t.
CANADIAN TRANSLATION OF FISHERIES AND AQUATIC SCIENCE1
No. 4643
The growth and life span of Sthenotheuthis pteropus
in the east-central Atlantic
by
G.V.Zuyev
Ch. M. Nigmatullin
V.N. Nikol'sky
Original Title:
Rost i prodolzhitel'nost' zhizni krylorukogo dal'mara
Sthenoteuthis pteropus v vostochno-tsentral'noi
Atlantike
From:
Zool. Zh. 58: 1632-1641, 1979.
Translated by the Translation Bureau (NDE)
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
18
pages typescript
•
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TRANSLATED FROM - TRADUCTION DE
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English
Russian
AUTHOR - AUTEUR
G.V. Zuyev, Ch.M. Nigmatullin and V.N. Nikolisky
TITLE IN ENGLISH - TITRE ANGLAIS
The growth and life span of"Sthenoteuthis« pterOpus in the
east-central Atlantic
TITLE IN FOREIGN LANGUAGE (TRANSLITERATE FOREIGN CHARACTERS)
TITRE EN LANGUE ÉTRANGÉRE (TRANSCRIRE EN CARACTÉRES ROMAINS)
Rost i prodolzhitel'nost' zhizni krylorukogo dal'mara
Sthenoteuthis pteropus v vostochno-tsentral'noi Atlantike
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,
Zoologicheskiy zhurnal, 1979, v. 58, No. 11, pp. 1632-1641
UDC 594.582.5:591.134+139(261.5/7)
(1632)*
The growth and life span of Sthenoteuthîs Pteropus in
Lu
7.`e!
0
..;u
,
the eastcentral_Atlantic
by G.V. Zuyev, Ch.M. Nigmatullin
and V.N. Nikolysky
E
(LI
{D.
0-,, (7)
c.
2 0
T)
e
C.:
L.)
0
k-
P
Institute of South-Sea Biology of the Academy of Sciences of
the Ukrainian SSR (Sevastopol)
The linear growth and weight gain of Sthenoteuthis pteropus
are studied by analyzing the dynamics ofthe size structure of the
catches in the east-central Atlantic, and its life span is deterr.
mined on the basis of this. The relative daily increases in mass
diminish from 15 to 0.5% in the course of development, and Its life
span does not exceed 1-1.5 years.
Despite the significant number of papers that have been devoted to various aspects of the ecology of cephalopods, data on their
growth and age are still extremely scarce. This is due to the fact
that cephalopods, with the exception of a few- small littoral species,
adapt very poorly to life in captivity.
The number of cephalopod
species presently reared in laboratory conditions does not exceed
15-20 (review: Boletzky, 1974 ).
Observations of cephalopods in
natural environments are associated with serious difficulties due
of these invertehrate,qy
high motor activity and the lack of sufficiently effective
to the
fishing techniques and equipment.
Mass tagging,-
a method of di-
rect grohirate analysis which ensures tag returns, has been
.
-•_ •
* The numbers
are
the
pages
of the
in the right-hand margin
Russian text - translator
SEC 5-25 (Rev. 6/78)
2
carried out only for Todarodes pacificus, the object of a specialized industry.
No. morp171ological.structureb that would characterize growth rate,
such as the scales, otoliths or bones in fish, have been detected so
far in cephalopods.
Clarke (1965) attempted to determine the age
and growth rate of Moroteuthîs ingens by the mandibles of the bees
on which he detected clearly defined structural formations resembling
those on fish scales; he called these structures "growth rings".
He associates the zones in which these rings come closer together
with the periods of overall growth inhibition as a result of yari'a.r
bility in biotic and abiotic factors.
Since we still do not know-
how long the squid are exposed to these factors, it is impossible to
establish the duration of each cycle of mandible growth. As a, Me7
suit, the author arbitrarily gives ît as 6 to 12 months. The e..7tempts to study the growth and age of cuttlefish by their shell
(sepium) were also unsuccessful, as the rate at which new septa
are formed depends to a great extent on the environment of theeftï
mals (Choe Sang, 1963; Richard, 1 9- 67, 1969; Boletzky, 1934aY.
It
has been established that, when cuttlefish are reared in laboratory
conditions, the rate at which new chambers are formed diminishes
drastically when there is a shortage of food, or when the tempera
ture drops. The number of septa can serve as an age indicator only
when the cuttlefish inhabit an unchangeable environment.
(1633)
The growth rates of cephalopods in natural populations are
most commonly analyzed with the use of empirical data on the modal
size classes and their time bias -
-_- (Summers, 1968; 1971;
Holme, 1974; Tinbergen, Verwey, 1945; Fields, 1965; Murata, Ishii,/
1977; Ishii,1977); however, this method can be used successfully
3
only in the cases where the spawnirig of cephalopods is of a welldefined seasonal nature, and usually In temperate zones.
The growth, age and life span of oceanic squid - are not as wellresearched as in the other groups of cephalopods. They-, preSumably,
grow more quickly than the neritic species.
For eXample, the Month4
ly increases in the size of some Todarodes- patificus (iweth. a, mantle
length of 15-25 cm) have been known to reach 2-4 cm during the grow.r
ing period in summer.
faster growth rate.
T. saggittatus is distinguished Byan even
During the summer growing period in the vicinity
of Iceland ànd in Norwegian waters, the average monthly Increases
in size amount to 5.2-7.6 cm with a length. of 2 0.. 30 cm Wridriksson e
According to our own data, the young of this
1943;Wiborg,72).
species measuring 19-24 cm in length grow by 4.5 cm in a month dur 7
ing June-July in the vicinity of Cape Blanc. The life span ofNem-r
mastrephes bartrami in the East Pacific is one year, and the 'maxi
mum increases in size for individuals 25-30 cm in length amount to
3-4 cm in a month (Murata, Ishii, 1977; Ishii, 1977 ). .
The present paper looks at the growth of one of the prolific
Atlantic species, Sthenoteuthis pteropus, which inhabits the upper
pelagic zone of the open waters in the tropical region of the ocean
from the coast of Africa to America.
The main difficulty in determining the age and growths rate og
this species lies in the year-round nature of its spawningi with a
weakly-defined seasonal fluctuation in the greater part of the reproductive region of the range. As a result of this, the population
is constantly replenished by individuals of younger age groups, and
so it is extremely difficult to establish the modal size classes.
4
The northern peripheral population of large squid (the distant
neritic population of the Canary Current). from the East-Central
Atlantic (EC Atlantic) was used as the modal one for analyzing the
growth rate.
Its range occupies the southeastern part of the nor
thern subtropical anticyclone system which consists of the water
masses of the Canary and Northern Equatoriallcurrents and the adjacent regions of the northern tropical cyclone system with "divergence
in its centre (Zuyev, Nigmatullin, 19771.. Some individuals of this
population grow to a size of 50 cm.
for calculating
The chbice Of this population
the growth rate was prompted by the fact that
the area inhabited by it occupies the part of the species- range
with the highest degree of seasonal variability in climatic and
hydrological conditions.
The calculation of growth rate was based on an analysis of the
size composition of the catches and the dynamics
of this composim
tion during the different months in the productive areas oe thé
populational range with seasonal migrations taken into account.
Analysiâyofthé samples from productive areas gives us the most
objective picture of the size structure of the whole population.
The modal size groups were established on the basts of the èamples
câught with hooks and lines Cspoon ,jgs for catching squie, Wè
analyzed a total of 2364 specimens 12-50 cm In length, grouped
into size classes differing by 3 cm. The main disadvantage of the
data obtained with the help of hooks and lines is that not enough
individuals of the smallest and largest classeè•(smaller
than 18 cm and larger than 40 cm)were caught.
5
Throughout the year, the northern boundary of the populational
Cand speciesY range in the East-rCentral Atlantic undergoes notice-.
able spatial changes, shifting by 5-7 ° lat. off the coast of Africa.
(1634)
Squid of this species are abundant near the Madeira Islands in summ.
mer (32-34 0 N lat.), and are usually encountered south of the tropics
(23-25 ° N lat.) in winter (Zuyev et al., 19761,
Simultaneously with the seasonal changes in the boundary of
the range, we observe the spatial shifting of the productive zones
(the areas of concentration of squid) in theoceah. During the cold
winter-spring period (February-June),
accumulations of squid in
the East-Central Atlantic are observed mainly south and southeast
of the Green Cape Islands; squid are scarce north of these islands.
During the summer-autumn period (July-January), this productive zone
shifts northward, to the area between the Green Cape Islands and
Cape Blanc. Besides that, another productive zone in the form of
a continuous strip between the Madeira and Canary islands is formed
in summer. Groups of squid gather at the ,-Madeira Islands eachlrear
in August-September, and disappear from the area shortly àfterwards
(Clarke, 1966).
A comparative analysis of the size structure and biological
condition of individuals from different regions of the populational
range permits us to conclude that the population of large squid is
represented by at least two biological forms, apparently of .a
populational class, which alté .Edistenguished by the pereods and
places of spawning (the summer-spawning and autumn-,spawning fonmsE,
The summer-spawning squid reproduce mainly in July-September south
of the Green Cape Islands. The autumn-spawning squid reproduce in
6
October-December north of the Green Cape Islands, and forage in the
productive waters between the Madeira and Canary islands in summer,
Taking into consideration the above-mentioned intrapopulational
differences in the periods and places of spawning, we shall studythe annual dynamics of the size structure of the population of large
squid in order to determine their growth rate. In December, the
dimenal
y series of the squid in the productkve zone between the Green
---
Cape Islands and Cape Blanc is represented by- a bimodal curve with
the modes 18-21 cm (27.5% ) and 39,42 cm (2.5 96Y (fïg! 1, al,
The
first modal size group consists mainly of males and immature females
at the 1st and 2nd stages of maturity.
The second modal group
cludes only pre-spawning females (3rd, 4th and 5th_ stages of,Imatu,
rity).
It can be assumed that the small immature females-were-born
during the summer_iand-represent the summer-spawning form; the large
mature females are autumn-spawning ones which reproduce at this
particular time.
In January, the productive zone is located at the latitude of
Aimensionalj
the Green Cape Islands and south of them. T1TTY, series of squid
is characterized by the presence of a single modal size group,
21-24 cm (27.0%), in which immature females predominate (fig. 1, b).
Apparently, these are juvenile summer-spawning squid.
The decrease
in the relative abundance of large individuals and the disappearance
of the second modal group, which is observed in December, should Be
the fact that the overwhelming majority of the autumnatribuedo
generation has by this time spawned and died.
7
30
a
1
_
5
0\0
10
.
ow 3°
5
0
P
o
o
o
ttl
0
z
to
l
(
t .
1
1------.------
a
p1 10
.
.
e
10
I
1
1
,
r-,
i
r-1
I
i
JO
10
I
ofc
1
,
,
.
5
M
M
q
1 1
H1
24
JO
36
Leneb, og_mant-la f
Fig. 1.
112
48
•gt
Changes in the size structure of squid according to
the months:
a - December (n=282); b - January (n=293); c - February
(n=573); d - March (n=404); e - April (n=194); f, g August (h=356 and 262)
The curve of the size composition of the squid from this area
in February is similar in form to the January curve
(fig. 1, c),
its only difference beihg the ablUttagg. , , nu of the modal size to
the 24-27 cm interval, which is due to the further growth of the
8
summer generation of squid.
In March, squid 27-30 cm In length:predominate (20%f in the
productive zone south of the Green Cape Islands, i.e. the mode continues to shift toward the larger sizes (lifig. 1, d ) .
At the eame
time, we observe an increase in the relative abundance of squid with
a length approximating the maximum one. The squid of the modal size
group are represented exclusively by Immature femalee;
the largest
squid consist of mature females. It can be assumed that the latter
are also representatives of the summer-spawning (More abundant)
(1635)
subpopulation, which were born the year before last, and are a
year older than the squid of the modal size group. A large number
of 12-15 cm squid belonging to this species, which cannot be caught
with hooks and lines, is also observed in this area in March (near
the surface at night).
According to our assumptions, these are
representatives of the autumn generation, which were born in OctoberDecember.
The fact that these young squid belong to the autumn
generation is indirectly confirmed by their relativelyôld-loving
nature (the water temperature in the area inhabited by them does
not exceed 22-24 °C).
In April, the curve representing the size composition of the
catches is fairly flat with two weakly-defined modes of 21-24 cm
and 30-33 cm (fig 1, e).
There is no doubt that the individuals
of the second modal size group belong to the summer generation.
As to the individuals of the first group, it can be assumed that
these are squid of the autumn generation some of the males are
mature, the females are all immature).
(1636)
According to the data of
visual observations, 18-21 cm squid are the most abundant in April
9
(near the surface at night).
Apparently, the first size group is
flot completery represented in the catches due to the use of hooks
and lines.
As we have already mentioned, another productive zone,apart
from the year-round productive zone in the vicinity of the Green
Cape Islands (the area of northern tropical divergence)- , develops
in August between the Madeira and Canary islands, and some of the
large squid migrate to this zone.
Analysis of the size composI
tion of the catches and the biological condition of individuals
from different productive zones has revealed significant differences.
The curve representing the size cdmposition of the squid catches
south of the Green Cape Islands is bimodal, with abundance peaks
in the 24-27 and 39-42 cm intervals (fig. 1, f ) .
Of special in-r
terest is the second modal group (including_mature females) which
should be identified with the summer generation.
The first modal
group apparently consists mainly of growing squid oe the autumn
generation, which should reach maturity in December-January. The
,dimensional)
series of the squid catches in the vicinity of the Canary and
Madeira islands at this time is altogether different. It is limited
to a fairly narrow interval
(24-42 cm) with a marked predominance
of the 30-33 cm size group (40% of the total numbers) (fig. 1, gl.
All the individuals in this area are immature, actively feeding
females. They should be regarded as the autumn generation whic h.
will be reproducing between the Green Cape Islands and Cape Blanc.
The growth of the large squid of different subpopulations in
the East-Central Atlantic and their conjectural age can be depicted
10
in the following way. As we can see from the table and fig. 1,
the growth ratepf squid belonging to different-subpopulations
are similar, and so a single curve can Be used to depict their
growth. We chose to use the Bertalanffy equation which is used to
depict the growth of animals belonging to different systematic
groups, including the mollusks (Zaika, 1972):
Lt = 1,03 (1— e - k(1-1 0),
where L t denotes the linear size of an animaliof age t; and Loo,
k and to are constants.
The values of L,ce, k and to based on empirical growth data
are usually determined by the graphic method (Shcherbich, Slepokurov, 1976), or the method of computation (Hohendorf, 196- 61, When
using either of these methods, we must know the size of the animal
during equal time intervals. Such data are not available to us.
Growth and age of Sthenoteuthis.pteropus,
AUtuMri-S pawning
Summer spaWning
mantle
conjectural
mantle conjectural
age, months
age, months
length,
length,
cm
cm
-
Month
August
December
January
February
March
April
August
1
18-21
21-24
24-27
27-30
48-51
30-33
39-42
0
39-42
4
5
6
7
19
8
12
12
■■■
(12-15)
3
18-21
30-33
4
8
In this case, the unlmowir ,parameters,can be computed with the help
- -
of the numerical methods used for finding _ the extremum, which
minimize some functional of quality (Yefimov, Igoshin, 1976).
search . procedure can be simplified considerably by reducing
The
11
3-parameter computation to the numerical search for one parameter,
(1637)
L,o0, which enables us to use a low-capacity compùter and to greatly
reduce CpuA. time.
Having converted equation (1) to
4
(2)
Lo
and having taken the logarithms of both parts, we get
1n(1---)
-F kt
Loe
After inserting
37/ ln
Lco
;
a
(3L
(4)
— k; b = kt o ,
we get a straight-line equation relative to t l in its general form:
(5)
Let us assume that we have a seties of empirical values of
the mean dimensions of animal l i , which correspond to the series
of values of age ti where i=1, 2....n.
any 1.1>max{r1i
The straight line (5)_ for
can be plotted as a line of regression by the least
squares method, having determined the values a and b.
k and to are derived from (4).
The values
Lis determined by one of the me-
thods of onel-dimensional searchal, including the simple methodof
trying out the possible values of L„with'a prescribedstep.
•Sej-v-P-s--J
The minimumistandard deviation -07-as the criterion of maximum
conformity of the plotted curve to the empirical values
SD=
l -I t (
1/_
. i -21
1i - yi
(_6)
Yi
n—2
e values calculated
by equation (1). The coefficients k and t 0' and then the standard
deviation)
y SD, are computed , d for each assigned value of L. The Loo
---
value corresponding to the minimum SD is chosen.
*
central processing unit - transl.
•
12
50
tr+
e
w.
F-1
w zo
z
5000
te000
rn
rn
ni
2000
6 8 10 12 111 15 18 20
AgQr YI)Q1
Fig. 2. Growth curves of squid: a - linear growth (circlesssummer-spawning, triangles - autumn-spawning); b - weight
gain (1 - increase in mass, 2 - absolute daily gains,
3 - daily gains in % of body mass); c - growth of Ommastrephidae: I - Sthenoteuthis pteropus, II-- Todarodes
sagittatus tafter Wiborg, 1972; L - large Ommastrephes
bartrami, S - small ones (after Ishii, 1977)]
13
The numerical values of the equation parameters for S. pteropus, computed on the MIR-2 computer, are
t o --- 0.18 with SD=1.3%.
LOg
=60.0 cm, k=0.091 1 and
The middle values of the modal size
intervais from the table for summen,spawning squid were used as
the basic data. The computed growth curve is given in fig. 2 which
also contains data on the growth of autumn-spawning squid for the
purpose of comparison.
In order to pass from the linear growth of S. pteropus• to
weight gain, we useC.the empirical equation for squid 1 to 4.5 cm
in length, W=0.099 L 2 ' 13 , and the one for individuals exceeding the
length of 4.5 cm, W=0.019 L 3 r 2 , where L denotes the length-)6fthe
mantle (cm), and W the weigh±ofbthe-D:atei:d4g).
The curve representing weight gain is Sn-shaped. The utaxaum
increments in mass occur at the age of 11-14 months and amount to
more than 12 g/day.
The relative growth rate diminishes rapidly,
i;e. the increment in mass amounts to 5.8% of the body mass per day
at the age of one month, 1.3% at 6 months, and approximately 0.5%
at the age of one year.
(1638)
The table contains data on the growth of squid larger than
18-21 cm, and so the plotted curve cannot be expected to depict
the growth of the early young very well. Instead of attempting to
describe the growth of the young, we shall limit ourselves to analyzing the possibility of such rapid growth at the age of one month,
when the specific growth rate is at the highest level. According
to equation (1), young squid measuring approximately 1 cm (0.1 g)
grow to 6 cm (4.7 g) in a month. Such growth is possibleiïftthe
mass of the squid increases by 14-15% daily, though the law of
14
growth will differ from that of the given equation (1).
(1639)
a
100
80
eo
60
ai
O 40
ni
rd
ZO
18
20
2e
22
e, 100 F
-rd
•
80
. x'sx ■
1-
6
x
■
/
x., x
60 -
t.-..
/J
-
e' /
4
cri
W
44
/
'
,.. / /2
\k•, .\.\
/
•
-/
//
.,-1
0 40 4
M
/
JO T°
//
'-x--. x,„ 1
x,,.x
/, \
114
O
28
25
-
20
-
_
\
i /
>Lx-----x-- 5
. ._,,,
.......
//
i
:55 10
-
i
1
20
I
111
JO
40
1
I
50
Length_of mantle cm
,
-f
Fig. 3. Abundance relationship of large . (shaded) and small sguid
dépending on temperature (a) and infestation of the squid with
parasites of different sizes (b): 1 - Anisakis sp.,
2 - Phyllobothrium sp., 3 - Tentacularia coryphaena,
4 - Nybelinia sp., 5 - Didymozoidae
Considering that the daily food rations of the young amount
to 40-180 9 of their body mass when they are reared (LaRoe, 1971;
Boletzky, 1974) and the value of the coefficient of food consumption for growth (K 1 ) in cephalopods is equal to 0.4-0.6 (MangoldWirz, Boucher-Rodoni, 1973), such rapid growth is quite possible.
It should be noted that the counting off of age in the given
case begins when the squid is 1 cm in length.
The duration of de-
velopment from the moment the larva leaves the egg to the time it
15
reaches this size is unknown, but it apparently does not exceed
one month.
We found that the weakly-defined seasonal fluctuation of reproduction made it impossible to study the growth rate of squid
belonging to the small equatoriarpopulation, which begin to mature
at a length of 18-20 cm and grow to a maximum size of 35 cm, by
analyzing the dynamics of the Size structure.
Nevertheless, it can
be assumed that the small squid are similar to the large ones in
their growth rates, but are distinguished by a higher rate of deve ,
reproductive system at the higher temperatues in the lopmentfh
equatorial zone. In connection with this, we analyzed the temperature selectivity of the large and small squid.
Temperature selectivity was based on the frequency of occurrence correlation between small and large mature females at differ ,
(1640)
ent temperaturesof the surface layer of water.
Only large mature
females were observed at low temperatures (18-22 ° C).
As the tem-
perature rose above 22 ° C, we began to observe small mature females
which became predominant at temperatures of 26-30 °C (fig. 3). Therefore, small squid are more thermophilic than large ones.
At the
same rate of growth, the mass maturation of small squid begins4at
the age of 6 months, and the life span of a generation is less than
one year.
Additional data on the infestation of squid with the larval
forms of cestodes, nematodes and trematodes were used in the com,
parative study of the relative age and life span of small and large
squid.
Cestodes and nematodes accumulate in the organism oe squid
throughout life, while the numbers of Didymczoidae larvae-
16
gradually diminish (Gayevskaya, Nigmatullin, 1976). Because of
the intensity and extensiveness of invasion
this,
can serve as the, biological indicators of age in a comparative
study of squid from different size groups. However, the rations
of the squid must be the same when using the degree of infestation
as an indicator of age.
Due to the absence of food ration data,
the indexes of fullness were used. They proved to be similar in
squid of the same size in both the small and large forms.
The extensiveness and intensity of invasion of squid belonging
to the large formsproved to be considerably higher than in the small
forms (fig. 3, b), which indicates that large 'forms of squid have
a longer life span.
Our data on the growth rates of Sthenoteuthis pteropus accord
well with the data available on the growth of other species of Ommastrephidae (fig. 2, c).
Apparently, all of the oceanic species
of this family are characterized by high growth rates and a short
life span (1-2 years), which, together with monocycly, results in
a very rapid succession of generations.
References
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17
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Fisket
18
I
Ilapil311T0X035111HIlbIe CB11311 apbumpyxoro KaJlbB Tp01111YOCK011 AT.111111TIIKe. MaTeplIKJIbI II Bcec. crimrio60JIC311111,1 mopcarix XIIIBOTHI,IX: 16-17, KaamllimpaR.
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GROWTH AND LIFE SPAN OF• ST ENOTEUT HIS PT'ERUP US«
IN THE EAST-CENTRAL ATLANTIC
.
G. V. ZUEV, Ch. M. NIGMATULLIN
and V. N. NIKOLSKY
Institute of Biology of South Seas, Academy of Sciences of the Ukrainian SSR
(Sevastopol)
Summary
The linear and weight growth of StIzenoteuthis pteropus was studied and its life.
span was determined on the basis of studying the dynamics of size structure 'of the
catches in the East-Central Atlantic. The growth of squids is satisfactorily described bythe equation
e-0,091(t-F 0 .1 8)),
L t --- 60 (1
where Lt is mantle length in cm and t is age in months. The values of relative daily increments of mass are reduced in ontogenesis from 15 down to 0.5%, the life span cloes
not exceed 1-1.5 years.