ISSN 10630740, Russian Journal of Marine Biology, 2010, Vol. 36, No. 3, pp. 157–166. © Pleiades Publishing, Ltd.

, 2010.
Original Russian Text © A.V. Zavolokin, 2010, published in Biologiya Morya.

ECOLOGY

Distribution and Abundance Dynamics of Jellyfish

in the Sea of Okhotsk1
A. V. Zavolokin
Pacific Fisheries Research Center, Vladivostok, 690950 Russia
email: zavolokin@tinro.ru
Received November 19, 2009

Abstract—The seasonal and interannual trends in the distribution and abundance of jellyfish (Scyphozoa and
Hydrozoa) in the epipelagic and mesopelagic areas of the Sea of Okhotsk during 1992–2005 were examined
on the basis of trawl survey data. The area of occurrence, biomass, and the numbers of Scyphozoa in the epi
pelagic layer were the smallest in spring; in summer and fall their abundance sharply increased and then
decreased in winter. In contrast to the epipelagic zone, the numbers of scyphomedusae in the mesopelagic
layer were significantly lower in the summer than in the winter and spring. This probably indicates that a part
of scyphomedusae winter in the mesopelagic. Hydrozoa in both the epi and mesopelagic areas were more
numerous in the winter and spring. Jellyfish biomass and abundance greatly changed from year to year. Thus,
in fall the biomass of scyphomedusae and hydromedusae in the epipelagic zone varied from 166 to 1271 and
from 6 to 49 kg/km2, respectively.

Key words: Scyphozoa, Hydrozoa, jellyfish, biomass, abundance, distribution, Sea of Okhotsk.
DOI: 10.1134/S1063074010030016

1
To date, little information is available on the jelly
fish of the Sea of Okhotsk. The most recent prominent
works dealing with the taxonomy and biogeography of
cnidarians of the Sea of Okhotsk were published about
a century ago [8, 9]. Various aspects of the ecology of
jellyfish, such as distribution, total biomass and abun
dance, nutrition, trophic relationships, etc., have been
studied insufficiently.
Some information on the distribution and abun
dance of jellyfish in the Sea of Okhotsk can be
obtained from trawl surveys, which have been annually
performed during the last several decades by the staff
of the Pacific Fisheries Research Center (TINRO
Center). The trawling gear used in these surveys is pri
marily designed for fish and squid abundance assess
ment; however, it permits an approximate estimation
of the occurrence and abundance of large jellyfish (>
3–5 cm).
Estimates of jellyfish abundance in the Sea of
Okhotsk based on trawl survey data were first provided
by Shuntov et al. [15]. These authors have shown that
gelatinous zooplankton have a significant biomass and
can obviously play a substantial role in the pelagic
communities. Results of the study of jellyfish species
composition, distribution, and abundance were after
wards supplemented by information on diets and the
trophic status of gelatinous zooplankton [2, 6, 7].
However, the published data are limited to shortterm
1 The article was translated by the authors.
observation series (1–5 years), usually relating to one
season and only to the epipelagic zone. In the present
paper, we summarize the results of the study of jellyfish
distribution and abundance in the epi and mesope
lagic zones of the Sea of Okhotsk in 1992–2005.
MATERIALS AND METHODS
This work uses trawl survey data on the quantitative
composition of jellyfish in the epipelagic and mesope
lagic zones of the Sea of Okhotsk in 1992–2005. A
total of about 4600 trawlings were performed (Table 1).
The epipelagic zone was studied most completely:
about 80% of the trawl stations were made in the
0⎯200 m layer. In the mesopelagic zone, the surveys
were limited to the 500 m depth. The region of study
included the entire northern Sea of Okhotsk, as well as
waters of western Kamchatka and eastern Sakhalin.
For a more correct analysis of the seasonal and inter
annual dynamics of jellyfish, it was divided into four
large areas: eastern Sakhalin, western Kamchatka,
northern Sea of Okhotsk, and Shelikhov Bay (Fig. 1).
Jellyfish were caught using RT80 and RT108
pelagic trawls (vertical opening 30–60 m) with a
smallsize meshed (1 cm) insertion at the cod end.
The abundance (biomass) of jellyfish was determined
by the formula
n ( b) ,
N ( B ) = 
vtak

157
158 ZAVOLOKIN

Table 1. The time of surveys and the number of trawlings in Since a significant proportion of jellyfish are sieved
the Sea of Okhotsk off through the trawl nets, an allowance for catchabil
ity is made in the estimation of jellyfish abundance and
Number
Year Period of survey
of trawlings
biomass. In our calculations, the catchability coeffi
cient was 0.1, a value generally accepted in biocoeno
1992 January–August 369 logical researches conducted by the TINROCenter
1993 July–December 268 [10–14]. Obviously, the catchability coefficient sub
stantially differs among the species and size groups of
1994 July–December 280 jellyfish; however, because no special study of this issue
1995 July–December 369 was made, we used a general allowance for the catch
1996 July–September 129 ability of the trawl nets.
1997 July–August 179 The horizontal opening of the trawl mouth was cal
1998 February–November 528 culated by the formula, which takes into account the
vertical opening of trawl, the velocity of towing, the
1999 March–October 383
depth of trawling, and the wire length [1].
2000 March–November 493
The species identification of jellyfish followed the
2001 April–November 485 works of Naumov [8, 9] and Kramp [18]. Because of
2002 March–December 484 incorrect identifications of jellyfish in some surveys,
2004 April–November 361 Aurelia limbata and Phacellophora camtschatica were
united into one group.
2005 April–November 282

RESULTS AND DISCUSSION

where N(B) is the abundance (biomass) of species,
Composition and Distribution
kg/km2; n(b) is the actual abundance (mass) of species
in a catch, kg; v is the velocity of trawl towing, km/h; During the study period, 13 species of jellyfish were
t is the duration of trawling, h; a is the horizontal recorded for the Sea of Okhotsk, 9 species belong to
opening of the trawl mouth, km; and k is the catch the class Scyphozoa (Aurelia aurita Linnaeus, 1758;
A. limbata Brandt, 1838; Atolla wyvillei Haeckel, 1880;
ability coefficient. Chrysaora helvola Brandt, 1838; C. melanaster Brandt,
1838; C. quinquecirrha Desor, 1848; Cyanea capillata
Linnaeus, 1758; Periphylla periphylla Peron et
62° Lesueur, 1809; and Phacellophora camtschatica
Brandt, 1838) and 4 species belong to the class Hydro
zoa (Aequorea forskalea Peron et Lesueur, 1809; Caly
SHB copsis nematophora Bigelow, 1913; Ptychogena lacteal
A. Agassiz, 1865; and Tima sachalinensis Bigelow,
59°
1913). Most of these species occur in both the epipe
lagic and mesopelagic layers at depths down to 300–
500 m. Mesopelagic species include scyphomedusae
A. wyvillei and P. periphylla, which are episodically
NSO encountered in the epipelagic [7, 9], as well as the
Kamchatka

56°
hydromedusa C. nematophora, which makes migra
WK tions towards the surface at night.
All species of the freeswimming jellyfish of the
ES class Scyphozoa recorded by Naumov [9] for the Sea
53° of Okhotsk were found in the trawl catches, except
Cyanea purpurea Kishinouye 1910. Only 4 out of
186 jellyfish species of the class Hydrozoa characteris
tic of the Sea of Okhotsk [8] were encountered in the
Sakhalin

catches. Evidently, due to their small size (mostly no

50° larger than 3–5 cm) and feeble body, the hydromedu
sae are washed out of the trawl sac during trawling. A
distinctive feature of species of the class Hydrozoa is
140° 145° 150° 155° 160° the short medusoid stage in their life history (lacking
in some species) [8], which decreases the probability
Fig. 1. A schematic map showing the region of study (Sea of their occurrence in trawl catches. Thus, the data on
of Okhotsk). Areas: ES, eastern Sakhalin, WK, western
Kamchatka, NSO, northern Sea of Okhotsk, SHB—She the distribution and abundance of hydrozoan jellyfish
likhov Bay. provided in this paper refer to a small portion of spe

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 DISTRIBUTION AND ABUNDANCE DYNAMICS OF JELLYFISH 159

62°
April–June, 2004 July–August, 1997

59°

56°

1
53° 2

4
50°
5
62°
September–November, 2003 November–December, 2002

59°

56°

53°

50°

140° 145° 150° 155° 160° 140° 145° 150° 155° 160°

Fig. 2. The distribution of biomass (kg/km2) of scyphomedusae in the Sea of Okhotsk. Designations: 1, no catch; 2, less than 50;
3, 50–300; 4, 300–2000; 5, more than 2000; dashed line, 200 m isobath.

cies of Hydrozoa, which have larger dimensions and a
more rigid body.
The distribution and occurrence of jellyfish mark
edly change during the year. In spring, the distribution
area of scyphomedusae is the smallest, their abun
dance is low, and the main scyphomedusan concentra
tions in this period are confined to the western Kam
chatka shelf (Fig. 2A). During summer and fall, scy
phomedusae are found almost throughout the region
studied (Figs. 2B, 2C), and their densities are mark
edly increased. In fall–early winter, despite their wide
occurrence, the abundance of scyphomedusae begins
to decrease (Fig. 2D).

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160 ZAVOLOKIN

62°
April–June, 2004 July–August, 1997

59°

56°

53° 1

3
50°
4
62°
September–November, 2003 November–December, 2002

59°

56°

53°

50°

140° 145° 150° 155° 160° 140° 145° 150° 155° 160°

Fig. 3. The distribution of biomass (kg/km2) of hydromedusae in the Sea of Okhotsk. Designations as in Fig. 2.

The distribution of hydromedusae does not change
as markedly as that of scyphomedusae from season to
season. The distribution area of Hydrozoa includes the
entire northern part of the Sea of Okhotsk (Fig. 3).
The largest concentrations are, as a rule, observed out
side the shelf in the central part of the sea. In contrast
to scyphomedusae, hydromedusan densities in winter
and spring are somewhat higher than in summer and
fall.
Seasonal Dynamics of Abundance
Comparison of average biomasses of jellyfish in the
epipelagic zone during the period of 1992–2005
showed that the seasonal dynamics of scyphomedusae

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 DISTRIBUTION AND ABUNDANCE DYNAMICS OF JELLYFISH 161

700 60
(a) (b)
600 50
Biomass, kg/ km2

500
40
400
30
300
20
200
100 10

0 0
Spring Summer Autumn Winter Spring Summer Autumn Winter
Chrysaora melanaster Tima sachalinensis
Cyanea capillata Ptychogena lactea
A. limbata + P. camtshchatica Other

Fig. 4. The seasonal dynamics of biomass (kg/km2) of scyphomedusae (A) and hydromedusae (B) in the epipelagic of the Sea of
Okhotsk.

(Fig. 4A) and hydromedusae (Fig. 4B) are in counter
phase. Species of the former class reach maximum
biomass in the warm season, while hydromedusae are
the most abundant during the cold season. In the epi
pelagic, C. melanaster and C. capillata dominate
throughout the year among Scyphozoa, T. sachalinen
sis and P. lacteal dominate among Hydrozoa (Table 2).
The increased abundance of scyphomedusae in the
epipelagic zone in the warm season well agrees with
their life history features. In spring and summer, the
active strobilation of polyps and the release of large
numbers of juvenile jellyfish take place, during sum
mer and fall the juveniles rapidly grow, reaching
high biomass toward the end of fall. However,
the highest concentrations of one of the dominant
species—C. melanaster—were recorded in summer,
while in fall its biomass decreased (Table 2).
Similar seasonal dynamics of C. melanaster were
observed in the Bering Sea, where the biomass of this
species in summer was almost two times higher than in
winter [5].
In the mesopelagic zone, the abundance of jellyfish
also changes seasonally (Table 3, Fig. 5); however,
unlike the epipelagic zone, their biomass here
decreases in the warm season but increases in the cold
period. Probably, due to the cooling of surface water,
in winter a part of jellyfish descend into greater depths
and in spring migrate back into the epipelagic zone.
This is suggested by the presence of large jellyfish in
catches in early spring [2, 6]. In addition, mature
specimens of C. capillata with welldeveloped sex
glands were repeatedly encountered in the northern
Sea of Okhotsk in spring 2007. The available data on
the abundance of jellyfish in the mesopelagic area in
summer and fall are very scarce and are only limited to
two surveys.

Table 2. The biomass of jellyfish in the epipelagic of the Sea of Okhotsk in different seasons (M ± SD, kg/km2)
Class, species Spring Summer Fall Winter
Scyphozoa 261 ± 272 434 ± 776 555 ± 595 481 ± 411
Chrysaora melanaster 111 ± 103 388 ± 772 142 ± 235 171 ± 197
Cyanea capillata 125 ± 254 40 ± 63 264 ± 340 261 ± 247
Aurelia limbata + 24 ± 48 5±7 133 ± 285 48 ± 65
Phacellophora camtschatica
Hydrozoa 33 ± 34 8 ± 13 17 ± 23 46 ± 62
Tima sachalinensis 16 ± 26 4 ± 11 9 ± 15 29 ± 54
Ptychogena lactea 13 ± 23 1±3 4±8 6 ± 10
N 6 12 9 4
Note: Here and in Tables 3–6: M is the mean value; SD, standard deviation; N, the number of surveys.

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162 ZAVOLOKIN

Table 3. The biomass of jellyfish in the mesopelagic zone of the Sea of Okhotsk in different seasons (M ± SD, kg/km2)
Class, species Spring Summer Winter
Scyphozoa 249 ± 376 38 ± 52 333 ± 414
Chrysaora melanaster 157 ± 241 38 ± 52 122 ± 121
Cyanea capillata 44 ± 79 – 47 ± 71
Aurelia limbata + 47 ± 125 – 43 ± 100
Phacellophora camtschatica
Hydrozoa 74 ± 95 37 ± 19 150 ± 134
Tima sachalinensis 55 ± 93 22 ± 16 133 ± 128
Ptychogena lactea 4±4 0.3 ± 0.6 3±3
N 6 3 3
Note: Here and in Tables 4–6: “–“ signifies no data.

In the mesopelagic zone, C. melanaster dominated
among Scyphozoa during the entire year (Fig. 5A). In
the spring and winter, the biomass of other species
increased substantially: C. capillata, A. limbata, and
P. camtschatica. Among Hydrozoa, T. sachalinensis
absolutely dominates in terms of biomass (Fig. 5B).
Interannual Dynamics of Abundance
The largest data series on the interannual dynamics
of jellyfish biomass and abundance exist for the fall
season in the epipelagic zone (Tables 4 and 5). The
abundance of jellyfish greatly varies from year to year.
In fall 1993–2005, the average biomass of scyphome
dusae in the northern Sea of Okhotsk varied from 166
to 1271 kg/km2 and of hydromedusae, from 6 to
49 kg/km2 (Table 4). The average abundance of scy
phomedusae was 173–1591 indiv/km2 and of
hydromedusae, 146–2789 indiv/km2 (Table 5).
The highest biomass and abundance values of jelly
fish were observed in the 1990s (Tables 4 and 5). In the
early 2000s, a dramatic decrease in the abundance of
jellyfish, especially scyphomedusae, occurred in all
areas. This might be accounted for by the changes in
the oceanographic conditions observed in 1998–2001.
By the values of nearbottom water temperature and
ice cover, the investigated period was the coldest over
the last decades [3]. The decrease in the jellyfish bio
mass in 1998–2002 in the Sea of Okhotsk was observed
both in the summer and in the fall (Fig. 6). In abnor
mally warm years (1996 and 1997), the jellyfish abun
dance was markedly higher than the average. In years
when ice cover was close to the normal, there was no
clearcut relationship between jellyfish biomass and
thermal conditions. Thus, the maximum biomass of

350 160
(a) (b)
300 140
Biomass, kg/ km2

250 120
100
200
80
150
60
100 40
50 20
0 0
Spring Summer Autumn Winter Spring Summer Autumn Winter
Chrysaora melanaster Tima sachalinensis
Cyanea capillata Ptychogena lactea
A. limbata + P. camtshchatica Other

Fig. 5. The seasonal dynamics of biomass (kg/km2) of scyphomedusae (A) and hydromedusae (B) in the mesopelagic zone of the
Sea of Okhotsk.

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 DISTRIBUTION AND ABUNDANCE DYNAMICS OF JELLYFISH 163

Table 4. The biomass of jellyfish in the epipelagic of different areas of the Sea of Okhotsk in the fall in different years
(M ± SD, kg/km2)
Areas 1993 1994 1998 1999 2000 2001 2002 2004 2005
Scyphozoa
WK 1188 1928 ± 5564 1310 ± 4177 2041 ± 2885 930 ± 1480 392 ± 1480 115 ± 113 82 ± 122 163 ± 334
SHB 460 ± 663 838 ± 1603 851 ± 991 174 ± 257 71± 69 488 ± 818 – 1146 ± 808 –
NSO 648 ± 986 2059 ± 7461 294 ± 570 117 ± 259 98 ± 246 122 ± 231 424 ± 558 921 ± 2738 –
ES – 260 ± 204 166 ± 342 465 ± 600 40 ± 62 82 ± 144 106 ± 162 61 ± 83 169 ± 207
All 766 1271 655 699 285 271 215 552 166
Hydrozoa
WK – 74 ± 150 97 ± 226 19 ± 52 15 ± 45 43 ± 92 9 ± 15 13 ± 45 6 ± 13
SHB – – 2±7 – – 1±4 – – –
NSO 6 ± 10 60 ± 137 29 ± 68 19 ± 58 31 ± 112 27 ± 75 22 ± 45 16 ± 49 –
ES – 12 ± 19 20 ± 55 4±8 3±8 8 ± 14 3±8 2±5 8 ± 17
All 6 49 37 14 16 20 12 10 7
Number of stations
WK 1 40 52 43 47 48 5 14 13
SHB 13 14 14 12 13 12 – 3 –
NSO 17 31 55 36 48 49 28 50 –
ES – 6 30 26 32 40 32 40 13
All 31 91 151 114 140 149 65 107 26
Note: Here and in Tables 5, 6, the names of the areas are the same as in Fig. 1.

Table 5. The abundance of jellyfish in the epipelagic of different areas of the Sea of Okhotsk in the fall in different years
(M ± SD, indiv/km2)
Areas 1993 1994 1998 1999 2000 2001 2002 2004 2005
Scyphozoa
WK 4159 230 ± 204 1834 ± 4153
1895 ± 4248 590 ± 799 501 ± 898 116 ± 88 79 ± 110 259 ± 428
SHB 207 ± 175 139 ± 182 1430 ± 1433
708 ± 1349 225 ± 245 518 ± 769 – 1863 ± 843 –
NSO 407 ± 354 440 ± 217 361 ± 569 113 ± 203 239 ± 366 163 ± 430 322 ± 336 2166 ± 6664 –
ES – 74 ± 88 151 ± 212
1099 ± 4205 53 ± 64 81 ± 104 82 ± 95 109 ± 133 358 ± 378
All 1591 221 944 954 277 316 173 1054 309
Hydrozoa
WK – 1153 ± 3382 669 ± 1750 536 ± 2449 915 ± 4042 9971 ± 42380 1025 ± 2272 206 ± 517 607 ± 1430
SHB – – 8 ± 31 – – 325 ± 1127 – – –
NSO 146 ± 306 1841 ± 8672 736 ± 2040 279 ± 611 1430 ± 5737 607 ± 1683 1604 ± 5354 771 ± 2313 –
ES – 1771 ± 2937 126 ± 405 102 ± 304 245 ± 642 254 ± 494 250 ± 765 211 ± 693 596 ± 1285
All 146 1588 385 306 863 2789 960 396 602

scyphomedusae was observed in 1992; the ice cover in
1991–1992 was rather close to the manyyear average
(Fig. 6).
The effect of climatic conditions on jellyfish abun
dance has been addressed in a review paper by Purcell
[20], who concluded, based on literature data, that the
abundance of most species of gelatinous organisms in
temperate latitudes depends on temperature condi
tions. With relatively higher temperatures, the size of
the population increases, as do the rates of sexual and
asexual reproduction. Evidently, in the Sea of Okhotsk
with harsh thermal conditions, the effect of the tem
perature factor on jellyfish abundance is fairly signifi
cant.
The pattern of change of total jellyfish abundance
in different areas of the Sea of Okhotsk was not the

RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 36 No. 3 2010

164
20
15
2 1200
Winter ice cover anomaly, % 10
5
0 800
1992 1993 1994 1995
–5
1
–10
–15
–20
–25
Fig. 6. Interannual variations in the average winter ice cover (1) [3] and the biomass of scyphomedusae in summer (2) and fall (3)
in the epipelagic of the Sea of Okhotsk. Standard errors are provided for mean biomasses of jellyfish.
same in different periods (Tables 4–6). Thus, a
marked decrease in the biomass of scyphomedusae at
the turn of the 21st century began in 1998 in the cold
est areas of that sea: in the northern Sea of Okhotsk
and off eastern Sakhalin. In Shelikhov Bay the
decrease in scyphomedusae abundance occurred a
year later. In the western Kamchatka area, which is
influenced by warm oceanic waters, a marked decrease
in jellyfish biomass was observed in 2000–2001. We
note that the interannual dynamics of abundance of
different jellyfish species within the same area also dif
fers substantially [7].
In terms of biomass, scyphomedusae absolutely
dominated over hydromedusae. The average biomass
values in the two classes in the epipelagic zone for the
fall 1993–2005 were 542 and 19 kg/km2. At the same
time, the abundance of scyphomedusae and hydrome
dusae was comparable: the average was 649 and
843 indiv/km2 for the investigated period, respectively.
In the epipelagic zone, the dominance of scyphome
dusae by biomass was observed in all seasons (Table 2,
Fig. 4). In the mesopelagic zone, where hydromedusae
were more abundant than in superficial water layers,
the biomass of the two jellyfish classes varied from
50 to 300 kg/km2 (Table 3, Fig. 5). The sharp yearto
year variation in jellyfish abundance observed in the
fall period also occurs in other seasons, both in the
epipelagic zone and in the mesopelagic zone. Thus, in
the spring the biomass and abundance of scyphome
dusae in the mesopelagic zone respectively varied from
7 to 943 kg/km2 and from 13 to 681 indiv/km2 (Table
6). The interannual fluctuations of biomass and abun
dance of hydromedusae were no less significant.
ZAVOLOKIN
1400
1000
Biomass, kg/ km2
1996 1997 1998 1999 2000 2001 2002 2004
3 600
400
200
0
When analyzing the abundance dynamics of jelly
fish, we took that fact that the time of the survey sub
stantially influences jellyfish biomass and abundance
estimates into account. For convenience, all available
materials were grouped by seasons (the data were
insufficient for further subdivision). However, due to
the high growth rates of jellyfish, their abundance even
within one and the same season, for example in the
beginning and at the end of summer, can greatly differ,
thus leading to increased variability of biomass and
abundance estimates. The different time of surveys is
not the only factor that determines jellyfish abundance
dynamics. Results from a 6year study performed in
the Bering Sea in one and the same season and in one
and the same area also indicated great yeartoyear
variations in jellyfish abundance [4]. When describing
periodic outbreaks of jellyfish, ctenophores, and
siphonophores in different parts of the world’s ocean,
Mills [19] uses the term “bloom,” which characterizes
the magnitude of their abundance fluctuations well.
The causes of the regular drastic changes in the
numbers of jellyfish in different regions of the world’s
oceans are unknown thus far. Few regional studies
exist that have revealed a fairly distinct relationship
between environmental factors and jellyfish abun
dance. Thus, it was suggested that the increase in the
numbers of scyphomedusae Chrysaora quinquecirrha
and Aurelia aurita in the Gulf of Mexico was a conse
quence of the increased trophic flow due to eutrophi
cation and overfishing [17]. However, in most cases, it
is impossible to find the causes of the observed
changes, even on the basis of longterm observation
series. Brodeur et al. [16], who had at their disposal
RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 36 No. 3 2010
 DISTRIBUTION AND ABUNDANCE DYNAMICS OF JELLYFISH 165

Table 6. The biomass (M ± SD, kg/km2) and abundance (M ± SD, indiv/km2) of jellyfish in the mesopelagic zone of different
areas of the Sea of Okhotsk in spring in different years
Areas 1992 1998 1999 2002 2004 2005
Biomass
Scyphozoa
WK 80 ± 86 7 ± 49 95 ± 416 5 ± 18 303 ± 625 900 ± 1100
NSO – – 68 ± 176 9 ± 27 49 ± 98 985 ± 1317
All 80 7 81 7 176 943
Hydrozoa
WK 56 ± 52 1 ± 13 316 ± 815 4 ± 19 90 ± 110 99 ± 122
NSO – – 114 ± 244 3 ± 12 45 ± 88 9 ± 18
All 56 1 215 4 68 54
Abundance
Scyphozoa
WK 179 ± 156 14 ± 83 158 ± 527 7 ± 24 488 ± 1096 661 ± 980
NSO – – 73 ± 158 18 ± 43 143 ± 218 701 ± 1201
All 179 14 115 13 316 681
Hydrozoa
WK 3240 ± 2486 144 ± 1081 6986 ± 22776 43 ± 198 3699 ± 6244 3012 ± 5620
NSO – – 3915 ± 9359 75 ± 216 3374 ± 6628 194 ± 108
All 3240 144 5450 59 3537 1603
Number of stations
WK 6 118 78 41 44 111
NSO – – 14 21 26 127
All 6 118 92 62 70 238

25year observation data on the abundance dynamics ACKNOWLEDGEMENTS

of jellyfish in the Bering Sea, came to the conclusion
that the exiting data are insufficient to reveal the fac
tors determining their abundance and biomass.
Variations in the abundance of jellyfish are evi
dently due, in addition to external influences, to their
lifehistory features. Most species of freeswimming
jellyfish found in the trawl catches are characterized by
the alternation of sexual and asexual generations, with
the former (medusae) occurring in the pelagic and the
latter (polyps) on the bottom. Without doubt, the
influence of one and the same environmental factor
on different lifehistory stages can vary, thus adding
further difficulty to the study of the causes that lead to
jellyfish abundance fluctuations. Therefore, jellyfish
abundance variations have been thus far associated
only with the extreme influences of external factors,
such as the abnormally cold period of 1998–2001 in
the Sea of Okhotsk or the strong eutrophication of
coastal waters in the Gulf of Mexico.
This study was performed with partial support by
Grant no. 090400769a from the Russian Founda
tion for Basic Research.
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Vol. 36 No. 3 2010