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Seasonal occurrence and vertical

distribution of appendicularians in
Toyama Bay, southern Japan Sea
MIKA TOMITA1,*, NAONOBU SHIGA AND TSUTOMU IKEDA
BIOLOGICAL OCEANOGRAPHY LABORATORY, FACULTY OF FISHERIES, HOKKAIDO UNIVERSITY,
1PRESENT ADDRESS: JOETSU ENVIRONMENTAL SCIENCE CENTER,

-- MINATO-CHO, HAKODATE -, JAPAN

SHIMOMONZEN, JOETSU -, JAPAN

To investigate seasonal variation in the community structure of appendicularians, vertical hauls

(0500 m) with a Norpac net were made at an offshore station in Toyama Bay at intervals of 24
weeks from February 1990 to January 1991. Additional samples were collected with MTD nets
at 1217 different depth layers between the surface and a depth of 600700 m at the same position
in June, September and December 1986, and March 1992, to examine the vertical distribution of
appendicularians. Twenty-one species (including two unidentified species) belonging to five genera were
found, and the dominant species were divided into three groups by their occurrence period. Oikopleura
longicauda, Fritillaria borealis f. typica and F. borealis f. sargassi occurred throughout most of the
year. Fritillaria pellucida, O. fusiformis and O. rufescens were found in summer and autumn. Oikopleura dioica was found in spring and winter. Oikopleura longicauda was overwhelmingly the most
abundant species throughout the year. This species was always distributed in the upper 100 m depth,
with a peak at a depth of 050 m that corresponded to the peak of chlorophyll a concentration during
the day and night in all seasons. The daynight vertical distribution patterns of F. borealis f. typica,
F. pellucida, O. fusiformis and O. rufescens were similar to that of O. longicauda. Seasonal variations in abundance of appendicularians are considered to be the result of biological factors rather
than physical factors such as temperature and salinity. In particular, O. dioica seems to be affected
by food availability.

I N T RO D U C T I O N
The establishment of in situ observations using SCUBA
and submersible (Alldredge, 1977, 1981; Barham, 1979),
and laboratory methods for rearing appendicularians
(Paffenhfer, 1973; Sato et al., 1999) has clarified some of
their biological properties. Their generation times are
very short; 224 days in tropical and temperate waters,
and their growth rates are higher than any other mesozooplankton (Greze, 1970; Paffenhfer, 1973, 1976;
Fenaux, 1976; Hopcroft and Roff, 1995; Nakamura et al.,
1997; Hopcroft et al., 1998). They are able to feed on
nanoplankton, picoplankton and even submicron colloids
by means of their house, a balloon-like gelatinous structure (King et al., 1980; Flood et al., 1992; Urban et al.,
1992; Bedo et al., 1993; Acua et al., 1996). In eutrophic
shallow embayments, the production of appendicularians
competes with that of copepods, which are major

components in secondary production (Nakamura et al.,

1997; Hopcroft and Roff, 1998). Therefore, the role of
appendicularians as secondary producers is unquestionably important in marine ecosystems (Tomita et al., 1999).
Quantitative studies of seasonal and vertical distributions
of appendicularians are essential to link their abundance
to their production. Knowledge of their diet and seasonal
vertical distributions is quite limited (Fenaux, 1968; Shiga,
1985, 1990).
Some seasonal distributions of appendicularians are
reported from the Japanese coastal waters facing the
Pacific Ocean and the Inland Sea (Omori, 1977; Itoh and
Iizuka, 1980; Shiga, 1985; Itoh, 1990; Nomura and
Murano, 1992; Uye and Ichino, 1995). No information,
however, is available from the Japan Sea except for Oikopleura longicauda (Tomita et al., 1999). The present study
investigates seasonal abundance and diel/seasonal
vertical distribution of appendicularians using samples

Journal of Plankton Research 25(6), Oxford University Press; all rights reserved

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*CORRESPONDING AUTHOR: tomita@jo-kan.or.jp

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collected at an offshore station in Toyama Bay, southern

Japan Sea.

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formalinseawater. For the samples collected with the

twin-type Norpac nets, only those from the 100 m mesh
net were examined in this study. Appendicularians were
sorted from entire samples or from aliquot subsamples
depending upon their abundance within the samples.
Conductivity, temperature and depth measurements
were taken to assess the vertical profiles of temperature
and salinity at the sampling site. For chlorophyll a (Chl a)
measurements, water samples were collected from 12
different depths between the surface and 400500 m.
These samples were taken to correspond with the MTD
net samples. Chl a concentrations were measured fluorometrically (Parsons et al., 1984).

R E S U LT S
Hydrography

The surface water in the upper 150 m was thermally

homogeneous (10C) from February to mid-April (Figure
2). During the warming period from April to midSeptember, a seasonal thermocline was formed around
50 m. The thermocline was most pronounced in August
and September when the surface temperature was
>26C. Surface cooling and subsequent vertical mixing
started to erode the seasonal thermocline in October, and
the upper 150 m of the water column was thermally
homogeneous by December (17C) and January (14C).
Below 150 m the seasonal variation of temperature was
small and was almost absent below 300 m. A permanent
thermocline existed around 150300 m, in which the
temperature decreased from 10C at the top of the
thermocline to 2C at the base.
Salinity in the upper 150 m also changed seasonally.
Salinity was consistently recorded as >34.2 at a depth
of 50150 m depth from June to February. Maximum
salinity >34.4 was found at 75100 m during August
and September. These high-salinity waters appear to be
the warm Tsushima Current, which is characterized by
high salinity [34.234.7; (Shuto, 1982)]. Low-salinity
water (<33.0) was observed in the upper 20 m
between August and November. Nagata and Nakura
observed that river discharge temporarily decreased the
salinity in the central part of Toyama Bay (Nagata and
Nakura, 1993). Therefore, the surface layer of the
Tsushima Current might be affected by the run-off river
water. Salinity below 300 m was homogeneous throughout the year. Water characterized by a salinity of
34.0534.10 and a temperature <1C is termed deep
water and is widespread throughout the entire Japan
Sea (Nishimura, 1969). In Toyama Bay deep water was
Fig. 1. (A) Location of Toyama Bay in southern Japan Sea. (B) The
sampling station (circled star) in Toyama Bay. The 200, 500 and 1000 m present at depths below 350 m during all seasons of the
year.
depth contours are superimposed.

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Samples were collected every 24 weeks from February

1990 until January 1991, aboard the RV Tateyama
Maru at an offshore station (3700N, 13714E; bottom
depth 1000 m) in Toyama Bay (Figure 1). Vertical hauls
were made from 500 m to the surface with a twin-type
Norpac net [45 cm mouth diameter, 330 and 100 m
mesh, (Motoda, 1957)] during predominantly daylight
hours at a speed of 1 m s1.
To study vertical distribution, samples were collected
by the RV Mizuho Maru using MTD nets [56 cm
mouth diameter, 350 m mesh, (Motoda, 1971)] from
1217 different depth horizons between the surface and
a depth of 600700 m at the same station as mentioned
above. Samples were taken on June 45, September 2,
December 56, 1986 and March 910, 1992, during both
day and night (Table I). The series of nets were towed
horizontally for approximately 3060 min at a ship speed
of 1.01.5 knots.
The volume of water passing through the twin-type
Norpac net and MTD nets was measured by a Rigosha
flow-meter mounted at the mouth of each net. All
samples were immediately preserved in 10% neutralized

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Table I: Sampling data with MTD nets at an offshore station (3700N, 13714E) in Toyama Bay
Date

Local time

Sampling depth (m)

1986
4 June

20:2521:59

5 June

07:3610:30

2 Sep.a

19:3822:08

0, 10, 20, 30, 50, 75, 125, 150, 200, 250, 300, 350, 400, 500, 600
0, 10, 30, 50, 75, 100, 150, 200, 250, 300, 350, 400, 450, 500, 550, 600, 700
10, 21, 31, 52, 77, 103, 129, 155, 187, 233, 280, 327, 373, 466, 560

2 Sep.

12:2614:52

0, 10, 31, 51, 76, 102, 153, 204, 255, 293, 342, 391, 439, 488, 537, 586

5 Dec.

18:4421:00

0, 10, 30, 50, 75, 100, 125, 150, 200, 250, 300, 350, 400, 500, 600

6 Dec.

06:4009:10

0, 10, 30, 50, 75, 100, 150, 200, 250, 300, 350, 400, 450, 500, 550, 600

1992
21:4300:20

0, 8, 24, 41, 60, 80, 100, 120, 160, 219, 263, 307, 350, 394, 438, 526, 613

10 Mar.

09:2912:00

0, 9, 25, 42, 63, 85, 106, 127, 169, 213, 256, 299, 341, 384, 427, 512, 597

Note no data at the surface on 2 September (night-time), 1986.

Fig. 2. Seasonal variations in temperature and salinity at the sampling station in Toyama Bay. Sampling dates indicated by triangles on top abscissa.

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9 Mar.

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Appendicularians

Seasonal occurrence
Total appendicularian abundance showed two prominent
peaks in February and June, respectively (Figure 3). The
maximum abundance [388 individuals m3 (ind. m3)]
occurred on June 15 and the minimum abundance (3.3
ind. m3) was recorded on October 2, 1990. Seven
dominant species are discernible, >10 ind. m3 from one
or more samples during the investigation, were divided
into three groups by their pattern of occurrence.
1. Oikopleura longicauda, F. borealis f. typica and F. borealis f.
sargassi occurred almost throughout the whole year
(Figure 3). The period of occurrence of these three
species was the same, whereas their seasonal change
in abundance was different. That is, O. longicauda was
numerically abundant from June to early July, with
maximum abundance (330 ind. m3) on June 15. The
maximum abundance (201 ind. m3) for F. borealis f.
typica was found on February 28. Fritillaria borealis f.
sargassi had two prominent peaks on July 17 (40 ind.
m3) and December 5 (44 ind. m3). Oikopleura longicauda was the most abundant species during the study
period (annual mean to the total appendicularian

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abundance: 50%, range: 991%). The numerical

peaks of the total appendicularian abundance in
June and February were composed of O. longicauda
(91% of the total) and F. borealis f. typica (74%),
respectively.
2. Fritillaria pellucida, O. fusiformis and O. rufescens were
found from July to January (Figure 3). The maximum
abundance for these species was recorded from late
August to October (23, 20 and 19 ind. m3, respectively).
3. Oikopleura dioica was found throughout the sampling
period except from August to November (Figure 3).
The maximum abundance (50 ind. m3) occurred on
June 3.
Most of the minor species only occurred for a short
time from July to January (Figure 4). and Oikopleura
labradoriensis and other Oikopleura sp. occurred irregularly
throughout the year.

Diel vertical distribution

Diel vertical distributions were observed in approximately
spring, summer, autumn and winter. Each vertical profile
of hydrography recorded on the sampling date reflected
the annual cycle of hydrography from February 1990 to
January 1991 (Figure 5). The maximum Chl a concentration in each sampling season was always found within
the upper 50 m of the water column and was low even
during the spring bloom (1.0 mg m3 in March).
Of the dominant species, O. longicauda, F. borealis f. typica,
F. pellucida, O. fusiformis and O. rufescens inhabited the upper
100 m throughout the year (Figure 5), suggesting that
their distribution is confined above the permanent
thermocline. Oikopleura longicauda had a peak in abundance in the top 50 m of the water column during both
day and night. This distribution pattern did not change
among the four seasons. Although occurrence periods of
F. borealis f. typica, F. pellucida, O. fusiformis and O. rufescens
were limited, their daynight vertical distribution patterns
were similar to that of O. longicauda. In March 1992, too
few F. borealis f. typica were collected in the night for
counting purposes. Considering that this species was
distributed in a very thin layer during the day, the MTD
nets which were towed at 1020 m depth intervals (Table
I), could not sample such a thin layer at night.
We calculated the weighted mean depth (WMD) as a
measure of the centre of vertical distribution [cf.
(Worthington, 1931; Pearre, 1973; Frost and Bollens,
1992); Table II]. The WMDs of O. longicauda, F. pellucida,
O. fusiformis and O. rufescens were deeper during the day
and shallower at night. WMD of F. borealis f. typica was
shallower during the day and deeper at night. Two other
dominant species, F. borealis f. sargassi and O. dioica, were

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A total of 21 appendicularian species (including two

unidentified species; Oikopleura sp. and Fritillaria sp.)
belonging to five genera occurred in Toyama Bay (Figures
3 and 4). Fritillaria sp. and Stegosoma magnum are not shown
in these figures, because Fritillaria sp. occurred only in
MTD net samples and S. magnum occurred only in 330 m
mesh Norpac samples. According to Tokioka, O.
labradoriensis and F. borealis f. typica are oceanic cold-water
species (Tokioka, 1955). Oikopleura dioica is an inlet/neritic
species and the remaining species, other than Appendicularia sicula and Oikopleura sp., are oceanic warm-water
species. Appendicularia sicula occurs in embayments in
temperate or tropical regions (Tokioka, 1955). Therefore,
it is likely that A. sicula is a warm neritic species. Oikopleura
sp. seems to be a new mesopelagic species.
Tokioka previously reported 21 appendicularian
species belonging to six genera, based on samples
collected at many stations from Maizuru Bay to New
Yamato Bank in the Japan Sea in April, July and September (Tokioka, 1951). Among them, O. cornutogastra
[synonym of O. fusiformis f. cornutogastra (Aida) Tokioka],
F. aberrans (synonym of F. magna Lohmann) and Pelagopleura sp. did not occur in the present collection. These
three species were collected from oceanic stations in
Tokiokas study (Tokioka, 1951). The present study is the
first to record F. borealis f. typica and F. messanensis in the
Japan Sea.

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Fig. 3. Seasonal variations in abundance of total appendicularians and seven dominant species in Toyama Bay. Open square indicates no occurrence.

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Fig. 4. Seasonal variations in abundance of 12 minor species in Toyama Bay. Open square indicates no occurrence.

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Fig. 5. Vertical distributions of the populations (%) of Oikopleura longicauda, O. rufescens, Fritillaria borealis f. typica, F. pellucida, O. fusiformis, O. labradoriensis and Oikopleura sp. during the daytime (open) and at night (shaded) in June, September and December 1986 and March 1992. For each species,
the sampling dates on which the population sizes were <10 ind. m2 are omitted, because this seemed to be insufficient to depict the vertical distribution. Figures at the bottom of each graph indicate the total number of specimens through the 0600 or 0700 m water column (ind. m2).
Asterisk denotes no data for 0 m at night. Vertical profiles of temperature, salinity and Chl a concentration are shown on the top panel. Note that
the depth scale is not even.

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Table II: Summary of weighted mean depth (WMD) and range of temperature, salinity and density
at mean WMD of dominant species
Species

WMD (m)

Jun

O. longicauda

Sept

Dec

Mar

Range of physical factors at mean WMD for each species

Ma

45

11

49

37

41

34

50

38

40

10.523.6

33.134.3

22.926.2

24

39

30

10.525.7

33.034.2

22.226.2

45

43

F. borealis f. typica

Temp (C)

Salinity ()

Density

71

61

60

12.819.7

33.234.4

24.125.6

38

19

30

10.525.7

33.034.1

22.226.1

O. rufescens

42

31

40

11.423.6

33.134.3

22.926.0

Blank columns show insufficient data or no data (no occurrence) to calculate WMD.
a Mean for each species was approximately estimated as the centre of the vertical distribution.

not collected in sufficient numbers with MTD nets for

their vertical distribution to be depicted.
Distributions of O. labradoriensis and Oikopleura sp. were
deeper than those of the dominant species (Figure 5).
Oikopleura labradoriensis was distributed between 10 and
350 m. Oikopleura sp. was absent from the epipelagic layer,
occurring between 150 and 700 m with its maximum
abundance below 350 m at each sampling date. The
distribution of O. labradoriensis was deeper during the day
and shallower at night, whereas Oikopleura sp. exhibited no
persistent pattern in its vertical distribution (daytime
WMD and night-time WMD: for O. labradoriensis 91 and
49 m in June; for Oikopleura sp. 447 and 369 m in June,
303 and 472 m in September, and 421 and 368 m in
December).
Unfortunately, statistical tests could not be performed
to measure any differences between daytime WMD and
night-time WMD, because of inadequate sample series on
each sampling date (only one pair of daynight samples).
Using Norpac net samples, a single regression analysis
was performed between abundance and the temperature,
salinity, or density to examine the seasonal variation for
the five dominant species. Habitat temperature, salinity
and density of five dominant species were represented by
the temperature, salinity and density at mean WMD
throughout the year (Table II). No significant correlations
between the abundance and habitat temperature, salinity,
or density were found for five dominant species.

DISCUSSION
Toyama Bay is a large area opening widely to the northeast. The bay is affected annually by the warm Tsushima

Current, especially from summer to autumn (Taniguchi

et al., 1997). The warm oceanic species, O. longicauda was
predominant throughout the year, while the other warmwater species increased in abundance during summer to
autumn. The large number of species (21) observed in this
study is the result of the occurrence of many warm
oceanic species (Table III). Succession from O. dioica to
F. pellucida, O. fusiformis and O. rufescens might be explained
to some extent by the Tsushima Current (Figure 3). Using
the same Norpac net samples as the present study, Onbe
and Ikeda indicated that two cladoceran species, Podon
schmackeri and Evadne spinifera, which occurred from June
to September, are indicative of the Tsushima Current
(Onbe and Ikeda, 1995). Terazaki also used the same
samples and reported that warm oceanic chaetognaths,
which are also indicators of the Tsushima Current,
occurred in autumn and winter (Terazaki, 1993).
The community structure of appendicularians on an
annual basis is reported from several localities within
Japanese waters (Table III). In spite of differences in
sampling depths and the mesh size of plankton nets used
in different areas, comparisons of dominant species and
their abundance are justified. The abundance of
dominant species in our results is expressed as the density
in the upper 500 m. It is reasonable that the abundance
should be expressed for the upper 100 m because the
distributions of dominant appendicularians were
restricted to this depth range throughout the year
(Figure 5). Their densities converted into the abundance
in the 0100 m water column is given in Table III. Oikopleura longicauda is more abundant, and O. dioica is less
abundant in Toyama Bay than in Omura Bay, Tokyo Bay,
or Fukuyama Harbour. In general, the appendicularian

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F. pellucida
O. fusiformis

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Table III: Comparison of appendicularian communities in several waters around Japan

Region

Toyama Bay

Suruga Bay

Bottom

Sampling

Mesh size Number of

Dominant

Maximum abundance Reference

depth (m)

depth (m)

(m)

species

species

(ind. m3)

1000

0500

100

21

O. longicauda

1650a

F. borealis f. typica

1005 a

O. dioica

248 a

5002000

0150

330

20

O. longicauda

This study

(Omori, 1977)

O. dioica
Volcano Bay

60100

0bottom

330, 100

11

(Hokkaido)

30

0bottom

330

12

(Suruga Bay)

Omura Bay

3010

0bottom

100

(Nagasaki)

1177

O. labradoriensis

216

O. longicauda

202

O. longicauda

83

O. dioica

64

O. fusiformis

38

O. dioica

7000

F. haplostoma

1200

(Shiga, 1985)

(Itoh, 1990)

(Itoh and Iizuka, 1980)

Tokyo Bay

26

0bottom

300

O. dioica

3088

(Nomura and Murano, 1992)

Fukuyama Harbour

78

0bottom

62

O. dioica

53200

(Uye and Ichino, 1995)

(Inland Sea of Japan)

a

Converted into the abundance in the 0100 m water column (see text).

fauna in the coastal waters around Japan is represented

by O. dioica, but is dominated in oceanic waters by O. longicauda, with the exception of Volcano Bay where the cold
Oyashio water has a seasonal affect. Thus, the appendicularian fauna of Toyama Bay indicates the presence of a
warm oceanic influence.
Fritillaria borealis f. typica, which is a so-called cold water
species (Tokioka, 1955), was the second most dominant
species on an annual basis in Toyama Bay (Table III) with
maximum abundance in February. The temperature
range for F. borealis f. typica was 1126C in Toyama Bay
(Table II) and 121C (optimum temperature) in Volcano
Bay (Shiga, 1985), indicating that this species can tolerate
high temperatures. Therefore, F. borealis f. typica is best
treated as a temperate species.
As mentioned above, the seasonal occurrence of
appendicularians is governed primarily by the intrusion of
the Tsushima Current into Toyama Bay. Temperature,
salinity and density, however, could not account for the
variation in appendicularian abundance. This implies
that the variation of appendicularian abundance is the
result of biological and chemical factors rather than of
physical factors. Oikopleura dioica is a good example of this.
In Toyama Bay, O. dioica was absent during the summer
when the temperature in the upper 50 m was between 19
and 28C. Itoh also found O. dioica to be absent during the
summer when the temperature was as high as 26C off

Miho Peninsula, Suruga Bay (Itoh, 1990). The decline in

the population of O. dioica in summer has also been
observed at other localities; Villefranche [>20C;
(Fenaux, 1961)], central Cantabrian coast [>16C;
(Acua and Anadn, 1992)], Tokyo Bay [no temperature
recorded; (Nomura and Murano, 1992)] and Inland Sea
of Japan [>25C; (Uye and Ichino, 1995)]. In contrast,
Fenaux found O. dioica within temperatures from 3.2 to
29.5C and considered it to be the most eurythermal
species of appendicularian (Fenaux, 1967). Furthermore,
O. dioica dominated at 2630C in tropical waters (Esnal
et al., 1985; Hopcroft and Roff, 1995). Fenaux et al. also
pointed out that O. dioica could adapt to various temperatures (Fenaux et al., 1998). Hence, the absence or
decrease in O. dioica populations cannot be explained
merely by high temperature in summer, but may in fact
suggest a biological cause.
Uye and Ichino considered that the mid-summer
decline in the population of O. dioica might be attributed
to predation by lobate ctenophores in Fukuyama Bay
(Uye and Ichino, 1995). Their data showed the appendicularian community to be composed mostly of O. dioica.
However, many appendicularian species coexisted in
summer in Toyama Bay. In the case of Toyama Bay, the
absence of O. dioica could not be attributed to selective
predation as O. dioica is similar in body to the other appendicularian species. Judging from the predominance of

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Off Miho Peninsula

F. borealis f. typica

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AC K N O W L E D G E M E N T S
We thank the captain and crew members of the RV
Tateyama-Maru, Toyama Prefectural Fisheries Experimental Station and the RV Mizuho-Maru, Japan Sea
National Fisheries Research Institute for their help in field
samplings. We also thank Dr P. Dalpadado for useful
comments on the manuscript.

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O. dioica in eutrophic coastal waters (Nomura and

Murano, 1992; Uye and Ichino, 1995), the food requirement of this species might be higher than those of other
appendicularian species. Therefore, food availability may
be one of the most critical limiting factors in the population dynamics of O. dioica. At most, Chl a concentration
was <0.5 mg m3 in Toyama Bay during summer and
autumn. It is likely that such a low food condition is not
optimal for O. dioica.
Few studies have been carried out on the seasonal variation in vertical distribution of appendicularians. In
Volcano Bay, species-specific depth distribution was
observed from spring to autumn when the thermocline
developed gradually, whereas a homogeneous distribution
was shown in winter when the water column was vertically mixed (Shiga, 1985). In the Mediterranean Sea,
oikopleurid abundance peaked between a depth of 0 and
25 m depth in summer and autumn and had an almost
homogeneous distribution within the upper 75 m in
spring and winter (Fenaux, 1968). In the central
Cantabrian Sea (Bay of Biscay), a summer temperature
gradient causes species-specific vertical distribution
patterns of appendicularians (Acua, 1994).
In contrast with these observations, most of the appendicularian species in this study had their numerical peak
in the upper 50 m of the water column both during the
stratification period ( June and September) and during the
mixing period (March and December) in Toyama Bay. For
diel vertical distribution, appendicularians did not seem
to exhibit a distinct difference in daynight distribution,
or if they did, diel vertical migration was short range
within the shallow depth. With respect to food conditions
in Toyama Bay, a Chl a maximum was consistently
observed in the top 50 m of the water column during both
the stratification period and the mixing period (Figure 5).
Vertical distribution of appendicularians corresponded to
the Chl a concentration. This strongly implies that the
vertical distribution pattern of appendicularians depends
to a large extent on food availability. The consistency with
which appendicularians remain within more preferable
food conditions throughout the day all year round may be
high in situations where food concentrations are generally
low (<1 mg m3), such as Toyama Bay.

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