Iranian Journal of Fisheries Sciences
15(4) 1410-1424
2016
Age and growth of bigeye kilka (Clupeonella grimmi
Kessler, 1877) in Iranian waters of the Caspian Sea
Janbaz A.A.1*; Fazli H.1; Pourgholam R.1; Kaymaram F.2; Khedmati K.3;
Parafkandeh F.2; Afraei Bandpei M.A.1
Received: January 2012
Accepted: April 2015
Abstract
Age and growth of Bigeye Kilka, Clupeonella grimmi, were studied in Iranian waters of
the Caspian Sea from early May 2006 to April 2007. The sagitta otoliths were removed
for each 5 mm length interval up to a total number of 262 fish. Fork length and weight
ranged from 92.5 to 142.5 mm and 4.2 g to 23.4 g, respectively. The largest lengthgroups was observed from December to February and the lowest in April, which was
coincided with spawning period and feeding during wintering, respectively. The overall
sex ratio (male:female) was 0.33:1 which differed significantly from the expected ratio
of 1:1. The length-weight regression was W= 0.0000744 FL3.14 for females and
W=0.0000341FL3.16 for males, indicating the Caspian Bigeye grew isometrically for
both sexes. Age determination based on otoliths readings showed that the population
was composed of six- groups from 2 to 7 years old which observed a rapidly growth
during the second year. In the age compositions, the four years old specimens with a
mean fork length and weight 115.5±7.8 mm and 11.9 ± 2.8 g were the most abundant
age group and accounted for 40.1%. The condition factor was 0.7-0.8 which varied in
difference months and coincided to the gonad development. Age-at-length data were
used to determine von Bertalanffy growth parameters for this population in both sexes,
indicating that K value and L∞ of females was more than that of males. The results
showed that C. grimmi is a rapidly-growing species in the southern parts of the Caspian
Sea.
Keywords: Age, Growth, Clupeonella grimmi, Caspian Sea, Iran
1-Ecological Research Center of the Caspian Sea, Iranian Fisheries Science Research
Institute, Agricultural Research Education and Extension Organization, P.O.Box, 961,
Sari, Iran
2-Iranian Fisheries Science Research Institute, Agricultural Research Education and
Extension Organization, P.O.Box: 13185-116, Tehran, Iran.
3-Inland Water Aquaculture Research Center, Iranian Fisheries Science Research Institute,
Agricultural Research Education and Extension Organization , P.O.Box: 66, Bandar
Anzali, Iran
*Corresponding author's email: aliasgharjanbaz@yahoo.com
1411 Janbaz et al., Age and growth of bigeye kilka (Clupeonella grimmi Kessler, 1877) in …
Introduction
Age and growth studies constitute an
extremely important field of commercial
fishery biology, providing the knowledge
which is required for the solution problems
of population dynamics, assessment of
fishing ground, and fishery management
techniques (Matisa, 1992; Mohd Nsir,
1994). The most abundant fishes of the
Caspian Sea are three small clupeids
known as “Kilka” which are anchovy
Kilka
(Clupeonella
engrauliformis
Svetovidov, 1941), common Kilka (C.
cultriventris caspia Bordin, 1904), and
bigeye Kilka (C. grimmi Kessler, 1877)
(Svetovidov, 1963).
Bigeye Kilka comprised about 1-7
percent of the total catch Kilka in the
Iranian waters (Besharat and Khatib, 1993;
Fazli et al., 2006). This species inhabits
the relatively deeper areas of the central
and southern Caspian Sea. Bigeye differs
from the two other species of Kilka in their
distinctive adaptation to deeper parts of the
water bodies (Prikhod’ko, 1981). For
instance, they have bigger eyes, structural
difference in their retinas and greater
transparency of the tissues (Prikhod’ko,
1981). The larvae of bigeye Kilka remain
further offshore in depths exceeding 70 m,
compared to the anchovy kilka. They are
stenohaline and live in the lower water
temperature (Prikhod’ko, 1981). This
species has a life span of 8 years in the
southern Caspian Sea with males and
females attaining sexual maturity between
65 mm and 75 mm, respectively
(Prikhod’ko, 1981). Bigeye Kilka, mainly,
have been recorded as native species in the
Iranian waters of the Caspian Sea and
rarely found in other regions of the Sea
(Prikhod’ko, 1981).
Some studies were carried out
previously on bigeye Kilka in Iranian
waters regarding to Kilka distribution (Svetovidov 1963; Besharat and Khatib,
1993; Rezvani et al., 1993; Naderi et al.,
1997), stock assessment methods by
hydroacoustic
during
1994-1996
(Pourgholam et al., 1996; Fazli and
Besharat, 1998) before entrance of
Ctenophorea to the Caspian Sea. Other
studies were on the age, growth and
biomass (Mamedov, 2006; Fazli et al.,
2009)
and
population
dynamics
(Parafkandeh, 2009).
Despite the economic and ecological
importance of C. grimmi in the Caspian
Sea, there is limited information available
regarding its age, growth and condition
factor in Iranian waters of the Caspian Sea
after observing the jellyfish (Ctenophora,
Mnemiopsis leidyi). Such information is
especially important because of the
ecological changes that are occurring in
the Sea at present due to the appearance of
Ctenophorea, in 1999 (Shiganova et al.,
2001). In the 1980s, this species was
reported in the Black Sea devastating
affected the whole ecosystem (Vinogradov
et al., 1989). The present study aimed to
investigate the biology, population
structure, age, growth and von Bertalanffy
population parameters of C.grimmi
compared with before observing the
Ctenophora M. leidyi in Iranian waters of
the Caspian Sea.
Iranian Journal of Fisheries Sciences 15(4) 2016
Materials and methods
This study was conducted in the Iranian
waters of the Caspian Sea between May
2006 and April 2007. The commercial
catches of Kilka fish is commenced at this
time of the year excepted from May to
June that catch of Kilka fish is banned. It
should be noted that this species is caught
only in the colder months (from December
2006 to April 2007). Fresh samples of C.
grimmi were collected using a conical
liftnets mobilized with 1500w underwater
electric lights to attract the fish in
Babolsar, Amirabad and Anzali stations in
Mazandaran and Guilan provinces,
respectively (Fig.1). The diameter of the
1412
hoop of a conical liftnet was 2.5 to 3 m
and the length of the net bag was at least
1.25 times greater than the diameter of the
hoop (Ben- Yami, 1976). The mesh size
between two knots of the net is 7-8 mm.
Age was determined using otolith reading.
For each sex we collect 5-10 otoliths from
each 5 mm size class. In this study, 151
otoliths were removed. To enhance
contrast and facilitate reading and
interpretation of growth marks, the whole
otoliths were put in the glycerin solution
for 24 hours and then observed with black
background under reflected light through a
microscope (Fig.2).
Figure 1: The landing places of kilka in Iranian waters (Amirabad-Babolsar and
Anzali harbors) of the Caspian Sea.
1413 Janbaz et al., Age and growth of bigeye kilka (Clupeonella grimmi Kessler, 1877) in …
Figure 2: Age determination of Clupeonella grimmi with otolith (annual rings in 4 years old)
After transferred specimens to the
laboratory, up to 200 specimens for Bigeye
Kilka were randomly selected and then
biological parameter as fork length, weight
and age were studied. Fork length and
weight of captured fish were measured to
mm and g, respectively.
The relationship between weight and
length was calculated using the
exponential regression:
W a Lb
where W is the total weight (g), L is the
fork length (mm); a is the regression
constant (intercept) and b is the regression
coefficient (slope) (Ricker, 1975).
The Fulton condition factor (CF) was
determined for each fish using the
following equation (Bagenal, 1978):
W
100
CF
b
L
where W is the total weight (g) and L is
fork length (mm). Length-at-age data were
fitted to the von Bertalanffy growth model
(Ricker, 1975) using non-linear leastsquares regression:
Lt L (1 e k (t t0 ) )
where Lt is the length of the fish at age t,
L∞ is the asymptotic length, K is a growth
rate per unit of time, and t0 is the
theoretical age at which the fish would
have length zero. Overall growth
performance index (Φ') of species can be
interpreted by the growth index (Munro
and Pauly, 1983):
K
L
log10
2 log10
Length-at-age data analysis was used to
FiSAT software proposes. Chi-squared
analysis was used to examine a significant
difference from an expected 1:1 sex ratio
for all fish.
Results
Length and weight relationship
Iranian Journal of Fisheries Sciences 15(4) 2016
Six hundred and twenty six specimens of
C. grimmi were totally collected and used
for this study. One hundred and fifty five
specimens were males and 471 were
females. The overall sex ratio (males:
females) in this study was 0.33:1 for adult
bigeye Kilka which differed significantly
from the expected 1:1 ratio (λ2 = 159.5,
p<0.05, Table 1). Females were generally
more abundant and significant difference
1414
was observed between males and females
throughout the year (p<0.05; Fig. 3),
except in January, there was no significant
difference between males and females (χ2
=2.8, p=0.096). Males were most abundant
in all the size classes lower than 102.5 mm
(p<0.001). Females predominated in the
size classes greater than 102.5 mm
(p<0.001; Fig. 4).
Table 1: Chi-square monthly test for Clupeonella grimmi sex ratio in the Iranian waters of the Caspian
Sea.
Month
Males
Females
Total
χ2
p
Dec.
14
113
127
77.2
0.001
Jan.
56
75
131
2.8
0.096
Feb.
64
189
253
61.8
0.001
Mar.
8
37
45
18.7
0.001
Apr.
13
57
70
27.7
0.001
Total
155
471
626
159.5
0.001
Fork length and weight ranged from 92.5
to 142.5 mm and 4.2 to 23.4 g (average
119.8±7.9 mm, 13.1±2.8 g, n=471) for
females and from 92.5 to 132.5 mm and
5.2 to 18.5 g (average 113.90±8.8 mm,
11.2±2.8 g, n=155) for males. A
significant difference was observed in
length and weight relationship between
males and females in various months
(ANOVA, F = 89.676, p<0.05). For both
sexes of all individuals, the relationship
between total length and somatic weight
was described as: W = 0.00000572 FL3 (r 2
= 0.91, n = 626);
1415 Janbaz et al., Age and growth of bigeye kilka (Clupeonella grimmi Kessler, 1877) in …
Figure 3: Monthly variations in sex ratio of Clupeonella grimmi in the Iranian waters of the Caspian Sea.
Figure 4: Sex composition of bigeye kilka Clupeonella grimmi in the Iranian waters of the Caspian Sea.
for females: W= 0.000007441 FL3.159
(r2=0.89, n=471); and for males:
W=0.0000034 FL3 (r2=94, n=155). There
were significant difference between sexes
in the slopes (b) of length–weight
relationship (t-test, t=38.736.102, p <0.05).
Length frequency
Fork length frequency of C. grimmi
samples varied between 92.5 and 142.5
mm. The largest length groups (from 92.5
to 142.5 m) was observed during
December to February and the smallest in
April (from 97.5 to 127.5 mm) which was
coincided with spawning period and
feeding during wintering, respectively
(Fig. 5). After a relatively low fork length
from the average 120.4 mm in December
to 117.7 mm in January, an increase in
fork length was recorded (up to the
average of 119.6 mm) in March. In April,
Iranian Journal of Fisheries Sciences 15(4) 2016
the fork length started to decrease steadily
to the average of 114.4 mm.
Condition factor
The Fulton's condition factor (CF) varied
from March (0.70) to December (0.79).
There was a decrease in growth rate from
December to March (0.79 to 0.7), an
increase in April (0.78), which was
coincided with development of gonads
(Fig. 6). Condition factor was 0.75 (Fig.
7).
Von Bertalanffy age-at-length catch curve
Age of 262 fish was determined using the
otoliths. The fish of the youngest group
was two years and the oldest was seven
years old. Totally, there were six agegroups (2 to 7 years old) (Fig. 8). The
mean age was 4±1 yr. Totally, there were
six age-groups (six age-groups for females
1416
with mean 4.1±1yr and also five agegroups for males with mean 3.9±1.2 yr). In
the age compositions, the four years old
specimens with a mean fork length and
weight 115.5±7.8 mm and 11.9±2.8 g were
the most abundant age group and
accounted for 40.1% (40% for males and
41% for females; Table 2). There was a
significant difference in age between
males and females (t-test, t=9.566,
p<0.05). The highest fork length growth
increments were 133 mm/yr for males and
144 mm/yr for females at four years old.
The von Bertalanffy growth parameters
were: L∞ = 148 mm, K = 0.29 year-1, t0 =1.123 years for both sexes (Fig. 9). These
parameters were as follows: L∞=144 mm,
K=0.31 year-1 for females; and L∞=132
mm, K = 0.27 year-1,
1417 Janbaz et al., Age and growth of bigeye kilka (Clupeonella grimmi Kessler, 1877) in …
Figure 5: Monthly length-frequency distribution of Clupeonella grimmi in the
Iranian waters of the Caspian Sea.
Figure 6: Mean (± S.E) of the condition factor of Clupeonella grimmi in the Iranian
waters of the Caspian Sea.
Iranian Journal of Fisheries Sciences 15(4) 2016
Figure 7: Annually changes of condition factor (mean ± S.E) of Clupeonella grimmi in the
Iranian waters of the Caspian Sea.
Figure 8: Age composition of Clupeonella grimmi in Iranian waters of the Caspian Sea.
Figure 9: Theoretical growth curve for fork length of Clupeonella grimmi in Iranian
waters of the Caspian Sea.
1418
1419 Janbaz et al., Age and growth of bigeye kilka (Clupeonella grimmi Kessler, 1877) in …
for males (Table 3). The growth
performance index (Φ') was 3.688 for both
sexes. The K value and L∞ of females was
more than that of males.
Table 2: Fork length (mm) means of Clupeonella grimmi in Iranian waters of the Caspian Sea.
Age groups
Fork length
(mm)
2
3
4
5
6
7
Female
N
91±2.91
10
104±4.3
6
114.5±7.8
90
125.8±4.5
44
130±5.9
55
Male
90±1.5
105.5±6.3
110.8±5.7
120.3±5.5
128±11.6
N
4
8
15
9
6
Total
N
90.2±1.4
14
105±8.7
14
110.2±7.8
105
124.4±9
53
130.5±8.4
61
134.9±5.8
15
135±8.4
15
Table 3: Von Bertalanffy Parameters of growth equation C. grimmi from various localities and at different
times.
Area
L∞
K
t0
Reference
Iranian waters
125.5
0.467
0.33
Pourgholam,1996
Azerbaijan waters
123.8
0.414
0.41
Mamedov, 2006
Iranian waters
Iranian waters
Iranian waters
143.6
142
148
0.267
0.280
0.290
1.74
1.39
1.12
Parafkandeh, 2009
Fazli et al., 2009
Present study
Discussion
In the present study, the exponent (b) of
length-weight relationship for both sexes
was 3.16. Bagenal (1978) indicated that
the exponent usually ranges from 2.30 to
5.50 and it is generally more than 3. The
relationship between fork length and body
weight was calculated for each sex. Values
of the isometric coefficient (b) were 3.14
and 3.16 for females and males,
respectively, which were not significantly
different from 3, indicating that the growth
of C. grimmi is isometric. Similar
isometric growth was reported by
Mamedov (2006), but with over b (3.56) in
Azerbaijan off waters. In contrast, Fazli et
al. (2009) reported a lower exponent of the
Von Bertalanffy equation (b = 2.89) and
also less than that (2.61) reported by
Parafkandeh (2009). It is thus difficult to
compare the values of previous studies
with those in the literature, as the recent
reported data are insufficient; especially
new data in the southern part of the
Caspian Sea is needed.
Length can vary with geographical
location that is probably related to some
factors such as climate, trophic status and
diet, destruction of natural habitats,
unsuccessful natural breeding, fishing
pressure (Koliev, 1997). During surveys
along the central Caspian off Azerbaijan
between 1995-2004, the fork length and
weight means were 91 mm and 8.5 g
Iranian Journal of Fisheries Sciences 15(4) 2016
(Mamedov, 2006). Pourgholam (1996) in
hydroacoustic project reported that fork
length and weight means of C. grimmi
were 108 mm and 9.5 g in Iranian waters.
During the years between 1995-2001, fork
length and weight means of C. grimmi
were 103.6 mm and 7.6 g, respectively, in
Iranian waters (Fazli et al., 2009).
Parafkandeh reported that fork length
means were 117.5 mm and most of them
were in range of 120-130 mm, while the
majority of the fish were 87 to 107 mm in
1997 (Fazli, 2002). In this study, the
average fork length and weight of C.
grimmi obtained as 118 mm and 12.7 g.
Therefore, at the present study an increase
in fork length and weight means of C.
grimmi was observed than the previous
years. This is due to the abundance of fish
less than 90 mm severe and even reached
zero and despite of big fish, the average
length increased. Comparing these data
suggest that the length range of the species
is more limited and most of the larger fish
are caught.
The length- frequency distribution has
the largest span in December to February
and then span changes from February to
April from maximum 142.5 to 127.5 mm.
This could be due to migration of females
and males to these areas for spawning.
Generally, the highest values of
condition factor were observed in
December and April. The increase
condition factor in December and April
could be due to gonadal development and
feeding
activity.
Condition
factor
decreased in winter (January) because of
decreased
food
intake
and
low
temperature. Similar fluctuation in
1420
condition factor was reported by Fazli et
al. (2009). In addition condition factor
increased to the highest levels in 2004 to
2006 rather than to previous years. After
Mnemiopsis appeared in 1999 in the
Caspian Sea (Ivanov et al., 2001), they fed
aggressively zooplankton, fish eggs and
fish larvae (Mutlu, 1999) and particularly
on zooplankton, which is the food for
zooplanktivorous kilka (Kideys et al.,
2001; 2005). According to Fazli et al.
(2007a), the biomass of the most abundant
species of Kilka, anchovy, and bigeye
Kilka decreased during 2000-2004. They
found that the trends for biomass and
condition factor were opposite, therefore
prior to the stock collapse, the species may
have been food-limited. Subsequent to the
collapse, perhaps reduced inter and intraspecific competitions may have led to
increased availability of zooplankton for
the diminished Kilka stocks in which their
condition increased.
Data on sex-ratio in different sizes
showed absence of males above 102 mm.
Females were the dominant sex and also
were dominant in length class above 102
mm. This indicates that females grow to a
larger size than males. The overall ratio of
males to females (male:female) was
0.33:1, significantly different from 1:1,
and there was a month difference in sex
ratios except for January. This estimate
differs from the ratio of 2.12:1 that
reported by Besharat and Khatib (1993)
who collected samples from 1990-1991
using a conical lift net with underwater
electric lights. During hydro-acoustic
biomass assessments which conducted in
1996 and 1997, some samples were
1421 Janbaz et al., Age and growth of bigeye kilka (Clupeonella grimmi Kessler, 1877) in …
collected by mid-water trawl in the Iranian
waters of the Caspian Sea (Fazli and
Besharat, 1998). In the shallow waters,
males were most abundant but in the deep
waters females predominated in all
seasons. These results confirm that the
behavior of females changed with respect
to underwater electric lights during the
year. Ben-Yami (1976) reported that fish
display different behaviors as they respond
to light. A key hypothesis is that fish are
attracted to light to feed. With the
development of the gonads, females
appeared to be less attracted to light. As
they approached to the spawning time,
they ceased feeding and disappeared from
the catches. In case of males, they
continued to feed during spawning and it
seemed that their response to light
remained unchanged (Ben-Yami, 1976).
Dominance of female may be attributed to
anyone or more the following factors as
reported by Del-Zarka and Sedfy (1970):
(a) segregation of the sexes through
various periods of the year including
segregation resulting from sex differences
in age and size at maturity, (b) gear
selectivity in relation to sex differences in
morphology and in physiological activity
and (c) differences in natural and fishing
mortalities between sexes and (d) greater
activity which in turn increase changes of
being caught. As the bigeye Kilka biomass
collapsed due to the combined effects of
fishing mortality and the decline in
available prey species (Fazli et al., 2007),
dominant females in the C. grimmi
population, probably a reaction to
declining fish stocks of the struggle for
survival through the spawning and
reproduction more.
Negative values of the t0 are frequent
among species with rapid growth during
the 1st year and reduced growth rates in
subsequent year (Peres and Haimovic,
2004). Difference in length-at age and
growth were observed between the males
and females of C. grimmi in the southern
of the Caspian Sea. Female fish often
attains greater size than males, usually
through a faster growth rate (Paker, 1992).
At the present study for the age group of 23 yr, the rate increments were 13 mm/yr
(20%) for females and 15.5 mm/yr (20%)
for males. Subsequently, growth rate
(length) in females and males trended to
decrease and the growth rate differed
significantly between sexes (about 5 mm/y
(10 %). Females of C. grimmi grow fast in
the second-third year of the life but, after
that, grow more slowly than females.
Several factors might be responsible for
this growth different between males and
females, for example, physiological
changes influenced by temperature
changes, feeding regimes and productive
cycles (Utagawa and Taniuchi, 1999).
There are few papers from previous
studies about growth parameters in C.
grimmi for comparing with the results of
present study in the southern of the
Caspian Sea. However, Table 3 gives
some comparison between previous and
present studies. Branstetter (1987)
categorized K value as 0.05-0.10/yr for
slowly growing species, 0.10-0.20/yr for
species with average growth, and 0.200.50/yr for rapidly growing species. The
results showed that C. grimmi in the
Iranian Journal of Fisheries Sciences 15(4) 2016
southern Caspian Sea have a rapid growth
rate. This could be due to available food
resources,
optimum
temperature
(temperate area) and salinity. Hong-Jing
and Cong-Xin (2008) reported that there
are several factors affecting growth rate
such as shortage of food, deprivation
caused by migration and changes of
temperatures. The K and L∞ ratio are
important parameters in the growth
patterns used by Pauly et al. (1988). In this
study, The K value and L∞ of females of C.
grimmi had relatively more than that of
males.
Consequently, the finding on age and
growth of C. grimmi from this research
will help to elucidate the distribution with
age of fish and their sustainable
management. Moreover, further research is
needed to complete the studies on age and
growth during annual cycle.
Acknowledgement
We are grateful to Iranian Fisheries
Organization for funding this study under
the project No. 2-032-200000-02-860186001. We are grateful to Dr.
Nasrollahzadeh for assistance and
cooperation in execution of this project.
We thank the following individuals,
organizations and cooperatives societies
responsible for their assistance in
obtaining samples for this project
including Dr. Ghasemi, Kor, Daryanabard,
and the rest of the staff at the Caspian Sea
Ecological Research Center.
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1426 Janbaz et al., Age and growth of bigeye kilka (Clupeonella grimmi Kessler, 1877) in …
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