Indian Journal of Geo-Marine Sciences
Vol. 45(9), September 2016, pp. 1086-1093
Biodiversity of Pelagic Copepod in the Yellow Sea and East China Sea
Hong-Ju CHEN1,2, Guang-Xing LIU1,2, *
1
Key Laboratory of Marine Environment and Ecology (Ocean University of China), Ministry of Education, Qingdao 266100, P.
R. China
2
College of Environmental Science and Engineering, Ocean University of China, Qingdao 266100, P. R. China
*[E-mail: [email protected]]
Received 27 January 2016; revised 08 March 2016
A total of 265 copepod species belong to 70 genera, 33 families and 5 orders were identified during 4 surveys in
the study area. Copepod diversity varied significantly in different seasons, and the distribution patterns were quite
different between Yellow Sea and East China Sea. Species richness, evenness and taxonomic diversity were
significantly higher in East China Sea, whereas taxonomic distinctness was significantly higher in the Yellow Sea.
Based on the present surveys combined with previous studies, the master list of pelagic copepods in the Yellow Sea
and the East China Sea were set up, and its hierarchical diversity was analyzed. Total Taxonomic Distinctness (sΔ+)
in four seasons were calculated. Funnel plots with 95% confidence limits for both Average Taxonomic Distinctness
(Δ+) and Variation in Taxonomic Distinctness (Λ+) and the ellipse plots with 95% probability contours for the joint
distribution of Δ+ and Λ+ of the pelagic copepods master list of the Yellow Sea and East China Sea were established,
and the theoretical mean value of Δ+ is 84.3.
[Keywords: Copepod, Biodiversity, Taxonomic distinctness, Yellow Sea, East China Sea]
Introduction
Zooplankton serve as secondary producers,
grazing on phytoplankton and providing food for
ichthyoplankton and carnivorous invertebrates1-2.
With short life cycles, zooplankton can respond to
and reflect the environmental condition changes
quickly; many zooplankton taxa are considered to
be indicator species whose presence or absence
may represent the influence of different water
types3. As important components of marine
zooplankton, copepods comprise the most
abundant and diverse marine communities, and
plays crucial roles in the ecosystem.
As a crucial biological parameter, biodiversity
affects the ecosystem structure and function4,
besides, the biodiversity might be an important
indicator of biological community variations in
response to hydro-climatic forcing5. The
knowledge of spatial and temporal distribution of
zooplankton biodiversity might be essential to
understand the mechanisms of which physical and
biological processes structure marine ecosystems.
The Yellow Sea and East China Sea surrounded
by China mainland, Korea Peninsula, Kyushu
Island, Ryukyu Archipelago and Taiwan Island,
are the marginal seas in the northwest Pacific
Ocean. The hydrographic conditions here are very
complex, because the presence of waters from
several origins. Generally, Coastal Water, Yellow
Sea Water, Mixed Water and Kuroshio Water are
the most fundamental waters in this area6-7, and
their properties vary seasonally with the variation
of air temperature, river discharge and wind
stress6,8. The complicated water properties are
believed to promote the prosperous biodiversity.
However, literature about zooplankton, especially
copepod biodiversity in the Yellow Sea and East
China Sea were inadequate. Studies usually
concentrated to some local area9-11, few papers
have concerned the whole shelf of the Yellow Sea
and East China Sea12.The year-round distribution
patterns of copepod biodiversity on the whole
shelf have not been clearly reported.
The aim of this work is to provide a
comparative and thorough description of the
distribution and biodiversity patterns of epipelagic
copepod of the eastern marginal seas of China.
This study will contribute to the research of
zooplankton ecology in the east China seas.
Materials and Methods
Four surveys were conducted at 37, 55, 60 and 104
CHEN et al.: BIODIVERSITY OF PELAGIC COPEPOD IN THE YELLOW SEA AND EAST CHINA SEA
stations in June-July 2006 (summer), JanuaryFebruary 2007 (winter), November 2007
(autumn), and April-May 2009 (spring),
respectively, in the Yellow Sea and East China
Sea on board R/V “Dong-Fang-Hong 2” (Fig. 1).
Zooplankton were sampled with a ring net (0.8 m
diameter, 505 μm mesh) hauled vertically from
bottom (or 200 m in deeper water) to sea surface at
a rate of 0.8-1.0 m s-1. Nets were washed after the
completion of each tow, and the samples were
preserved in 5% formalin (in seawater) for further
analyses. The filtered water volume was
determined by rope length multiplying mouth area.
N
40
In the laboratory, all copepod taxa present in the
samples were identified to species level when
possible and enumerated under a stereo
microscope (Leica S8APO). For this purpose, a
subsample was obtained from each sample with a
Folsom Plankton Splitter. Data were standardized
to abundance per 1 m3 based on the filtered water
volume determined from the rope length
multiplying mouth size.
Data on temperature, conductivity and depth of
the water were obtained using a CTD (Sea-Bird
SBE 9) profiler.
N
40
a
1087
b
Korea
35
35
China
30
30
25
25
115
N
40
120
125
E 130 115
c
N
40
35
125
E 130
120
125
E 130
d
35
30
30
25
25
115
120
120
125
E 130
115
Fig.1—Survey area and sampling stations in the Yellow Sea and East China Sea, and the horizontal distribution of surface
temperature (full line, °C) and salinity (dotted line, psu) in the study area (a. spring; b. summer; c. autumn; d. winter)
INDIAN J. MAR. SCI., VOL. 45, NO. 9 SEPTEMBER 2016
1088
Due to the characteristic of the net mesh used
(500 μm), smaller genera (e.g. Oithona and
Oncaea) and young copepodite stages were likely
to be poorly sampled13-15, and therefore, the
absolute species richness was probably
underestimated. Because samples came from the
top 200 m in deeper water, only epipelagic
zooplankton have been considered here. The
deeper community will contain other taxa which
may inhabit surface waters at seasons or times of
day that were not sampled14.
Biodiversity is a multifaceted concept which
can be considered to have three major
components: the number of taxa (richness), the
distribution of abundance within taxa (evenness)
and the relatedness of different taxa (taxonomic
distinctness)16.
Individual components of diversity were
calculated. Richness was represented by the
numbers of species. Sample evenness was
calculated using Pielou’s17 evenness J',
S
H' = －
 P log P ,
i
2 i
i 1
J' = H'/log2S,
where H' is the Shannon-Wiener diversity index,
and Pi is the proportion of the ith taxa and S is the
number of taxa in the sample.
Beside, Warwick and Clarke18 defined two
taxonomic diversity indices, capturing the
structure not only of the distribution of abundance
amongst species but also the taxonomic
relatedness of the species in each sample.
Taxonomic Diversity (Δ) includes all three aspects
of diversity. It is based on Simpson’s index with
an additional distinctness component, and it could
be thought of as the average taxonomic “distance”
between any two organisms, chosen at random
from the sample. Taxonomic Distinctness (Δ*) is
the average path length between any two randomly
chosen individuals, conditional on them being
from different species. Clarke and Warwick19-20
defined Average Taxonomic Distinctness (Δ+),
Variation in Taxonomic Distinctness (Λ+) and
Total Taxonomic Distinctness (sΔ+) for the special
case where the data consist only of
presence/absence information. Both Δ+ and Λ+ are
independent of the species number in the sample,
so they will not be influenced by sample size and
sampling efforts20. These indices were calculated
as follows:
   X X   0 X ( X  1) / 2 ,
  X X   X ( X  1) / 2
   X X ,
=
 X X
  ,
=
ij
i j
Δ=
i
i
i j
Δ*
i j
i j
Δ+
Λ+ =
sΔ+ =
i j
j
j
i
ij
i
i
j

i
ij
S ( S  1) / 2
i j
i
i
j
S ( S  1) / 2
i  j (ij   )2

i
i
ij
S 1
,
,
where Xi is the abundance of the ith species and ωij
is the distinctness weight between species i and j,
S is the number of species. For these analysis, two
species at the greatest taxonomic distance apart is
set to ω = 100. Thus, the path between species was
ωspecies = 25, ωgenera = 50, ωfamily = 75, ωorder = 100.
In the absence of a genetic phylogeny, the
published morphological and phylogenetic
literatures for copepods21-27 were used to make the
master list of pelagic copepod to calculate those
taxonomic diversity indices.
To compare the biodiversity among four
seasons, the K-dominance curve28 was plotted by
ranking the species in decreasing order of the
abundance.
Data were analyzed using the statistical package
Plymouth Routines In Multivariate Ecological
Research (PRIMER v6.1.10, PRIMER-E Ltd
2006)28.
Results
The
hydrological
characteristics
varied
substantially in different seasons (Fig. 1). Surface
temperature ranged from 8.2 °C to 26.4 °C, 17.9
°C to 29.7 °C, 14.8 °C to 26.0 °C, and 4.2 °C to
24.7 °C in spring, summer, autumn and winter,
respectively. Surface salinity ranged from 19.2 to
34.6, 21.6 to 34.2, 28.7 to 34.5, and 29.9 to 34.6 in
spring, summer, autumn and winter, respectively.
CHEN et al.: BIODIVERSITY OF PELAGIC COPEPOD IN THE YELLOW SEA AND EAST CHINA SEA
According to the morphological and
phylogenetic literature published and the present
survey data, a total of 406 copepod species
belonging to 109 genera and 44 families were
recorded in the Yellow Sea and East China Sea
(Table 1). Among these species, Calanoida species
comprised 70.7% of the total species richness,
followed by Cyclopoida species (comprised
24.4%).
Table 1—Number of each taxonomic level in the master list
of pelagic copepods in the Yellow Sea and East China Sea
Order
Family
Genus
Species
Calanoida
Cyclopoida
24
7
76
17
287
99
Harpacticoida
11
14
18
Monstrilloida
1
1
1
Mormonilloida
1
1
1
Total
44
109
406
A total of 265 copepod species belong to 70
genera, 33 families and 5 orders were identified
during the 4 surveys. Calanoida and Cyclopoida
species comprised 183 and 73 species,
respectively. In spring, 234 species belonging to
67 genera, 30 families, and 5 order were recorded,
and the number in summer, autumn and winter
were 135 species belonging to 47 genera, 25
families and 3 order, and 209 species belonging to
61 genera, 28 families and 4 order, and 166
species belonging to 55 genera, 25 families and 3
order, respectively.
The copepod species richness varied
significantly in different seasons (One-Way
ANOVA, p<0.01). As was shown, the copepod in
the Yellow Sea was less than 10 species per
station all-year-round, except in autumn, the
number in the southeast part of the Yellow Sea
reached up to 30 (Fig. 4-a). In contrast, the species
richness in the East China Sea was significantly
greater and variable, with acute seasonal variation.
The distribution patterns were similar in 4 seasons,
and the richness was higher in the offshore area
than that in the inshore. The Kuroshio Current area
possessed the highest species richness in spring
and autumn (Fig 2-a, Fig 4-a).
Similar with that of species number, the
evenness index value (J’) was higher in the East
1089
China Sea than that in the Yellow Sea. In the
Yellow Sea, the J’ value was 0.4~0.5, except that
in summer (<0.3). In the East China Sea, a similar
pattern with that of species number was detected.
It was notable that, the lowest J’ value (0.04) was
found in the adjacent water off the Changjiang
River Estuary in spring, as a result of the dominant
species Calanus sinicus “bloom” (14523.0
ind/m3).
In like manner, the distribution pattern of Δ was
similar with S and J’. The values were much
higher in the East China Sea than that in the
Yellow Sea (Fig. 2~5, c). In summer, the Δ value
in the Yellow Sea got the rock bottom (Fig. 3-c).
Inconsistently, the Δ* showed a diametrically
opposite distribution pattern. The values were
higher in the Yellow Sea than that in the East
China Sea (Fig. 2~5, d).
According to the summarized master list of the
pelagic copepods, we generated the 95%
confidence funnel for Δ+ and Λ+, and the
theoretical values were 84.3 and 256.9,
respectively. Both the two indices showed no
dependence of its mean value on the sample size
(except for very small samples) (Fig. 6). The
values of sΔ+ were 1966, 1137, 1772 and 1388 in
spring, summer, autumn and winter, respectively.
The K-dominance curves showed the
cumulative abundance (percentage) against species
rank (logarithmic) in decreasing order of the
abundance in four seasons (Fig. 7). The curves in
autumn and winter with low initial dominance
reached the top slow indicated homogeneous
abundance and high diversity in these seasons.
Whereas the curve in spring with a very high
initial dominance, which show the low evenness
despite the highest species richness.
Discussion
Compared with the other large-scaled surveys
conducted in the world, for example, the Scotia
Sea (67 species)29, the north Atlantic coast of
southern Morocco (78 species)30 and the
southwestern Atlantic Ocean (113 species)31, the
pelagic copepod species richness in the Yellow sea
and East China Sea were higher. Chen26 indicated
that, most of the pelagic copepod species in China
seas belong to warm-water species and tropic
INDIAN J. MAR. SCI., VOL. 45, NO. 9 SEPTEMBER 2016
1090
species. Zheng et al.32 reported that, the Kuroshio
Current played a controller role in the horizontal
distribution of planktonic copepods in China seas.
Zooplankton communities in the Kuroshio area
were characterized by high species richness and
low abundance10-11. As was shown in the present
study, the horizontal distribution of pelagic
copepod diversity showed a remarkable decrease
gradient from the south to north and from the off
shore to inshore area, which implied the weaken of
the Kuroshio influence. The semi-closed Yellow
N
40
N
a
40
Sea was not evidently affected by the Kuroshio,
and the biodiversity showed no acute variation
year round (Fig.2-Fig.5). The Kuroshio water was
originated from the North Equatorial Current in
the tropic Pacific Ocean. The thermal structure in
the tropics and subtropics water provided greater
number of niches for the organisms than that in
they other sea areas14, simultaneously, the great
stability of the environment allow the organisms
more ecologically
N
b
40
N
c
40
35
35
35
35
30
30
30
30
25
25
115
120
125
25
E130 115
120
125
d
25
E130 115
120
125
E130 115
120
125
E130
Fig.2—Horizontal distribution of pelagic copepods diversity indices in spring (a, species number; b, evenness index; c,
taxonomic diversity index; d, taxonomic distinctness index)
N
40
N
a
40
N
b
40
N
c
40
35
35
35
35
30
30
30
30
25
25
115
120
125
25
E130 115
120
125
d
25
E130 115
120
125
E130 115
120
125
E130
Fig.3—Horizontal distribution of pelagic copepods diversity indices in summer (a, species number; b, evenness index; c,
taxonomic diversity index; d, taxonomic distinctness index)
N
40
N
a
40
N
b
40
N
c
40
35
35
35
35
30
30
30
30
25
115
25
120
125
E130 115
25
120
125
E130 115
d
25
120
125
E130 115
120
125
E130
Fig.4—Horizontal distribution of pelagic copepods diversity indices in autumn (a, species number; b, evenness index; c,
taxonomic diversity index; d, taxonomic distinctness index)
CHEN et al.: BIODIVERSITY OF PELAGIC COPEPOD IN THE YELLOW SEA AND EAST CHINA SEA
N
40
N
a
40
N
b
40
N
c
40
35
35
35
35
30
30
30
30
25
25
120
115
125
E130 115
25
120
125
1091
d
25
E130 115
120
125
E130 115
120
125
E130
Fig.5—Horizontal distribution of pelagic copepods diversity indices in winter (a, species number; b, evenness index; c,
taxonomic diversity index; d, taxonomic distinctness index)
Fig.6—Measured Δ+ and Λ+ of pelagic copepods in four cruises, plotted against the 95% confidence funnel in the Yellow Sea and
East China Sea.
Cumulative Dominance %
100
spring
summer
autumn
winter
80
60
40
20
0
1
10
100
1000
Species rank
Fig.7—K-dominance curve for copepod abundance in four
seasons.
specialization14,33. And, accompany with the stable
primary productivity, the zooplankton biodiversity
of the Kuroshio keeps in a steady high position.
The study of the influence of Kuroshio Current
and its branches (i.e., the Taiwan Warm Current
and the Yellow Sea Warm Current) on the
zooplankton distribution was always an important
component of the biodiversity study contents in
the China seas34,35.
The distribution of species number, evenness
index and taxonomic diversity showed similar
patterns, decreasing from the south to the north
and from the off shore to the inshore area. WooddWalker et al.14 indicated that, the taxonomic
diversity of pelagic copepod in the sea surface
water with warm temperature in the low latitude
area was high and stable. In the Kuroshio area, the
results support Woodd-Walker’s hypothesis. The
taxonomic diversity included the aspects of
species richness and evenness, so the distribution
pattern was greatly affected by these two indices.
The distribution pattern of taxonomic distinctness
was significantly different from the former 3
indices; the highest value was always located in
the middle part of the south Yellow Sea, and much
1092
INDIAN J. MAR. SCI., VOL. 45, NO. 9 SEPTEMBER 2016
higher than that in the East China Sea (Fig.2~Fig.5
d). Δ* only showed the average taxonomic
discrepancy in the community, the correlativity
with the other indices was not significant. Totally,
no more than 30 species of the pelagic copepod
were recorded in each season in the Yellow Sea.
The relatively rich in the high taxonomic
categories (eg., Family) and limited in the low
taxonomic categories (eg., Species) caused the
paradox of the high taxonomic distinctness and
low species richness in the Yellow Sea. The
increase in taxonomic distinctness of pelagic
copepod from the subtropical waters to the
subantarctic waters in the south-western Atlantic
shown by Berasategui et al.15 was consistent with
our data. The measures of Δ and Δ* are relatively
insensitive to disparities in sampling effort19, so,
these
indices
might
provide
more
intuitive/comprehensive measures of biodiversity
than conventional indices15.
According to the established master list of the
pelagic copepods of the Yellow Sea and East
China Sea, the species number was quite high
(406). As a result of the limitation of survey area
coverage, time, net mesh selected and the
literatures referred, the leave out of few species
was unavoidable. Whereas we all the same
consider this master list established was roundly
imaged the species composition of copepods in the
Yellow Sea and East China Sea. The hierarchical
diversities were not sensitive but robust, the
addition of a small number of newly recorded
species to the inventory was unlikely to have a
detectable effect on the overall mean and
confidence funnels20. So the present study
provided a reasonable and concernful result for the
background materials of biodiversity research in
east China seas.
At the level of both seasons and stations, the Δ+
values showed no significant departure from the
expected value, but the Λ+ values were
significantly higher than the “mean” value (Fig.6),
especially the departure of stations in the East
China Sea were more distinct. In the East China
Sea, the species richness was high, the species list
in which there were several different families
represented only by a single species, and also
some families which are species-rich, that pricked
up the variation of taxonomic distinctness.
According to Clarke and Warwick20, an expected
Δ+ value with a greater than expected Λ+ may be
characteristic of an island fauna, and the elevated
Λ+ is due to a reduction in habitat diversity.
Obviously, it was not in reason to explain the
present result with this conclusion. Clarke and
Warwick’s deduction was based on benthic fauna,
such as free-living nematodes, or other small
polychaete taxa, which lived a settled life. Pelagic
copepod drifted along with the water, and the
appearance and distribution was driven by the
current. The great disparity of living environment
and life style between the pelagic and benthic
organisms, might affect the reflecting of
biodiversity indices on the ecological significance.
Acknowledgement
This study was supported by National Key
Basic Research Project (2014CB953701) and
National Natural Science Foundation of China
(41210008). Authors also acknowledge Mr. DongHui XU, You-Song HUANG, Jian LIN and XiaoFeng CHEN for helping with the sampling work
on board.
References
1
2
3
4
5
6
Mackas, D.L. and Tsuda, A., Mesozooplankton in the
eastern and western subarctic Pacific: community
tructure, seasonal life histories, and interannual
variability, Prog. Oceanogr., 43 (1999) 335-363.
McKinnon, A.D., Duggan, S. and Déath, G.,
Mesozooplankton dynamics in nearshore waters of the
Great Barrier Reef, Estuar. Coast. Shelf S., 63 (2005)
497-511.
Mackas, D.L., Thomson, R.R. and Galbraith, M.,
Changes in the zooplankton community of the British
Columbia continental margin, 1985-1999, and their
covariation with oceanographic conditions, Can. J. Fish.
Aqua. Sc., 58 (2001) 685-702.
Hooper, D.U., Chapin, IIIFS., Ewel, J.J., Hector, A.,
Inchausti, P., Lavorel, S., Lawton, J.H., Lodge, D.M.,
Loreau, M., Naeem, S., Schmid, B., Setälä, H.,
Symstad, A.J., Vandermeer, J. and Wardle, D.A.,
Effects of biodiversity on ecosystem functioning: a
consensus of current knowledge, Ecol. Monogr., 75
(2005) 3-35.
Beaugrand, G. and Ibanez, F., Monitoring marine
plankton ecosystems. II: Long-term changes in North
Sea calanoid copepods in relation to hydro-climate
variability, Mar. Ecol. Prog. Ser., 284 (2004) 35-47.
Su, Y.S., A survey of geographical environment
circulation system in the Huanghai Sea and East China
CHEN et al.: BIODIVERSITY OF PELAGIC COPEPOD IN THE YELLOW SEA AND EAST CHINA SEA
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
Sea, J. Ocean University of Qingdao, 19(1) (1989) 145158. (in Chinese with English summary)
Feng, S.Z., Li, F.Q. and Li, S.J., An introduction to
marine science. Higher Education Press, Beijing, Pp
470-484 (1999). (in Chinese)
Li, G.X., Han, X.B., Yue, S.H., Wen, G.Y., Yang, R.M.
and Kusky, T.M., Monthly variations of water masses in
the East China Seas, Cont. Shelf Res., 26 (2006) 19541970.
Liu, H.B., He, D.H. and Wang, C.S., Preliminary study
on distributions and communities of zooplankton in the
Kuroshio area of the middle-southern East China Sea.
In: Essays on the investigation of Kuroshio, volume 3.
Ocean Press, Beijing, Pp 305-313 (1991). (in Chinese
with English summary)
Shih, C.T. and Chiu, T.S., Copepod diversity in the water
masses of the southern East China Sea north of Taiwan,
J. Marine Syst., 15 (1998) 533-542.
Hong, X.G., Zhang, X.L., Yu, J.L. and Meng, F., Study
on biodiversity of zooplankton in the Kuroshio in the
north of the East China Sea, Acta Oceanol. Sin., 23(2)
(2001) 139-142.
Zuo, T., Wang, R., Chen, Y.Q., Gao, S.W. and Wang,
K., Autumn net copepod abundance and assemblages in
relation to water masses on the continental shelf of the
Yellow Sea and East China Sea, J. Marine Syst., 59
(2006) 159-172.
Gallienne, C.P. and Robins, D.B., Is Oithona the most
important copepod in the world’s oceans?, J. Plankton
Res., 23 (2001) 1421-1432.
Woodd-Walker, R.S., Ward, P. and Clarke, A., Largescale patterns in diversity and community structure of
surface water copepods from the Atlantic Ocean, Mar.
Ecol. Prog. Ser., 236 (2002) 189-203.
Berasategui, A.D., Ramĭrez, F.C. and Schiariti, A.,
Patterns in diversity and community structure of
epipelagic copepods from the Brazil-Malvinas
Confluence area, south-western Atlantic, J. Marine
Syst., 56 (2005) 309-316.
Purvis, A. and Hector, A., Getting the measure of
biodiversity, Nature, 405 (2000) 212-219.
Pielou, E.C., An introduction to mathematical ecology.
New York: Wiley-Interscience, 286 p. (1969).
Warwick, R.M. and Clarke, K.R., New ‘biodiversity’
measures reveal a decrease in taxonomic distinctness
with increasing stress, Mar. Ecol. Prog. Ser., 129 (1995)
301-305.
Clarke, K.R. and Warwick, R.M., A taxonomic
distinctness index and its statistical properties, J. Appl.
Ecol., 35 (1998) 523-531.
Clarke, K.R. and Warwick, R.M., A further biodiversity
index applicable to species list: variation in taxonomic
distinctness, Mar. Ecol. Prog. Ser., 216 (2001) 265-278.
Chen, Q.C. and Zhang, S.Z., The planktonic copepods of
22
23
24
25
26
27
28
29
30
31
32
33
34
35
1093
the Yellow Sea and the East China Sea. I. Calanoida,
Studia Marina Sinica, 7(1965) 20-131. (in Chinese)
Chen, Q.C. and Shen, J.R., The pelagic copepods of the
South China Sea. II, Studia Marina Sinica, 9 (1974)
125-137. (in Chinese with English summary)
Chen, Q.C. and Zhang, S.Z., The pelagic copepods of the
South China Sea. I, Studia Marina Sinica, 9 (1974) 101116. (in Chinese with English summary)
Chen, Q.C., Zhang, S.Z. and Zhu, C.S., The planktonic
copepods of the Yellow Sea and the East China Sea. II.
Cyclopoida and Harpacticoida, Studia Marina Sinica, 9
(1974) 27-76. (in Chinese with English summary)
Chen, Q.C., The pelagic copepods of the South China
Sea III. Contributions on Marine Biological Research of
the South China Sea 1. Marine Press, Beijing, pp 133138 (1983). (in Chinese)
Chen, B.Y., A preliminary study on fauna of planktonic
copepods in the China seas, Acta Oceanol. Sin., 5(1)
(1986) 118-125.
Huang, Z.G., Marine species and their distributions in
China seas. Ocean Press, Beijing, Pp 496-516 (1994).
(in Chinese)
Clarke, K.R. and Warwick, R.M., Change in Marine
Communities: An Approach to Statistical Analysis and
Interpretation, second ed. Primer-E Ltd. 172p. (2001).
Ward, P., Grant, S., Brandon, M., Siegel, V., Sushin, V.,
Loeb, V. and Griffiths, H., Mesozooplankton
community structure in the Scotia Sea during the
CCAMLR 2000 survey: January-February 2000, DeepSea Res. II, 51 (2004) 1351-1367.
Somoue, L., Elkhiati, N., Ramdani, M., Lam Hoai, T.,
Ettahiti, O., Berraho, A. and Do Chi, T., Abundance and
structure of copepod communities along the Atlantic
coast of southern Morocco, Acta Adriatica, 46(1) (2005)
63-76.
Berasategui, A.D., Menu, M. S., Gómez-Erache, M.,
Ramírez, F.C., Mianzan, H.W. and Acha, E.M.,
Copepod assemblages in a highly complex hydrographic
region, Estuar. Coast. Shelf S., 66 (2006) 483-492.
Zheng, Z., Li, S. and Li, S.J., Species composition and
geographical distribution of plankton copepods in China
seas, J. of Xiamen University (Natural Science), 18(2)
(1978) 51-63. (in Chinese)
Valentine, J.W., Evolutionary paleoecology of the
marine biosphere. Prentice-Hall, Englewood Cliffs, NJ.
511p (1973)
Xu, Z.L., Cui, X.S. and Chen, W.Z., Species
composition and dominant species on pelagic copepods
in the East China Sea, J. of Fisheries of China, 28(1)
(2003) 35-40. (in Chinese with English summary)
Xu, Z.L., The past and the future of zooplankton
diversity studies in China seas, Biodiversity Science,
19(6) (2011) 635-645. (in Chinese with English
summary)