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Seasonal population dynamics and

production of Microsetella norvegica, a
widely distributed but little-studied
marine planktonic harpacticoid copepod
S. UYE, I. AOTO AND T. ONB
FACULTY OF APPLIED BIOLOGICAL SCIENCE, HIROSHIMA UNIVERSITY,
CORRESPONDING AUTHOR: E-MAIL:

- KAGAMIYAMA CHOME, HIGASHI-HIROSHIMA -, JAPAN

suye@hiroshima-u.ac.jp

Microsetella norvegica is a widely distributed marine planktonic harpacticoid copepod, which is poorly
known from the biological point of view. We investigated the seasonal population dynamics and
production of M. norvegica in the central part of the Inland Sea of Japan. It occurred throughout
the year, whilst its reproduction was confined to the warm season between May and November. The
proportion of ovigerous females, which carry a single egg sac, was low (mean: 23.1%) in August
and September, and high (53.6%) in October. Their brood size attained a maximum (mean: 15.8
eggs per sac) in July and August and declined gradually to 6.2 eggs in November. Duration time
from egg laying to moulting to adulthood was temperature-dependent; it was 31.9 and 14.3 days at
20 and 27C, respectively, under excess food conditions in the laboratory. An enormously high population abundance (7.32  104 individuals m3), which accounted for 86.5% of the total copepods, and biomass (69.6 mg C m3) gave an annual maximum production rate of 4.90 mg C m3
day1) in October. Nauplii and copepodites disappeared in December, and the overwintering population was represented by adults, mainly large females. Associations of M. norvegica with marine
snow aggregates, which have often been found in oligotrophic waters, were not observed in the foodrich environment of the Inland Sea of Japan.

I N T RO D U C T I O N
Harpacticoid copepods are primarily benthic. A very
small proportion (0.5%) of harpacticoid species inhabit
the pelagic ocean permanently (Boxshall, 1979; Huys and
Bttger-Schnack, 1994). These pelagic harpacticoids
show relatively active swimming ability (e.g. Euterpina acutifrons), unique structural features, such as an elongated
worm-like body and caudal setae that slow their sinking
velocity (e.g. Microsetella spp. and Macrosetella spp.), or close
association with floating substrates, namely the colonial
cyanobacterium Trichodesmium [e.g. Macrosetella gracilis;
(Calef and Grice, 1966; Tokioka and Bieri, 1966; ONeil,
1998)] or suspended organic aggregates [e.g. Microsetella
norvegica; (Alldredge, 1972; Ohtsuka et al., 1993; Green
and Dagg, 1997)].
Microsetella norvegica is distributed widely in subtropical
(Gonzalez and Bowman, 1965; Goswami, 1979), temper-

Oxford University Press 2002

ate (Mori, 1937; Boxshall, 1979; Bradford-Grive et al.,

1999) and boreal (Sars, 1911; Davis, 1949) waters of the
world oceans. This small species (usually < 0.5 mm in
body length) often represents one of the numerically
dominant copepods in coastal waters (Fish, 1955; Yamazi,
1956; Anraku, 1975; Hirota and Hara, 1975; Dugas and
Koslow, 1984). Compared to the vast information available on the population dynamics and production of
marine planktonic calanoids, few studies have been
focused on marine planktonic non-calanoids (Paffenhfer,
1993; Sabatini and Kirboe, 1994; Uye and Sano, 1995,
1998). The only exception among the planktonic
harpacticoids is represented by Euterpina acutifrons, which
ubiquitously occurs in tropical, subtropical and temperate
coastal waters and frequently dominates the local mesozooplankton community (Haq, 1972; Zurlini et al., 1978;
DApolito and Stancyk, 1979; Carlotti and Sciandra,
1989). Very few studies have centered on M. norvegica.

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In the Inland Sea of Japan, M. norvegica occurs in the

plankton throughout the year. Hirota investigated the seasonal occurrence of copepods in Hiuchi Nada, a central
part of the Inland Sea of Japan, and found that
M.norvegica was one of four numerically dominant species,
the others being Oithona davisae, O. similis and Paracalanus
sp., and was particularly important in summer and
autumn, comprising > 50% of the total copepod abundance (Hirota, 1964). However, even in the central part of
the Inland Sea of Japan, this species was very scarce in
less-saline Fukuyama Harbour, where salinity was often
diluted < 30 due to freshwater runoff [(Uye and Liang,
1998), and unpublished results]. Stenohaline nature of
M.norvegica, compared to that by Acartia omorii, O. davisae
and Paracalanus sp., was also reported by Yamazi (Yamazi,
1956), who found that this species occurred primarily near
the mouth, rather than near the head, of inlets and bays.
The annual average carbon biomass of the M.norvegica
population over the entire Inland Sea of Japan was 1.24
g C m3, which ranked fourth after Paracalanus sp.,
Calanus sinicus and Oithona spp. biomass (Uye and Shimazu,
1997).
The aim of this study is to fill gaps in our knowledge of
the population dynamics and production of M. norvegica.
We conducted weekly samplings for a year to reveal how

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the population varies seasonally. In addition, rearing

experiments were conducted from egg to adulthood under
controlled laboratory conditions. These field and laboratory results were combined to estimate the population production rate, and the trophodynamic role of this species
in the central part of the Inland Sea of Japan was assessed.

METHOD
Sampling and sample analysis
Weekly samplings of mesozooplankton were conducted
during daytime from 22 December, 1982 to 24 December,
1983, from a pontoon in Numakuma, Hiroshima Prefecture (St. A, Figure 1). Copepodites and adults of M.
norvegica were collected by towing a 94 m mesh plankton
net (with a flowmeter at the mouth) vertically from the
bottom to the surface (through a depth of 810 m). Its
nauplii were sampled with a Van Dorn bottle (6 l) at each
1 m depth (total water volume: 108132 l) and filtered
through a 40 m mesh hand net. Neutralized formalin
was immediately added to the samples (final concentration: 510%). Additional net samplings were made at
an offshore station (St. B, depth: 2022 m, Figure 1), ca.
10 km southeast of St. A, from 18 May to 13 October,

Fig. 1. Map showing the location of the sampling stations (St. A and St. B) in the central part of the Inland Sea of Japan.

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1983, in order to supplement data on the occurrence of

adult females and their fecundity.
Microsetella norvegica individuals were staged, sexed and
counted (all naupliar stages were counted collectively)
under a dissecting microscope. Egg sacs, most of which
were detached from the female genital segment in formalin-preserved samples, were also counted. The body
length of each developmental stage (Figure 2) was
measured to the nearest 5 m for at least 10 individuals
using an eyepiece micrometer. Usually 50 egg sacs were
picked up from each sample, and dissected with a needle
to count the number of eggs per sac.

Laboratory rearing experiment

The stock cultures were derived from adult females and
males collected at St. A and maintained in a temperature
controlled room (24 1C; with a light periodicity of 12
h light : 12 h dark). In the preliminary culture experiments, we tested eight phytoplankton species (namely
chlorophyte: Dunaliella tertiolecta; chrysophyte: Isochrysis
galbana; diatoms: Chaetoceros sp., Ditylum brightwelli, Skeletonema costatum, Thallasiossira weissflogii; dinoflagellate:
Heterocapsa sp.; and raphidophyte: Chattonella marina), and
found that I. galbana and Chaetoceros sp. yielded the highest
population increase in the cultured copepods. Hence, a
mixture of cultured I. galbana and Chaetoceros sp. (cell diameter: 45 m for both species) was given as standard food
(cell concentration of each species 1.21.7  105 cells
ml1, total carbon concentration ca. 10001400 g l1).
The surface of the medium in culture vials (i.e. 500 or
1000 ml glass bottles) was covered with a watchglass to
remove air space and the vials were placed on a rotary
shaker (Taiyo Co., 30 r.p.m. of horizontal rotation) to give
water turbulence. Half of the medium was replaced with
freshly prepared medium at 3-day intervals.
Ovigerous females from the stock culture were introduced individually into glass crystallizing dishes containing ca. 60 ml of rearing medium and were kept in a water
bath establishing four different levels of temperature
(17.1, 20.7, 24.1 and 26.7C). At 24 h intervals, egg
hatching and production of new egg sacs were monitored
for 14 females at each temperature. Fully matured egg sacs
were removed from the females and incubated individually in glassfibre-filtered (GF/C) sea water to measure the
duration from hatching to development into naupliar
stage (N) II, a first feeding stage.
Batches of ca. 200300 NI obtained from the stock
culture were reared in 500 ml glass bottles at 20.7, 24.1
and 26.7C. The stage composition was monitored daily
by subsampling 2030 specimens. Naupliar stage was
identified for formalin-preserved subsamples under a
binocular microscope. Copepodite stages were identified
in live subsamples, which were returned to the bottles after

inspection. The time of entering each stage was defined

by the time when 50% of specimens had moulted into that
stage [see (Uye, 1980)].
Live specimens of each developmental stage were subsampled from the stock culture or were freshly caught
animals at St. A and their carbon contents were measured
using a total carbon analyser (Oceanographic International Co., Model-524).

Growth rate and production rate

Combining the data from the laboratory experiment and
field surveys enabled the determination of the specific
growth rate and population production rate of M. norvegica
at St. A. Assuming that there was no food limitation, the
average specific growth rates over naupliar and copepodite stages (gN and gC day1) were calculated from:
gN = (lnCCI lnCE)/DN

(1)

gC = (lnCCVIF lnCCI)/DC

(2)

where CE, CCI and CCVIF are carbon weights (g) of an egg,
copepodite stage I (CI) and adult female, respectively, and
DN and DC are duration (days) of the naupliar and copepodite stages, respectively.
Since adult males did not grow beyond copepodite
stage V (CV), their net production was assumed to be negligible. We also assumed that the net production of adult
females was attained only by egg production. Hence, the
specific growth rate of breeding females (gCVIF, day1) was
given by:
gCVIF = {ln[CCVIF + (CE  BS)] lnCCVIF}/(DN + DI) (3)
where BS is brood size (eggs per sac) and DI is the duration
(days) from the hatching of an egg sac to the production
of a new one.
The population production rate (P, mg C m3 day1)
was given by:
P = (gN  BN) + (gC  BC) + (gCVIF  BCVIF)  OF

(4)

where BN, BC and BCVIF are biomass (mg C m3) of

nauplii, copepodites and females, respectively, and OF is
the proportion of total females that are ovigerous.

R E S U LT S
Seasonal environmental variables
Due to a relatively strong tidal current at St. A, vertical
differences in temperature and salinity between the
surface and bottom were always less than 1.0C and

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Fig. 2. Body length (BL) measurements applied to nauplius and copepodite of Microsetella norvegica.

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0.4 psu, respectively. Average temperature of the water

column varied from 9.1C in February to 27.4C in
August. Average salinity ranged from 30.9 to 32.9 psu.
At the offshore station, St. B, water column stratification was more prominent (maximum vertical differences: 4.0C and 2.5 psu), from May to September.
However, there was no difference in the surface temperature and salinity from those at St. A on each sampling
date.

Seasonal variation in abundance

The population of M. norvegica occurred almost continuously throughout the year, but the occurrence of the larval
stages was confined to warm seasons (Figure 3). The
nauplii did not occur until May, and they increased gradually thereafter to form a peak (2.50  104 individuals m3)
in early October. All copepodite stages appeared almost
simultaneously in July, and increased to a maximum (7.32
 104 individuals m3) in early October. After their disappearance from the samples in late June and early July,

Fig. 3. Seasonal variation in abundance of different developmental

stages of Microsetella norvegica at St. A from December 1982 to December 1983. Shaded area indicates the abundance of ovigerous females.

adults were resurgent and increased to a peak (6.25  104

individuals m3) also in early October. The larval stages
almost disappeared by the end of December. The adults
overwintered and remained in the plankton until the
resurgence of the next generation in May. The occurrence
of ovigerous females was confined to the period from
August to November. In early October, M. norvegica monospecifically dominated, accounting for 86.5% of the total
copepod (copepodites and adults) abundance.
At St. B, although adult females became very scarce
(<5 individuals m3) in June, they never disappeared from
the plankton. The variation pattern of adults was similar
to that at St. A, but their peak (8.3  104 individuals m3
in late September) was much lower than that at St. A.

Seasonal variation in body length

Mean body lengths (BL) of copepodites and adult males
were shortest in summer and slightly increased in the
autumn, but this seasonal pattern was not apparent for adult
females. The lengths of adult females and males increased
gradually during overwintering (Figure 4). The relationship
between BL and water column average temperature at the
time of sampling (T ) was examined for each stage, and a
significant (P < 0.05) negative correlation was obtained
excluding adult females (annual mean BL 639 m):
CI: BL = 4202.61T (r = 0.637)

(5)

CII: BL = 4913.74T (r = 0.711)

(6)

CIII: BL = 4952.57T (r = 0.534)

(7)

Fig. 4. Seasonal variation in body length of Microsetella norvegica at

St. A.

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CIV: BL = 5854.39T (r = 0.717)

(8)

CV: BL = 6655.64T (r = 0.697)

(9)

CVIM: BL = 6043.49T (r = 0.776)

(10)

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The eggs of M. norvegica were spherical or slightly ovoid.

There was no seasonal variation in egg size; the mean
equivalent spherical diameter was 63.2 m and egg
carbon weight was 0.025 g.

Development rate and body length : carbon

weight relationship

Seasonal variation in fecundity

Adult sex composition skewed towards male domination
during early reproductive period (e.g. July and August),
and was almost even during the peak reproductive period
(e.g. September and October). Thereafter, females usually
outnumbered males (Figure 5).
During the breeding period, the mean composition of
ovigerous females (OF ) was 23.1% in August and September, increased to 53.6% in October, and declined in
November to < 2.8%.
Mean brood size (BS ) was largest (15.8 eggs per sac) in
August and early September at St. A, and gradually
decreased to 6.2 eggs per sac toward the end of the breeding season (Figure 8). At St. B, the spawning commenced
in May and the brood size attained near-maximum level
(15.4 eggs per sac) in July (Figure 6). After the midsummer peak (16.3 eggs per sac), it declined in a similar
manner observed at St. A, but reproduction had ceased by
the end of October at St. B.

The time from hatching of an egg sac to the bearing of the

next (DI) was independent of temperature, with a mean of
0.39 days (SD = 0.17 day, n = 40). However, the mean
times for eggs to develop into NI were 5.7, 3.1, 2.2 and 1.6
days at 17.1, 20.7, 24.1 and 26.7C, respectively (Figure
7). The best fit Blehrdeks function [see (Corkett and
McLaren, 1970)] to describe the relationship between
temperature (T, C) and the duration of egg stage (DE,
days) was:
DE = 269(T 7.9) 1.75 (r = 0.999)

(11)

The duration of naupliar and copepodite stages also

increased exponentially with the decrease of temperature
(Figure 7). Assuming the equiproportional development,
where the curvature of the regression line is uniform for
any life stage (Corkett, 1984), the Blehrdek functions to
describe the naupliar (DN, days) and copepodite (DC, days)
durations were:
DN = 1034(T 7.9)1.75 (r = 0.999)

(12)

DC = 1198(T 7.9)1.75 (r = 0.941)

(13)

It took about a month for M. norvegica to develop from egg

to adult at 20C and half this time at 27C. Stage duration was shortest in NI, roughly the same from NII to CII,

Fig. 5. Seasonal variation in sex composition of adult Microsetella

norvegica at St. A.

Fig. 6. Seasonal variation in brood size of Microsetella norvegica at St. A

() and St. B (). Vertical lines denote SD.

Fig. 7. Relationship between temperature (T, C) and duration (D,

days) of egg (E), naupliar stages (N) and copepodite stages (C) of
Microsetella norvegica.

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Table I: Relative duration of each developmental stage in total duration from egg to adulthood of
Microsetella norvegica at three different temperatures
Temp.

Developmental stage

(C)

Egg

NI

NII

NIII

NIV

NV

NVI

CI

CII

CIII

CIV

CV

20.7

10.5

3.5

9.6

6.3

4.9

5.1

10.9

5.3

5.9

12.1

10.3

15.6

24.1

11.5

3.4

6.3

8.5

8.1

6.7

6.5

4.6

7.5

9.6

11.9

15.4

26.7

10.3

3.1

5.7

7.4

7.1

4.5

12.2

6.5

6.5

13.9

11.4

11.4

Mean

10.8

3.3

7.2

7.4

6.7

5.4

9.9

5.5

6.6

11.9

11.2

14.1

and was comparatively longer in CIII, CIV and CV

(Table I).
Body carbon weight (C, g) increased exponentially
with the increase of body length (BL, m), expressed by a
single regression equation for entire life stages (Figure 8):
C = 2.65  106  BL1.95 (r = 0.987)

(14)

Seasonal variation in population biomass

and production rate
The biomass of the M. norvegica population was higher
than 1.0 mg C m3 only during the period from September to December, with an enormous peak (69.6 mg C m3)
in early October (Figure 9). The mean contribution of
nauplii to the population biomass was only 8% during this
culminating period.

Fig. 9. Seasonal variation in biomass of Microsetella norvegica population

at St. A.

The specific growth rates ( g) of nauplii, copepodites

and breeding females were calculated on each sampling
date (Figure 10). These rates were temperature-dependent, as shown in Figure 11, since the stage or inter-clutch
duration was also temperature-dependent (Figure 7). To
express the relation to temperature, a linear regression
was applied to nauplii:
gN = 0.023T 0.276 (r = 0.993)

(15)

since the correlation coefficient was slightly higher than

that of the non-linear form (r = 0.976). For copepodites

Fig. 8. Relationship between body length (BL, m) and carbon weight

(C, g) of Microsetella norvegica.

Fig. 10. Seasonal variation in specific growth rate of nauplii, copepodites and adult females of Microsetella norvegica at St. A.

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DISCUSSION

Fig. 11. Relationship between the specific growth rate ( g, day1) of

nauplii, copepodites and adult females of Microsetella norvegica and temperature (T, C).

and adult females, the following non-linear functions were

given, respectively:
gC = 0.0016e0.178T (r = 0.987)

(16)

gCVIF = 0.0023e0.176T (r = 0.972)

(17)

The production rate of the M. norvegica population was

significant only in September, October and November
(Figure 12). The highest production rate (4.90 mg C m3
day1) was recorded in early October. The contribution of
copepodites (52.0%) was largest to the production during
this period, followed by adult females (30.2%) and nauplii
(17.8%).

Fig. 12. Temporal variation in production rate of Microsetella norvegica

population at St. A from September to November, 1983.

One of the successful strategies employed by pelagic

harpacticoids is to develop associations with floating substrates. It has been documented that M. norvegica uses them
for buoyant support and/or food source (Alldredge, 1972;
Ohtsuka et al., 1993; Green and Dagg, 1997). However,
this species may never use these substrates, at least in the
central part of the Inland Sea of Japan. Microsetella
norvegica was neither observed attached to macroaggregates in the contents of the large-volume (2 l), clear
Plexiglas cod-end of plankton nets after they were towed
gently (present study), nor on discarded larvacean houses
collected by SCUBA diving (Ohtsuka et al., 1993).
Ohtsuka et al. (Ohtsuka et al., 1993) found that there was
no incidence of attachment (out of 276 houses examined)
in the eutrophic Inland Sea of Japan (near our St. B, mean
surface chlorophyll a concentration was : 3.14 g l1),
whereas the attachment was observed (five out of 272
houses) in an oligotrophic inlet (Honmura Inlet,
Kagoshima, mean surface chlorophyll a concentration:
0.74 g l1). A similar offshore increase in attachment was
described by Green and Dagg (Green and Dagg, 1997)
along an inshoreoffshore transect in the northern Gulf of
Mexico, where eight genera of copepods were observed
on marine snow aggregates. At their most offshore station
(water column average chlorophyll a concentration: ca.
0.2 g l1), on all collected aggregates (n = 48) M. norvegica
was observed. These facts suggest that the major role of
attachment to marine snow aggregates is to obtain food in
a relatively oligotrophic environment. Marine snow particles and their associated microbial communities are
actually a significant source of nutrition. If suspended
food particles are relatively abundant, like in the central
part of the Inland Sea of Japan and in the vials of our
laboratory culture, M. norvegica does not need floating
aggregates and lives as a free swimmer.
In still water, copepodites and adults of M. norvegica
were observed to sink slowly to the bottom of the rearing
vials, with occasional upward jerks. Preliminary experiments showed that the mortality was extremely high in
still water with an open air surface, due to trapping in the
airwater interface and entangling with sedimented
phytoplankton and faecal pellets on the bottom.
However, when the open air surface was removed and
water turbulence was provided by a rotary shaker, the
mortality was significantly reduced. Its extraordinary
elongated caudal setae may easily be caught in the
surface film in small vessels, while they may help in
slowing the rate of sinking [cf. (Huys and Boxshall, 1991)]
in the water column. In the field, water turbulence may
be important for this poor swimming species. Although
the relationship between the geographical distribution

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and the water turbulence was not substantiated, M.

norvegica often aggregates in areas with relatively high
water turbulence, such as the mouths of inlets, narrow
straits and in waters around dotted islands (Yamazi, 1956;
Hirota and Hara, 1975). In more stratified parts of inlets,
the genera Acartia, Oithona and Paracalanus are more dominant (Yamazi, 1956). In this study, St. A adjoined Abuto
Strait, where the tidal current is much stronger than at
the open water St. B, where Paracalanus sp. was more
abundant than M. norvegica in the autumn (present study).
To our knowledge, no information is available about
the feeding of M. norvegica in nature. We used a mixture of
I. galbana and Chaetoceros sp. (cell diameter: 45 m) as standard food, since these two species sustained the highest
population growth of M.norvegica. Thallasiossira weissflogii
(12 m) and Skeletonema costatum (810 m) also resulted in
high population increases, but the population crashed
earlier than with I. galbana and Chaetoceros sp. Dunaliella
tertiolecta (8 m) and Heterocapsa sp. (25 m) showed inferior
food values, since the population scarcely increased. Chattonella marina (30 m) and Ditylum brightwelli (62 m) were
probably not utilized even by adult M. norvegica, since the
copepods died within 5 days in their suspension. The
above findings lead us to conclude that the major food for
M. norvegica is nano-size (i.e. 220 m) particles.
Microsetella norvegica is a typical warm-water copepod
and does not produce any resting eggs for overwintering
in the Inland Sea of Japan. A similar seasonal occurrence
was found in the North Sea off Northumberland, UK,

where the breeding was confined from spring to late

autumn and the overwintering population was composed
only of adults (Evans and Diaz, 1978; Diaz and Evans,
1983). In our study area, surviving females after overwintering commenced spawning in May, as detected at St.
B. At St. A, our data failed to show the spawning by overwintered females in early summer, whereas newly produced nauplii were found in May. This discrepancy may
be explained either by the spawning of overwintered
females that were too scarce to be collected by our samplings at St. A or the transport of nauplii from offshore by
advection. This newly formed population developed to
adulthood by mid-July, and then had matured by August.
The offspring derived from these ovigerous females
showed a remarkable increase in September. Over the
reproductive period from May to November (temperature
range: 16.827.4C), the development time from egg
laying to moulting to adulthood varied from 14.3 to 31.9
days, assuming no food limitation. Adding the time necessary for sexual maturity of young females to lay first eggs,
the generation time was slightly longer [e.g. 1520% for
Euterpina acutifrons, (Haq, 1972)]. Judging from these estimates, probably five or six generations were produced
annually in the central part of the Inland Sea of Japan,
and probably at least three generations at St. A based on
the temporal sequence of stage-specific abundance
(Figure 3).
The specific growth rates of M. norvegica are low compared to those of seven sympatric copepod species at

Table II: Comparison of the specific growth rates of Microsetella norvegica to those of sympatric
copepod species at 20C
Species

Specific growth rate (day1)

Calculated from

Nauplii

Copepodites

Females

Acartia omorii

0.42

0.37

0.65

Uye (1981), Liang and Uye (1996a)

Calanus sinicus

0.46

0.44

0.15

Uye (1988, unpublished)

Centropages abdominalis

0.34

0.45

0.65

Liang et al. (1994, 1996)

Paracalanus sp.

0.21

0.34

0.42

Uye and Shibuno (1992),

Pseudodiaptomus marinus

0.25

0.25

0.15

Liang and Uye (1997a, b)

0.21

0.21

0.28

Uye and Sano (1995, 1998)

0.22

0.27

0.28

Calanoids

Liang and Uye (1996b)

Cyclopoid
Oithona davisae
Harpacticoids
Euterpina acutifrons*

Zurlini et al. (1978),

Carlotti and Sciandra (1989)

Microsetella norvegica

0.18

0.056

0.078

* Growth parameters are determined for specimens from the Mediterranean Sea.

This study

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20C, an intermediate seasonal temperature (Table II).

Another planktonic harpacticoid, Euterpina acutifrons, also
occurs during summer and autumn in the central Inland
Sea of Japan, but at much lower densities than M. norvegica
(Hirota, 1964; Checkley et al., 1992). As no study has
measured the growth rates of E. acutifrons from our study
area, the growth data are borrowed from other studies
conducted for specimens from the Mediterranean Sea
(Zurlini et al., 1978; Carlotti and Sciandra, 1989). The
growth rates of M. norvegica are lowest among them, particularly in copepodites and females. The lower growth
rates in copepodites and females were not surprising
because of their lower feeding activity. Our laboratory
observation revealed that the nauplii frequently moved
appendages both to swim and feed, although the copepodites and adults swam intermittently and stayed calm,
without moving their feeding appendages for most of the
time.
In the central part of the Inland Sea of Japan, the water
temperature exceeds 25C in mid-summer, which is lethal
or sublethal to the dominant copepod species, such as
Acartia omorii, Calanus sinicus, Paracalanus sp., which decline
the population size. Instead, M. norvegica increases in late
summer and autumn, due to temperature-induced higher
fecundity and probably using the phytoplankton (particularly nanophytoplankton judging from our feeding experiment) food supply elevated by the autumn water mixing
(Endo, 1970). Our St. A may represent a suitable habitat
for M. norvegica, which attained a biomass as high as 69.6
mg C m3 in early October. This mono-specific biomass
was much higher than the average copepod biomass over
the entire Inland Sea of Japan in October, 1993 [16.3 mg
C m3; (Uye and Shimazu, 1997)]. Due to elevated
biomass coupled with temperature-driven higher specific
growth rates, M. norvegica plays a significant role as a
secondary producer in the Inland Sea of Japan in the late
summer to autumn.

Fish, C. J. (1955) Observations on the biology of Microsetella norvegica. Pap.

Mar. Biol. Oceanogr., Deep Sea Res., 3 (Suppl.), 242249.

AC K N O W L E D G E M E N T S

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We would like to dedicate this paper to the late Dr M. M.

Mullin of the Scripps Institute of Oceanography, for his
wonderful memory during a cruise of the joint Japan-US
Science Program in the Inland Sea of Japan (1985) and
for his valuable comments in many aspects of our plankton research.

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