative characters (body shape, coloring,
nature of spots, etc.), age and growth, fecundity, and sex ratio. A
brief description is given of the habitats and the mode of life of
arctic char.
The shortage of spawning areas and the poor food supply
for fingerlings are responsible for the absence of non-anadromous lake
char on Bering Island and the presence of comPlexly structured populations of anadromous char.
The intrapopulation groups (typical anadro-
mous, river and dwarf males), obviously have a single gene pool and
are not genetically isolated from each other. Comparison of arctic
char with chars of other parts of the range shows its affiliation to
the species.a. alpinus. Some differences noted in the number of
vertebrae (Kamchatka population) and in the number of scales and
*The numbers in the right-hand margin indicate the corresponding
pages in the original. - Transl.
SEC 5-25T (Rev. 6/78)
/1847*
2
vertebrae (Sakhalin population) are not grounds for distinguishing the
Commander char as a separate subspecies.
The ichthyofauna of the bodies of water of the Commander
Islands is practically unstudied. Fragmentary information about the
fishes of this region is given only in the papers by Grebnitskii
(1897), Suvorov (1912) and Kurenkov (1970). At the same time,
information on the fishes which populate the Commander Islands is
important to solve a number of problems of taxonomy, zoogeography and
evolution, and also to organize the efficient exploitation and
conservation of fish resources.
This article is devoted to the most
numerous and ubiquitious fish species on the Commander Islands arctic char (Salvelinus alpinus). The aim of the authors is to refine
the taxonomic status of arctic char of the Commander Islands and to
study its population structure by analysing some mophobiological
features.
Material was collected from August 27th until September 4th,
1974 in the Sarannaya River on Bering Island. A fixed gill net with a
40 mm mesh and a fishing rod were used as a gear.
specimens were analysed.
In all, 100 fish
The mophometric indices were studied
according to Pravdin (1966), qualitative characters were described
(body shape, coloring, nature of spots etc.), and age (by otoliths),
fecundity and sex ratio were determined.
The Sarannaya River issues from Sarannoe Lake, which is
situated on the northeastern part of Bering Island.
It joins the lake
3
with the sea and is 1.4 km long. This body of water was of interest
for the following reasons: in the first place, we wanted to establish
whether there are resident freshwater populations of lake char on the
Commander Islands, and in the second place, the short length of the
river made it possible to study the structure of a char population
group maximally approaching the
At the time the material was
collected, the spawning run of arctic char from the sea into rivers
had finished, the groups of fishes reproducing in the few little rivers
flowing into the lake had gone for spawning, and of the small local
population lingering the Sarannaya River, most had already reproduced
or had prepared for spawning. We were not able to find freshwater
char in Sarannoe Lake.
Despite the considerable size of the lake
(length 7.7 km, width 5.1 km, maximal depth 31 m, surface area 31
km2 - Kurenkov, 1970), there apparently is no lake char in it,
probably because of the shortage of spawning grounds and the poor food
supply for fingerlings (Kureonkov, 1970).
In the lake itself, only
anadromous char is found going into rivers or migrating to the sea
after reproduction, and periodically foraging in the lake. Thus, the
object of study was a small local population of the anadromous form of
the char. However, for a local char population of a small river
belonging to one intraspecies form (Savvaitova, 1971), it proved to be
exceedingly varied. Firstly, this appeared in the varied mode of life
of the individuals comprising the population.
3
groups were
*In the original this sentence ends with a misprint ("demu") which
offers no clue to the intended meaning. - Transl.
/1848
11
distinguished in the population of the Sarannaya River: (1) properly
anadromous, relatively large chars of both sexes, coming from the sea
for spawning, (2) small river males and females, maturing initially in
fresh water and (3) small dwarf males, maturing in fresh water without
going out to sea.
Given below is a description of these groups.
Table 1
Polymorphism of coloring the population groups of anadromous char
(Salvelinus alpinus) of the Sarannaya River (% frequency of
occurrence).
THŒwnio. npox0Anan
•
Q, rt=--42
„BoHa Teza
cepe6plierz?ie
"2, cTanbaibie
3seaeHoHaTue..
24,0
_
.
,4xpaéuble
'-.1-1J1aBHHKII
cepta
-..1Hpacmie
opaH)HeHige.
:
Uxpyrnnie:
ThlTna
imHoro
(Ao.30);-:y:
de, n=23
. 67,0
0.11HBKOBble
• Bpioxo
16eme
lopaHHçeHoe
• 3HpacHoe
,:çf,:ry15b1
7:-. I cepue •
z opaHHceaue ; •
v_ynepHue
:.
•
poaoHble . •
pe‘masi
.
'22;0 ;.•„-
86,0
12,0
2,0
9,0
62,0
38,0
17,0
83,0
52,0
38,0
- 39,0
••• .56,0
5,0
0
•54,0
, 46, 0
<...! 52,0
26,0
r22,0 :
-= 56,0
44,0'
- 35,0
65 0
ri60,0
40,0
Key: 1 - character; 2 - group; 3 - typical anadromous; 4 - river;
5 - dwarf males.
a - side of body : 1 silvery; 2 - steel; 3 - greenish; 4 olive.
b - abdomen: 1 - white; 2 - orange; 3 - red;
c - lips: 1 - grey; 2 - orange; 3 - black;
d - spots: 1 - rose; 2 - red; 3 - orange;
e - fins: 1 - grey; 2 - red; 3 - orange; 4 - rose;
f - spots: 1 - large; 2 - small;
g - spots: 1 - many (30-50); 2 - few (up to 30).
Characteristics of the intraDoDulation groups of anadromous char of
/1849
the Sarannaya River
Coloring
Typical anadromous sexually mature char, coming from the sea,
have a light gray coloring and white or pale-rose spots on the sides
of the body. The number of spots below the lateral line varies
greatly, from 30 to 50, and they can be relatively large (2-3 mm) or
small.
The number of spots and their size are not correlated. With
length of time spent in fresh water, the chars darken, the sides take
on a steel, greenish or olive color and the spots become brighter.
At
spawning, a "nuptial dress" appears, mainly in males. The lips and
abdomen, fins and spots become red or orange.
Although these changes
represent the pattern, they are hardly observed in all individuals in
the population, and the degree of their manifestation varies (Table
1).
All coloring characters are polymorphous, that is there are
several phenotypes as to coloring within a single crossbreeding
population. The combination of different coloring characters varies
and they show no particular correlation with each other.
The overwhelming majority of river chars have olive sides with
small red spots.
Individuals with many (from 30 to 50) and few (up to
30) spots are found approximately in equal proportions.
Fish with red
6
fins predominate.
The lips are grey in 66% of the females, orange in
70% of the males. The abdomen is white in females, orange in 60% of
the males.
Sexual dimorphism is thus manifested to some extent in the
coloring of the lips and the abdomen (Table 1). River males and
females do not differ as to other characters.
Dwarf males of the anadromous char population have an olive
body.
There are small, mainly few, red or orange spots on the sides,
and the fins are also mainly red or orange. The lips are grey in
most.
The abdomen is white in half the males, orange in the rest
(Table 1).
Feeding
Anadromous char at the IV and V stages of maturity, that is,
at spawning, did not feed. Anadromous char at the II and VI stages of
maturity ate chironomid larvae, Trichoptera and mollusks. River char
fed on mollusks and aerial insects.
The absence of food in the
stomachs of some fishes is probably explained by a diurnal rhythm in
feeding and is not connected with reproduction.
Dwarf males fed on
chironomid larvae, Trichoptera and aerial insects (56%).
44% of stomachs were empty.
The other
The fingerlings of the different groups
of anadromous char fed on benthos and aerial insects.
Age and growth
The age of sexually mature anadromous females caught varied
from 4 to 8 years, that of males from 4 to 7 years.
and 6-year-olds predominated (Table 2).
In both sexes, 5-
7
The body length of anadromous char females is 27-54 cm, weight
180 - 1650 gms, length of males 26.5 - 41.5 cm, weight 120 - 680 gm.
The rate of growth of these chars, judging from observed lengths, is
relatively low (Table 3).
Besides anadromous char, reproducing in
1974, chars of similar size, also coming from the sea, but omitting
spawning, were also studied. Their age varied from 3 to 6 years.
In
size and rate of growth, they were the same as the rest of the stock
(Table 4).
The age of mature river char caught was 3-5 years, but 3- and
4-year olds predominated (Table 2).
The body length of females was
24-26 cm, weight 120-150 gm, length of males 16.5 - 25.5 cm, weight 60
- 140 gm.
/1850
The rate of growth of these chars was lower than in the
typical anadromous chars. Males apparently grow somewhat more slowly
than females (Tables 3 and 5).
The age of dwarf males varied from 2 to 5 years.
years olds predominated (Table 2).
3 - and 4 -
The body length of dwarf males was
12.5 - 21.0 cm, weight 10-70 gm. Weight growth is slow (Table 5).
Reproduction
Anadromous char is found year round in the bodies of water of
Bering Island.
According to Suvorov (1912) it spends the whole winter
in lakes and rivers, while in the spring, in May, it descends to the
sea and again ascends the rivers in June - August for spawning. It is
possible that there exists a spring run of anadromous char in May
(Berg, 1948).
According to our observations, the spawning run of
anadromous char into the Sarannaya River finishes in mid to late
/1851
8
August. For the bulk of population, spawning takes place in August
and early September. It takes place directly in the river, over its
entire length from the mouth to the sources, across the entire
In fact, the Sarannaya River presents a continuous spawning
channel.
grounds for char. It has a regular, pebbly-sandy bottom, its depth is
less than 1-1.5 m, usually 0.5 m; the rate of flow is 0.3 - 0.8 m/s;
and the water is transparent.
It is essentially the same as typical
salmon rivers.
Typical anadromous char reach maturity at the age of 4-5
years. Spawning does not take place every year, and some fish omit it
(Table 6).
The bulk of the population apparently spawns in a
relatively short period.
The others spawn later, at lower
temperatures.
Anadromous char build nests and bury their roe in the bottom.
Several males of different size spawn with a single female. Females
predominate considerably (65%) in the typical anadromous group. The
shortage of large anadromous males in this group is compensated by the
participation of dwarf and large river males in spawning.
The
fecundity of typical anadromous females, reckoned in gonads of the IV
stage of maturity, fluctuates from 537 to 4536 eggs and depends on
age, length and weight (Table 7).
The diameter of roe is 3.2 -
4.8 mm, average 3.8 mm, color pale-yellow.
The coefficient of
maturity of gonads is low, 4.6 - 13.4, average 8.2.
River char reproduce at the same spawning grounds as
anadromous char, and at the same time (Table 6).
Apparently, there
/1852
9
is no reproductive isolation between them, and in a number of cases,
spawning takes place jointly. They mature at the age of 3 years.
The
fecundity of river females varies from 249 to 484, average 353 eggs.
The roe is relatively large, its diameter is 4 mm on average, and its
The coefficient of maturity of gonads is higher
color is pale yellow.
than in typical anadromous individuals and varies from 10.7 to 15.6,
average 12.7.
Dwarf males reach maturity at 2-3 years of age.
They
reproduce simultaneously (Table 6) with large anadromous females.
Table 2
Age composition (in %) of catches of brood stock of different groups
of anadromous char in the Sarannaya River.
•
2 Bcapaer, roAL,
Fpynnupon;iu
—
%tame *
■
: ap.rumosbie
camuu
Key:
2
j
_
npoxoestibie
.
: non_
3
4
5
6
40,0
—
10,0
11,8
10,8
40,0
35,0
29,5
32,4
20,0
40,0
41,2
40,6
—
60,0
27,0
6,5
7
8
•
ce
d
y 9, ed
dd
—
—
—
6,5
1 - group; 2 - sex; 3 - age, years;
5 - river*; 6 - dwarf males.
4
5,0 10,0
17;5 —
10,8 5,4
—
—
—
—
20
17
37
15
15
- typical anadromous;
*Because of the small amount of material, information on both sexes
has been combined.
10
Table 3
Linear growth (based on observed lengths, cm) of brood stock of
different groups of anadromous char in the Sarannaya River
3
rpyruntpoBna
"
.1M MHO
a
npoxo,uHme
?-9
-
de
-
Ç
617
— 26-31 27-41 33-4.5
46-47
46,5
30
32
38
— 36-38 26-36 33-42 27-35
31
37
30
38
— 29—E8 26-41 33-45 27-35 46-47
31 3 38
47
3 .1
21 24-28 25-26
26
26
20-25 21-26
—
22
23
20
20-25 21-28 20-96
92
94
24
14-18 17-19
—
17
18
16
•
—
Penise
dd
—
Y-9,dd —
6 Kapormame
camut
5
314
2
.9,dd —
.
Bcpaer. roAm
floi
dd
12
Key: 1 - group; 2 - sex; 3 - age, years;
5 - river; 6 - dwarf males.
25
44
5
10
-
:15
15
4 - typical anadromous;
Table 4
Linear growth (based on observed lengths, cm) of adult anadrolous
char, omitting spawning in 1974.
2 Boapacr, roz,ht
non
3
L
1•
dô
V?-dd
. 20
7
26
25
20-26
23
Key: 1 - sex; 2 - age, years.
I
6
4
21-31
95
9 -1-27
96
20-31
30
28-33
30
39
31
28-32
30
39
11
Table 5
Weight growth of brood stock of different groups of anadromous char
in the Sarannaya River (in gins).
3
'
17p'yfirotpo3-
,
Hoa
...
1,
T1unitnio
npoxo-
.
5
4
3
2
Bo3pacr rc;Abl
6
8
7
1000-1150
370-750 470-900 —
—
14
1075
670
—
563
250
—
420 - 550 160-290 500-660 150-400
—
—
dd
• 12
• --1
225
• 553 .. - 2 7 - . 485 .
4
150
- :250-550
160--750
500-670
150-40010009 9 -,-.1 -d. —
,1075 - 26
648 - - . 2"./ - •
400- - -2150 • -—..
----- -- ..:. —
-—'
—
120-160 140-180
yy - 4
—
—
160
—
140
—
—
—
—
70-140
— 50-130
dd
—
9
—
—
105
90
9 2,
—
—
70-160
140-180
50-130
—
d
d
.
,
13
—
—
.160
—
117
90
9
• lime
`(
;
.
.
.1)e.glibie
5-
1::l11ntlio'Bble
camuEn
7_
9
—
—
. 6d
10 20-60
37
—
40
35-50
44
3 - age, years,
5 - river; 6 dwarf males.
Key: 1 - group; 2 - sex;
—
—
—
—
—
—
1C
4 - typical anadromous;
Table 6
Stages of maturity of gonads (in % frequency) in representatives
of different groups of the population of anadromous char in the
Sarannaya River
Cra,31111 3penocrn
Hon
rpyonapowln
II
I I I— II I
III
I III —I V
IV
IV— V
7,0
31,0
43,5
66,0
80,0
-81,0
-4,3
,3
10., 0
12,0
40,0
V VI —VII
.
28,5
TmIPiH rrpoxoAume
•
z.
.1:ap.-1111(oBble cam`Ki?!
ci
ced
c ce
21,6
12,0
8,8
2,5
17,0
7,0
Key: 1 - group; 2 - sex; 3 - stages of maturity;
anadromous; 5 - river; 6 - dwarf males.
19,0
17,5
17,0
42
23
6
10
46
4 - typical
12
Table 7
Fecundity of anadromous char as a function of age,
length and weight
2-> -A6comornan
TOCTb
11.T10g01311-
(nhcno iircincion)
rionanarenb
npegenht nal
ponaHlül
Bo3pacr, rop,b1
4
.e)
5
8
)1„mttia. Tau, c.tt
30-35
35-40
40-45
'
45-50
50-55
. Bec, e
•
250-500
500-750
750-1000
1000-1250
1500-1700
.
1
4
cpenx.
i
630-1400
815--1608
1320-1660
853
997
1490
400-660
• 7
450-936
12
720-1440
8
3 1320-1660
545
716
1092
1520
4536
400-815
12
10
450-1170
924-1608
6
2 1320-1660
592.
801
1273
1529
4536
2
'
Key: 1 - index; 2 - absolute fecundity (number of roe); 3 - limits
of variation; 4 - average; 5 - age, years; 6 - length,* cm;
7 - weight, gm.
*Presumably "standard length" (length of the body without the tail) Transi.
Morphometry
The typical anadromous char has the the following indices:
D
III-IV 9-11, average 9.9; A III-IV 7 - 9, average 8.5; P I 11-14,
average 12.6; # 120-155, average 134.6; right branchiostegal rays
(r.br) 10-13, average 11.3; left branchiostegal rays (r.br) 10-12,
average 11.0;
gill rakers (sp. br ) 20-24 m, average 21.8; pyloric
caeca (P c) 19-35, average 27.8; vertebrae (vert.)
63 - 68, average
64.5 There are no differences between the sexes as to meristic
characters (Table 8).
13
The head of anadromous char is relatively small, average 21.2% of
Schmidt's length, conical, triangular (5 )4 %) or rounded (44%).
Among
females, individuals with rounded head predominate (62%), among males,
triangular head (65%).
The upper jaw in the majority of fish reaches
the level of the posterior orbital margin or barely passes it (68%).
In the rest it goes far beyond the orbital margin.
It is straight and
narrow in 66% of fish. In almost all individuals studied, the jaws
are of equal length, only in 6% does the upper jaw overhang the lower.
The growth of connective tissue on the lower jaw and the notch on the
upper jaw are present in 34% of fish, absent in the rest.
However,
they are rare in females (17%) and comparatively frequent in males
(65%) and in this population their development is slight.
The body is
shallow as a rule in representatives of both sexes. The caudal
sediment is long (88%) and broad (89%).
The relative values of
plastic characters of the typical anadromous group are shown in Table
Sexual dimorphism is not evident in the body proportions.
8.
River char have the following diagnosis: D III-IV 9-11;
average 10.0; A III-IV 8-10, average 8.7; P I 12-13, average 12.5; ie
124 147, average 134.4; right branchiostegal rays 10-12, average 11.6;
-
left branchiostegal rays 10-12, average 11.7; gill rakers 20-23,
average 21.5; pyloric caeca 22-35, average 28.6; vertebrae 60-67,
average 64.6.
The head of river char is relatively small (22% of L 1 ),
smaller in females than in males, more often rounded in both sexes
14
(63%).
In most, the upper jaw terminates at the level of the
posterior orbital margin (63%) or barely passes it (31%).
In
/1853
approximately half of the specimens studied, it is straight (56%) and
narrow (69%).
In the rest, it is curved (44%) and broad (31%).
upper jaw overhangs the lower jaw in 63% of individuals.
The
The growth
of connective tissue on the lower jaw and the notch on the upper jaw
are absent in females and present in 40% of males. The body is of
average depth in the majority of fishes (81%).
long in 63% and broad in 81%.
The caudal peduncle is
Sexual dimorphism in body proportions
is not very pronounced (Table 9).
The relative size of the upper and
lower jaws and pectoral fin is somewhat greater in males. The body of
females is deeper.
/1 8 5 14
Dwarf males have the following indices: D III - IV 9-11,
average 9.9; A III-IV 8-10, average 8.5; P I 11 -14, average 12.7;
W
120-145, average 130.5; right branchiostegal rays 11-12, average 11,5;
left branchiostegal rays 10-12, average 10.6; gill rakers 20-23,
average 21.1; pyloric caeca 19-34, average 26.5; vertebrae 63-68
average 65.4.
The head of dwarf males is relatively small (average 22%),
rounded (69%) or conical (31%).
The upper jaw in the majority of fish
terminates at the level of the posterior orbital margin or barely
passes it (63%).
In the rest it does not reach the orbital margin.
The upper jaw is straight (75%) and broad (56%).
The upper jaw
overhangs the lower in the majority of cases (63%).
A growth of
connective tissue on the lower jaw and a notch on the upper jaw are
absent in 100% of individuals.
The body is of average depth (94%),
the caudal pecuncle is short (69%) and broad (87%).
The relative values of plastic characters are shown in Table
10.
/1855
Table 8
Morphometric characteristics of typical anadromous char in the
Sarannaya River.
n-42
311),!Veti
'
Wm
M=m
20-55
32,9+1,05
cr:S. n=23
lint
M=M
I
25--45
33,4+1.0
I
Cf
'
o
yltd0'4
h
H; :
h
;
pr.
AD'
tiV
•:,;;FiA
VA •
iD
h
I)
li
D
.1)
r. b I
r. b II
br
vert
•
6,8
no
.38-44
38-44
43-51
63-73
25--31
16--23
8-13
11-16
7-11
11-16
- 10-16
9-13
B % Or :1:1111ibl Te.la
72,8+0,24
1,6
21,0±0,18
1,1
4,3±0,09
0,6
3,4+0,00
0,4 ;
9,74,0,12
0,8
13.3±0,11
0,7
6,8=09
0,6
11,7±0,12
0,8
7,3=-0,12
0,8
1,4±-0,05
0,3
11,7±-0,15
1,0
18,5+0,23
1,8
7,0 ±0,08
0,5
0,3
19.0±0,13
1.1
41.0±0,18
1.9
41.0±0,18
47. 0 ±0,25
1;6
06,5.+0,14
1,8
27,7+0,20
1,3
10,7±0,24
1,6
1,2+04
0,9
13,5+0,14
0,9
9,0+0,12
0,7
1,2
13,3+0,13
13,4+0,16
1,0
10,9+0,14
0,9
120-155
9-11
7-9
11-14
10-13
10--12
20-24
19-34
63-68
Alepncr tilecKne npn3nalm
134.7±1,00
6,4
125-150
9,9±0,07
0,5
9-11
8,5+0,09
0,6
8--9
12,5+0,12
0,7
12--14
11,4±0,11
0,7
10--12
11,1+0,10
0,7
10--12
21,91E0,17
1,1
20--23
27,4+0,55
3,3
20--35
64,4+0,22
1,4
63-68
69.+70
- 18-24
3-6
2-5
8-12
11-15
5--9
9-14
6--10
0,5-3
• 0-14
'13-23
6-8
9,6
Cmnry
- 71,4=0,33
19-26
ILLS
8--12
12-16
5-9
10-15
7-10
1--3-
10-17
15--23
6-8
17-92
38-45
37--43
45-53
63-71
25-29
16--21
10-14
12--17
7--11
11--17
11-18
9-14
21,7+0.30
5,0+0,20
.3,4=0,09.
9,9+0.16
13,6+0.17
11,8=0.22
8,4+0,17
1,5+0,06
13,2=0,33
18,8+0,36
6,9=0.10
19,0+0,19
41,2+0.32
48:9-0. 9 5
06,3+0.35
27,1+0.18
18,8±0,23
11,6+0,18
14,4+0,20
8,8=0,15
13,7+0,25
14,6+0,31
.11,7+0,23
136,1±1,17
9,9±-0,09
8,6±0.10
12,8.2=0,15
11,2=0,12
11,0=0,1-1
21,7±0.13
28,8=0:78
• 64,5 =0,26 •
1,8
1,5
1,0
0,4
0,8
0,8 ,
0,6
1,1
0,8
0,3
1,6
1,7
0,5
0,9
1,6
1,4
1,2
1,8
0,9
1.1
0,9
1,0
0,7
1,2
1,5
1,1
5,6
0,4
0,5
0,7
0,6
0,6
0,9
3.2
1,2
Key: characters; 2 - in % of Schmidt's length; 3 - Meristic
characters. L1 is Schmidt's length, cm; 11 is the
torso length**;
- is the length of the head; r is the
**"Distance from the posterior border of the gill cover to the end of
the scale covering." - Transl.
length of the snout; o is the eye diameter; HCO is the depth
of the head at the eyes; HCZ is the depth of the head at the
occiput; io is the interorbital distance; op is the
postorbital distance; 1mac is the length of the maxillae; hmax
is the depth of the maxillae; 1md is the length of the lower
jaw; H is the greatest body depth; h is the least body depth;
is the length of the caudal peduncle; AD is the distance
from the end of the snout to the insertion of the dorsal fin;
PD is the postdorsal distance; AV is the distance from the end
of the snout to the insertion of the pelvic fin; AA is the
distance from the end of the snout to the insert of the anal
fin; n. is the distance from pectoral to pelvic fins; VA is
the distance from pelvic fins to anal fin; 1D is the length of
the base of the dorsal fin; hD is the depth of the dorsal fin;
1A is the length of the base of the anal fin; hA is the depth
of the anal fin; j_p is the length of the pectoral fin; 1v is
the length of the pelvic fin; fl is the number of scales in the
lateral line; D is the number of rays (soft) in the dorsal fin;
A is the number of rays (soft) in the anal fin; P is the number
of rays (soft) in the pectoral fin; r, b I is the number of
branchiostegal rays on the right; r, b II is the number of
branchiostegal rays on the left; sp. br in the number of gill
rakers; vert is the number of vertebrae; 10 is the number
of pyloric caeca. The same symbols apply in—Tables 9 and 10.
Comparative characteristics of intrapopulation groups of
anadromous char in the Sarannaya River.
These groups of char are characterized by a number of specific
features. They differ somewhat in appearance.
In addition to size
the typical anadromous char stands out either in its typical "marine"
(silver) coloring or bright breeding colors and corresponding changes
in body shape.
River chars and dwarf males have a characteristic
freshwater coloring, a predominately olive body, very bright spots and
fins, and many have an orange abdomen and lips.
Parr marks are
preserved on the sides of the body in small fish. This coloring is
typical of the river populations of the char when not spawning as
well.
Sexual dimporphism as to color and shape of the head is less
17
Table '9
Morphometric characteristics of river char of the Sarannaya River.
00, n.=.6
e1 ± m
•
L1
I
20-28
24,77E0,11
-
, Cjip
NCO'
HC3
lo
op
Imax
lumu
lmd
H
h
pl
AD
PD
AV
AA
PV
VA
1D
hD
IA
hA
lp
lv
3-5
3-5
9-12
12-15
6-8
11-13
7-8
1-2
11-14
18--23
6-8
15-20
37-44
37-43
44-51
61-70
25-31
11-21
lt-13
11-16
8-10
12-15
13-15
1t-13
li
D
A
r. b I
r. b II
sp. br
P
vert
124-147
9-11
• 8--10
12--13
10--12
10--12
20--22
22--33
62--65
Kev: 1 - character;
characters.
J
0,25
.
OT R.1111ibl
71,3±0 ,28
o
nt
Um
21,4=0,20
4,14-0,22
3,77E0,15
10,6=0,32
13,67E0,30
6.7770,11
11.777:J .1.21
1,5=0,12
12,0 E70,29
19,9-'0,78
7,2±0,08
18,07E0,62
41 ,3=9,90
40 , i =0,69
47,97E1,14
66,17E1,21
27,9-H0,87
19,51E0,75
11,7±0,21
14,7=06
9,0=0,13
13,5=0,22
14,37E0,06
12,1±0,32
I
20-25
22,47E0,6
0,2 .
.
Te.ia rio C.■ Eirry
.69-74
0,6
71,0=E0i99:
0.5 '
.20-25 •
22,37E0,39
3-6
4,77E0,15
0,5
• 3,97E0,13
0,4.
3-5
10,67E0,26
0,8
9--12
0,7
12--15
13,97E0,25
0.3
6-8
6,77E0,17
11,47E0,27
10-13 •
tj.5
7--11
8,17170,2•J
0,3
1--3
1,87E0,12
0,3
11--17
0,7
13,57E0,39
16-20
18,5±-0,43
1,9
6,97E0,17
0,20
6--8
.
18,27E0,31
1,5
15--20
2,2
42,27E0,48
40-45
• ,7
40,57E0,37
38--43
44-50
47,27E0,57
2,7
64,57E0,75
60--68
3.0
25-29
2,2
27,67E0,86
15-21 .
18,87E0,45
1,8
11,0-2-0,41
10--13
0,5
15,17E0,23
14--17
1,0
8,77E0,14
8--10
0,3
14,04-0,13
0,5
13--16
15,97E0,78
14--18
0,2
0,8
11--14
12,97E0,28
MeplicTwiecKlie npumaEll
134,57E3,53
8,6
133--143
9,87E0,31
0,8
9--11
8,8=0,31
0.8
S--10
12,37E0,25
12--13
0,6
11,6=0,31
11--12
0,8
11,37E0,31
10--12
0,8
20,6 4-0,31
0,8
20-23 .
27,8-H2,16
4,8
24-35
63,7=0,18
0,4
60--67
134,3+1,88
10,1±0,18
8,7+0,21
12,5+0,15.
11,6+0,15
10,9+0,18
22,0+0,30
29,11b1,09
65,2+0,66
2 - in % of Schmidt's length;
2,2 .
1,2
0,5
0,4
0,8
0,8
0,5
0 ,9'
0,9.
0,4 •
1,2.
1,4
0,5
1,0
1,5
1,2
• ,8
2,4
142
1,4
1,3
0,7
0,44,
0,4
0,8 0,9 ,
5,.9
0,6
0,7
0,5
0,5
0,6
0,9
3,3
2,11
3 - meristic
1,1
18
Table 10
Morphometric characteristics of dwarf males of the anadromous char in
the Sarannaya River (n = 16).
AI
1>c:tau.
• ,
.
•
,
.„
,
.•
9, or ;;:nui Tem no Curry
1,8
07-75 71,5±0,42
0,9
20-25 22,0±-0,21
0,5
4,4+0,12
3-s 6
0,4
4,4±0,10
3--6
.o
0,6
9-12 10,87E0,14
fe0
4,0
12-17 14,1±0,24
11 C3
0,7
6,3±-0,16
5-8
• io
0,8
10-14 11,9±-0,13
op.: .
0,5
8,2±-0,12
7-10
Inzax
0,3
1,8±0,08
1-3
hiiidt0,6
11--14 • 12,67E0,15
Irnd
1,0
19,1+0,26
17-21
H
0,5
7,07E0,13
6--8
h
1,2
16-21 48,1±0,28
1,3
40-45 42,07_1:0,30
AD
1,4
.36-43 40,1370,33
PD
1,4
46-51 48,74-0,34
À1./
-
M ± rn
Ilpit3tian
.2., In
PV
VA
ID
IA
!IA .
lp .
lu
II
D
A
r. b I
r. b II
sp. br
P
myt
63.4270
26-32
16-22
0-14
14-18
6-11
12-16
14-18
11-15
• 29,0-4-0,31 •
19,3+0,27
11,7:L-0, 9 2
15,5+0,9:4
8,6±0,21
14,17E-0,23
15,6-27.0,24
12,7±0,20
1,3
1,1
0,0
1,0
0,0
0,9
1,0
0,8
MeinicTntlecnne niniznalm
. 15
130 ,5
120-145
0,97±0.11
9-11
8,5-'41.14
8-10
12,7±0.19
11-14
11,5+0.12
11-12
10,6±0,16
10-12
21,1±-0,28
20-23
26,5±-1,09
19-34
65,4±-0,35
63-68
0,5
0,0
0,8
0,5
0,7.
1,2
4,4
1,4
1 - character; 2 - in % of Schmidt's length;
6,8
3 - meristic characters.
pronounced in the river group than in the typical anadromous group.
However, none of the char groups has characters wich are absent in the
others. Rather they differ only in the degree of their manifestation.
The relatively fast rate of growth of the typical anadromous
char and its great fecundity are directly related to the regular
migration to the sea and the better food supply.
River chars and
dwarf males, which for some reason have not migrated to the sea and
which have initially matured in fresh water, have a low rate of growth,
••
19
while river females have low fecundity.
The freshwater groups reach
sexual maturity earlier and their life span is evidently shorter.
Unlike the typical anadromous chars, they feed while spawning and are
infested with freshwater parasites.
Spawning of these groups takes place at the same time and in
the same places at the same spawning grounds. River chars and dwarf
males pawn together with typical anadromous chars, and freshwater
average-sized small dwarf males apparently play a very large role in /1856
the reproduction of the typical anadromous group.
There is no
reproductive isolation between the groups. This is also indicated by
the data on meristic characters: they show no differences. Only in
dwarf males is the average number of vertebrae somewhat greater.
However, this is easily explained when one considers the long spawning
period, the considerable fluctuations in temperature due to winds
(Kurenkov, 1970) and the small sampling volume.
There is no doubt that the intrapopulation groups of the
anadromous char of the Sarannaya River form a single spawning stock, a
single local population.
They have a single gene pool and are not
genetically isolated from each other. Such a population structure is
known for other salmonids, in particular sea trout (Salmon trutta)
(Berg, 1948), Black Sea salmon (Salmo trutta labrax) (Barach, 1952),
Kamchatka trout (S. mykiss) (Savvaitova et al., 1973), sockeye salmon
(Oncorhynchus nerka) (Krokhin, 1967) and others.
The initial
maturation in fresh water of river char males and females and dwarf
males does not mean that, subsequent to first spawning, they do not
migrate to the sea and do not return to the river as "typical
20
anadromous" fish, and thus the segregation of all the groups discussed
here is to some extent artificial. Their ratio in the population can
change over the years.
A complex structure of the stock is apparently typical of the
majority of the populations of anadromous char in other rivers of the
Commander Islands.
It is an adaptation to the specific conditions of
existence in short small rivers with limited area of spawning grounds
and food supply. It is only due to the extreme intrapopulation
diversity and the absence of strict isolation between groups
that
the char achieves such great abundance here and occupies all the
ecological niches available to it. In other areas of the range of the
species, in regions where there are bodies of water of diverse nature
and of sufficient size (for example, on Kamchatka), the intraspecies
structure of char is different. Here the species is represented by
intraspecific forms which are considerably more differentiated as to
mode of living and genetically(anadromous, lacustrine, lake-river and
stream forms) (Savvaitova, 1973) each of which is capable of
maintaining itself independently. There is no doubt that genetic
exchange between forms does take place, but it is more limited. There
are also dwarf forms in the populations of anadromous char on
Kamchatka, but their abundance and role in reproduction are
considerably smaller.
Taxonomie rank of the Commander char.
The intrapopulation groups of char described have analogues in
other regions.
The anadromous form of char is very widespread, and is
found all over the Holarctic region reaching far to the south along
21
the Asian and American coasts.
A number of authors assign this form
to one species, S. alpinus (Barsukov, 1960); Savvaitova, 1961a, 1973;
Volobuev, 1973; Gritsenko, 1975).
Some investigators, following Berg
(1948) hypothesize that an indigenous species of char, S. malma,
inhabits the Asian coast of the Pacific.
American authors (McPhail, 1961; Behnke, 1972; Vladykov, 1954;
Delacy and Morton, 19 )1 2) consider that there are two species in the
bodies of water of North America, S. alpinus and S. malma. However,
none of these investigators give what we would consider convincing
arguments for the existence of a species S. malma distint from S.
alpinus (Savvaitova, 1969).
We believe that the anadromous char of
the Commander Islands also belongs to S. alpinus. " It ià
similar to
anadromous char from other regions, in particular, the Kamchatka
population which is the closest geographically (Table 11).
Table 11
/1857
Meristic characters of anadromous char from different regions of the
Asian coast*
Panolim
flpn 3naK
• gY10•rK
•
r5
11-1-12
D
A
13-11
P•
11
r. br
sm br
yen!
a •
10--13
91_95
.
67-69
Baccenn
p. Oxor
(f
9-12
8-10.
11--14
125-147
9-13
18-25
18-34
65-67***
KamvaTma,
9-11
8--9
128-140
9--12
18-25
25--34
64-69
13-13
Bepullra „.
Caxammt
9-11
7-9
11--14
120--155
10-43
20--24
19-35
63-68
-115-136
10--13
13-25
18--35
58-6.5
Key: 1 - character; 2- regions; 3 - Chukchi Peninsula; 4 - Basin
of the Okhota River; 5 - Kamchatka; 6 - Bering Island;
7 - Sakhalin.
* The table is compiled on the basis of Barsukov's (1960), Volobuelfis
(1973), and Gritsenko's (1975) data and those of the authors of this
paper.
** Volobuev (1973) counted the vertebrae without the urostyle. To
bring his data into line with those of other authors, we enlarged his
figures by one.
22
Table 12
Meristic characters of the river (stream) form in different parts of
the range*.
pafio.b,
Pa flOii
ripumem
•
Kamgalsa
0-13
1/
?
-I
D
A
»
9-12
7-10
. 11-13
127-143
Bepintra
9-11
8-10
12-13
123-147
. ,.
'Caxaaux,
n p ,,,,,i,„
Ka NitiaTKEI
Caxamiu,
npilmopbe
((
4
;
9-11
7-9 • 11-14
115-137
0-B
B ep;ra
r. b
sp. br
PC
vert
10-13
16-24
21-35
02-70
Key: 1 - character; 2 - regions; 3 - Kamchatka;
Islands; 5 - Sakhalin, Primorle+.
10-12
20-23
22-35
60-67
510-13
14-25
20-32 .
58-64
4 - Bering
*The table has been compiled on the basis of the data of Savvaitova
and Romanov (1969), Taranets (1936), Gritsenko (1975) and the authors
of the present paper.
+Primor'e includes the Maritime Territory and the southern part of
Khabarosk Territory (USSR).- Transl.
Some differences have been noted in the number of vertebrae
(C.D.* 0.7) (Kamchatka population) and in the number of scales
(C. D. 0.8) and vertebrae (C. D. 0.9) (Sakhalin population).
*C.D.- "Coefficient of difference" (Mair, 1971).
.,
23
The small river form S. malma morpha curilus distinguished
previously (Taranets, 1936; Berg, 19 )1 8) in our opinion belongs to the
speciesâ. alpinus. It is very widely distributed from the Gulf of
Anadyr and Kamchatka to Korea. It is present in Alaska, the Aleutian
Islands, in the Kuriles, on Sakhalin Island, in the lower reaches of
the Amur River, in Primor'e and Japan.
This form typically has small
size, slow growth and bright coloring when not spawning.
In a number
of regions, the river form is apparently not isolated from anadromous
char.
Berg (19 )1 8) proposed that the stream form be considered a
morph of anadromous char.
In his opinion, it is biologically
equivalent to the trout. However, in some places, the degree of
isolation of small river char is relatively great, for example, in
stream char in the basin of Azabach'ee lake on Kamchatka (Savvaitova,
Romanov, 1969) and in the more southern regions of the range
(Taranets, 1936); Gritsenko, 1975).
The river chars of the Commander
Islands most likely correspond to this small river, stream morph.
They differ from the Kamchatka population in the number of vertebrae
(C. D 0.8) and from the southern population in the number of scales in
the lateral line (C. D -1.0) and vertebrae (C.D -1.2) (Table 12).
The dwarf males of the Commander Islands are similar to the
dwarf males described in the populations of anadromous char of
Kamchatka (Savvaitova, 1960).
meristic characters.
They have the same coloring and similar
24
Differences in intrapopulation groups of anadromous Commander /1858
char from the chars of other regions appear mainly in the average
number of vertebrae.
As is known, the southern char which lives at
the mouth of the Amur River, in Primor'e, in Peter the Great Bay, on
Sakhalin, in the Southern Kuriles and in Japan, and which differs from
the typical form in the lesser number of vertebrae and larger scales,
was distinguished by Taranets (1936) as the subspecies S. malma
krascheninnikovi Taranetz.
Later Gritsenko, (1975), assigned it to
L. alpinus. The characters on the basis of which this subspecies was
described are subject to clinal variability. We have discussed these
findings previously (Savvaitova, Movchan, 1973).
At that time,
following Mair (1971) we concluded that if geographic variability is
clinal in nature, it is usually best not to distinguish suspecies.
Commander char somewhat violate
the cline rule. They differ
from populations situated at the saine latitude and forming a
continuous series of clinal variability with respect to vertebrae
(Okhotsk, Kamchatka populations). The smaller average number of
vertebrae in Commander chars and the great variability of this
character are determined by the specific temperature conditions during
reproduction.
During spawning, the temperature is twice as great
(12.50) as during spawning, for example, of Kamchatka anadromous
char.
This specific feature is largely due to the location of the
Commander Islands and the nature of the rivers. It is relatively warm
here in August-September and the temperature in the rivers can change
suddenly and drastically, due to strong winds, which accounts for the
•
•:,,
25
great variability of vertebrae on the whole for the population (lim
60-70, = 1.72).
Commander char are probably isolated to a large
extent from the chars of other regions. Although their foraging
grounds at sea are not known, one can assume that they do not go far
from shore (Darda, 1964).
Thus there are [some] formal grounds for
distinguishing a new subspecies in this case. One can make the
The number of vertebrae is a very
following objections to this: 1.
variable character, which reacts strongly to changes in external
conditions.
It is possible that different generations of Commander
char will differ with respect to average number of vertebrae.
The
number of vertebrae in Commander char varies in the same range as in
Kamchatka char.
The dwarf males of the population of anadromous char
of the Sarannaya River do not differ in number of vertebrae from the
dwarf males of Kamchatka char.
2.
In no cases does the value of the
coefficient of difference reach the subspecies level of 1.28 (Mair,
1971).
3.
The characters on the basis of which the southern
subspecies has been distinguished have independent directions of
geographic variability.
For example, the number of scales in the
lateral line in Commander chars is the same as in Kamchatka char, that
is, it bits in with the general pattern of clinal variability of this
character from south to north.
The larger number of scales is
apparently explained by the fact that they are formed later than
vertebrae and at lower temperatures.
This once again confirms that
the number of elements of these characters represents an adaptive
reaction to local climatic factors.
26
Considering the above, the segregation of Commander char as a separate
subspecies would hardly be justified, and thus it is best only to note
the specific features of this char, without according it taxonomic
intraspecies status.
References
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The role of brook trout (Salmo trutta fario)
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•
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13. Savvaitova, K.A., Maksimov, V.A., Mina, M.V., Novikov, G.G.,
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(Salmonidae, Clupeiformes), Gidrobiol. zh., 9,4: 94-98.
15. Savvaitova, K.A., Romanov, N.S., 1969.
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(Kamchatka), Nauchn. dokl. Vyssh. shkoly, Biological
sciences, 8: 16-28.
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there:
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,,,•,..::
S ÀÉVELINUS ALPINUS (SALMONIFORMES, SALMONIDAE) FROM WATER
BODIES OF THE BERING ISLAND (COMMANDER ISLANDS)
..,
.
,
i\... A. SAVVAITOVA and V. A. AfAKSIMOV
•
•
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■
,
9epartinent of Iclztiology, State University of Moscow
Summary
The Arctic char of the Commander Islands (Bering Island) is described for .the first
ne, 'fhere were under study: morphometric indices, qualitative features (body form,
lour, spot pattern etc.), age and growth, fecundity, sex ratio. The habita':,, are
spa-::1-,.clerized and the \-,•ày of life is describe(i. The deficiency of areas suitab:e
ig
the poor food supply...of young fish determine the absence of non-anadromous
ctic char on the Bering Island and the presence of complex-structured polations of
adromous char. The intrapopulation groups (typical anadromou1, river and dwarf ma>) represemt, apparently, united gene pool and are not isolated genetically frorn each
ter. A comparison of the Commander char with the chars from different parts of the
1ge suggests its status as S. alpinus. Some differences noted in the number of vertebra
amehatka population) and of scales and vertebra (Sakhalin population) .do not allow
establish the Commander Islands char as a separate subspecies.
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