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Journal of Earth Science and Engineering 2 (2012) 1-14
General Patterns of Spatial-Temporary Distribution of
the Integral Characteristics of Pelagic Macrofauna of the
North-Western Pacific and Biological Structure of Ocean
Igor V. Volvenko
Pacific Research Fisheries Center (TINRO-Center), Vladivostok 690091, Russia
Received: September 20, 2011 / Accepted: December 14, 2011 / Published: January 20, 2012.
Abstract: The integral properties of large multispecies assemblage from northwest Pacific pelagial are investigated. There are total number and overall biomass, average animal size (mean individual weight), species diversity (Shannon’s index) and its components: species richness and evenness (Pielou’s index), i.e. generalized parameters describing macrofauna as a whole. For the first time it was possible to correlate all listed integrative characteristics with each other and a location of research object in space and time, having described essence of the occurring phenomena proceeding from action of an extreme small set of general principles of organization of life in pelagic water layer. One of the consequences of the discovered regularities: the reasons for large-scale long-term changes in pelagic cenosis abundance, composition and structure are not in the rise of temperature, but in water exchange regime shift.
Key words: Pelagic macrofauna, relationships between integral characteristics, biodiversity, species richness, species evenness, abundance, animal’s size, biological structure, northwestern Pacific.
1. Introduction
According to the data of 20 thousands pelagic trawl
stations, carried out by TINRO research vessels in the
northwestern Pacific in the waters of an area of 6
mln·km2 (Fig. 1) in 1979-2005, general integral
characteristics of the macrofauna, which includes
almost 1000 species of fish and invertebrate with the
body size ≥ 1 cm (Table 1), have been explored. For
each station, one-degree trapezoid, statistical area and
water body, for different ranges of depths, for the
macrofauna as a whole and separately for
macroplankton, nekton, fish, cephalopods, for different
seasons and plurannual periods the following data have
been calculated: species diversity H (Shennon’s index)
[1] and its components—species richness S and species
evenness on the abundance J (Pielou’s index) [2], total
number N and overall biomass M of all individuals, as
Corresponding author: Igor V. Volvenko, associate
professor, Ph.D., main research fields: general biology, ecology, hydrobiology, biogeography, applied statistics, data bases and GIS. E-mail: [email protected].
well as their average individual weight W [3-10]. It
turned out that with any method of sampling and
pooling of data all these characteristics are interrelated
in a special way—neither of them varies in space and
time independently of the others [11-13].
2. Methods
It is commonly known that H is expressly
determined by its two components H = J·LogS and M
= N·W, therefore, to demonstrate the discovered
regularities, it is enough to analyze the
four-dimensional space with N, W, J and S as
coordinate axes. Visually it can be represented by six
plane projections—2D-graphs or by four 3D space
projections—solids (Fig. 2).
Bivariate plots show that the population density of
animals is inversely proportional to its size or body
weight, evenness of the species distribution is in
negative correlation with abundance, while the species
richness is maximum for average size classes and
average abundance value. Therefore, triangles made of
DAVID PUBLISHING
D
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
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Fig. 1 Map of 20 thousands pelagic trawl stations, 882 one-degree trapezia, 48 standard areas of averaging of biostatistical information and the surveyed sectors of four water bodies in the northwestern Pacific region with integral characteristics of macrofauna calculated.
Table 1 Composition of the studied fauna (number of species).
Biotopic group Ecological forms Taxonomic groups
Population of the pelagic zone (814)
Nekton (780)
Vertebrates (672) Mammals, birds and reptiles (0)
Fishes and cyclostomata (672)
Invertebrate (142)
Cephalopods (71)
Crustaceans (37)
Plankton (34)
Jellyfish (24)
Ctenophora (2)
Others (8) The group “cephalopods” includes squids, cuttlefish, and pelagic octopus; the “crustaceans” are shrimp and prawn; the “others” are pteropods, nudibranchs, pyrosoms, salps, and cyclomaria. During a pelagic trawling, all these organisms are caught in a trawl with a fine mesh (10-12 mm) inserted in its end.
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
3
Eve
nnes
s
Log Number
Log
Num
ber
Log ind. Weight
Log
ind.
Wei
ght
Evenness
Log ind. Weight
Spe
cies
ric
hnes
s
Log Number
Spec
ies
rich
ness
Evenness
Spe
cies
ric
hnes
s
Fig. 2 4D virtual space (with LogN, LogW, J and S as coordinate axes) visually represented by six plane projections—2D-graphs or by four 3D space projections—solids.
segments, the length of which is inversely
proportionate to S, appear on 3D projections. The
uppermost segment turns into a point with coordinates
equal to the maximum number of species, average
evenness, average logarithms of abundance and body
weight. On the fourth projection, if we introduce an
additional S axis in an arbitrary direction, for example,
equidistant from the rest of the axes, we get the same
result (Fig. 3). This is the variables inter-correlation
structure. All these bindings are not strictly functional,
but statistical, therefore, the points, which correspond
to certain measurements, are placed not only on the
lines, but form a cloud along the segments (Fig. 4). It
eventually results in a multidimensional mount, the
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
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Fig. 3 Changes in the segment, describing the relationship of the variables in the space of LogN-J-LogW, with an increase of S. This scheme shows that all the 3D projections shown in the previous figure are of the same type in principle.
shape of which doesn’t depend on the fact, where and
when the samples were collected (Fig. 5).
Further research showed that it is typical not only for
the macrofauna of the pelagic region, but for any
assemblages of animals or plants, whether marine,
freshwater, or land [14]. However, consistent with the
topic of the report, let us confine ourselves to what it
gives for the description of the northwestern Pacific
region pelagic macrofauna.
3. Results and Discussion
In the virtual space of integral characteristics, there
are 3 extreme points A, B, C (and also B1—the same as
B, but for a single-species system, when S = 1, J = 0
and variations of N and W are maximum) (Fig. 6). In
the real space-time continuum, they correspond with
the states of biocoenotic systems, which can be
observed: A—in shore at the water area periphery, B
and C—offshore near the ocean center. The difference
between B and C is conditional on the share of
numerous small specimens of mesopelagic fauna.
This fauna is characterized by daily feeding migration
(it stays at great depth, where the sunlight does not
Fig. 4 The points corresponding to individual measurements form a cloud along the segments—here is the joint range of definition of the four variables.
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
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Fig. 5 3D distributions describing interrelation between logN, J, and S of pelagic macrofauna in northwestern Pacific for the different (A) water bodies; (B) seasons; (C) long-term periods; (D) water layers; (E) samples collected over various depths and (F) on various distances from the shoreline. Each point on the graphs corresponds to one sample (a trawl station).
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
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B
AB1
C
A
C
B
A
B
CB1
B
AA
C
Fig. 6 Four 3D projections of mutual interrelations for integral parameters of pelagic macrofauna—the same that are shown in Figs. 2 and 4. The letters A-C here mark the points which describe extreme states of biocenotic system, typical for some positions in real space and time. The first and last figures are given in section for display of their internal structure. The color intensity shows the change in diversity H (decrease from dark to light).
penetrate, in the daytime, and rises to the upper layers
in the nighttime), therefore, on the scheme, the arrow
between B-C marks the light-dark transition, i.e. either
surface-depth, or day-night.
At the ocean periphery near the continents (A) the
macrofauna abundance is maximum, but J, S and H are,
correspondingly, very low. Moving away from the
shores, the population density declines, while H
increases. Closer to the ocean center, if the samples
were collected in the light—in the daytime at shallow
depth, S is low, while J is high (B), or, where the
population density is very low, there turns out to be
only one species in the sample (B1). If the samples were
collected in the dark—at depth or near the surface at
night, S turns out to be maximum, while J and
logarithm of abundance is average (C). Points A, B, B1,
C mark the limit state of the system, while all real
intermediary situations are between them within this
figure. The transition from A to B-C corresponds to the
centripetal direction in the ocean space, which explains
the circum-continentality of the abundance, the
contradirectional circum-continentality J and H, and
extrazonality S.
In full accordance with these regularities, the widely
known, at least in Russia, concept of biological
symmetry of the oceans by Zenkevich-Bogorov [15-18]
is further proved and supplemented. According to this
concept, the life in the Ocean spreads consistent with 3
symmetry planes (Fig. 7). Most stable space
regularities correspond to three zonality types:
latitudinal, circum-continental, and vertical. Primary
type of geographic (horizontal) zonality—latitudinal
zonality—is conditioned by irregular incoming of solar
energy to the Earth. Secondary
zonality—circum-continental zonality, which appears
in aquatic Environment—is conditioned by shallow
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
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Fig. 7 Biological structure of the ocean. At the left (A) symmetry planes are shown: I—equatorial, II and III—meridional (from Ref. [18]). On the right (B) by shading density the symmetry of Pacific waters productivity is shown (from Ref. [16]).
depths and changed water circulation near the land. It is
well known that neritic and shelf regions are
characterized by higher rates of primary production,
biomasses of phyto- and zooplankton, benthos, fish and
seabirds [16, 17, 19-25].
Integral characteristics of the northwestern Pacific
pelagic macrofauna differ in the display of horizontal
zonality in the following way: N and M display
circum-continental zonality on the regional level (d in
Fig. 8), J and H display circum-continental zonality on
the global level (b in Fig. 8), W displays two opposite in
direction types of circum-continental zonality on the
global level (b and c in Fig. 8)—for various animals
and data smoothing levels, S doesn’t display any
particular zonality (neither latitudinal, nor
circum-continental). This list proves: (1) different
degree of zonality display for different characteristics;
(2) total absence of any latitudinal zonality display in
the region (a in Fig. 8)—even such well-known
generalizations as Humboldt-Wallace’s law and
a b c
d e f
Fig. 8 Examples of hierarchical levels for geographic patterns: (a) latitudinal zonality on a global level: a parameter decreases from the equator to poles following the solar energy change; (b) circumcontinental zonality on the global level: a parameter decreases from the center to the periphery of the ocean, similarly distributed species evenness and diversity; (c) the same, but a parameter decreases toward the center of the ocean; for example, the average size of the animals are distributed so; (d) circumcontinental zonality on a regional scale; example—population density; (e) the local level, where zoning is not observed; an example—diversity; (f) sublocal level, where it is difficult to distinguish or do not see any pattern; the same example.
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
8
Bergman’s rule are not valid here.
The absence of any latitudinal zonality display in the
region is conditioned by the fact, that meridional (not
latitudinal) air and water mass transport prevail here,
unlike in southern regions. Collision and interaction
between northern and southern, cold and warm waters
creates a number of local whirlpools, fluid fronts,
eddies (Fig. 9). This mixed mosaic picture is very
changeable in time and space. Apparently, this is
exactly what impedes the formation of stable
latitudinal gradient of species richness and animal size
in this region. This is where Neustruev’s provinciality
law [12, 26] proves itself. The above schemes (Fig. 2)
explain the reason for the absence of
circum-continental zonality of species richness display
in the region: S is connected with N and M so that the
species richness cannot be high either by high
population density, or by a low one, either in shore, or
far in the ocean. The average abundance is necessary
but insufficient condition for maximization of species
richness.
On the basis of “ecosystem = biocenosis+biotope”
ratio, let us proceed from biocoenotic and
biogeographic level of phenomena description to
ecosystem level. To do so, let us take notice of the fact,
what essential biotopic (abiotic) environmental factors
Fig. 9 Schemes of air and water masses moving in the surveyed region (an illustration from various papers and books).
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
9
change in space according to the circum-continental
zonality principle. It results in the following
supplement to Zenkevich-Bogorov’s concept of
biological structure of the ocean.
In the directions from the ocean center to the
periphery, the environment stability declines, while the
water exchange intensity increases, whereby the
environment ecologic capacity grows up. Accordingly,
the primary production and biogeochemical cycle
intensity as a whole increases, as well as N, M and
variations of W, numerical and weighting prevalence of
few dominant species over the others. At the same time,
H decreases.
Let us now analyze the vertical distribution of
integral characteristics (Fig. 10). Specific fauna have
formed in the deep-water biotope. This fauna includes
a great number of species, which are characterized by
medium on the logarithmic scale size of individuals
(small sizes on the linear scale), average population
density and corresponding average evenness of the
species structure. Exactly this kind of integral
characteristics combination can be observed in point C
of all figures, represented in Fig. 6. Consequently,
changes in integral characteristics with depth are
largely determined by the share of mesopelagic species
in the fauna. Another consequence of the mesopelagic
fauna exceptional position in the integral
characteristics space is the analogy between some
Fig. 10 Vertical distribution of the integral characteristics. On axes of ordinates—depth, m. On axes of abscissa: N—number
of animals, ind./km2; M—their biomass, kg/km2; W—mean weight of an individual, kg/ind.; J—evenness; S—species riches;
H—diversity, bit/ind. Values of three former variables are presented by their decimal logarithms.
0.0 0.2 0.4 0.6 0.8 1.0
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
10
features of mesopelagic and tropical fauna. In the
pelagic region, the depth increase corresponds to the
movement from the poles to the tropics [27]. Note that
as the depth increases, the environment conditions
stability grows, the water exchange intensity and
productivity decline, so the result of the depth increase
is the same as the geographical latitude decrease. Thus,
if we draw a parallel in the sequence
chorology-geography-ecology (depth-geographical
position-environment factors) one could make the
following conclusion.
The environment conditions stability decreases and
the water exchange intensity increases in the depth to
surface direction, the same as by moving from the
equator to the high latitudes and from the ocean center
to its periphery. The environment ecologic capacity
and intensity of the biogeochemical cycle as a whole
increase. As the depth decreases, the primary
production, variability of the population density and W
of animals, numerical and weighting prevalence of few
dominants over the others increase respectively, while
S and H decrease.
It is important that regular change of biocoenotic
system status in integral characteristics space takes
place not only depending on its position in real space
(latitudes, longitudes, depths, the distances to the
shores), but also depending on time.
The daily changes of the system have been described
before. In the seasonal dynamics of integral
characteristics, one could trace the following
tendencies. The winter-spring period is characterized
by S, J and H reduction, as well as reduction of
hydrobionts abundance and increase of their W. The
summer-autumn period is characterized by opposite
transformations. Two extreme points, situated in the
opposite sections of each coordinates system,
correspond to the directionality of such changes in all
analyzed spaces of integral characteristics (Fig. 11).
S-A
W-S
S-A
S-A
S-A
W-S
W-S
W-S
Fig. 11 Seasonal redistribution of point’s density inside figures along inclined axis. By letters are designated extreme points of the system tendencies: W-S in winter-spring, S-A in summer-autumn period.
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
11
Let us now take notice of the fact that in the analyzed
region there are 3 periods [28]: 1980-1990—sardine
and pollack epoch of fish abundance,
1991-1995—transition period of sharp abundance
reduction, 1996-2005—the period of low level and new
growth of fish capacity. The corresponding tendencies
of long-term dynamics of all analyzed indices of the
system are generally represented in Fig. 12. The dark
arrows mark the general tendencies of the change that
took place by the beginning of 1990s: the macrofauna
abundance reduced, J grew considerably, W increased,
while the changes of S turned out to be indefinite (an
increase in some water areas, and decrease in the
others). These changes correspond to the centrifugal
tendency from point A. As it is shown above, it
describes the state of offshore shelf and continental
slope biocoenotic systems with high productivity, high
population density, and marked prevalence of
dominant species. Since the 2nd half of 1990s all the
changes, marked on the schemes with light arrows,
have been described by centripetal motion to point C,
that is to the tops of the “mounts” and the center of the
“cylinder”: increase of S, further moderate increase of J,
minor growth of abundance and reduction of W.
The main ecological sense of these processes is as
follows: the centrifugal tendency from point А
corresponds to the reduction of dominants abundance
(Walleye pollack, Pacific sardine, Pacific herring and
other relatively large nekton representatives), while the
centripetal tendency to point C corresponds to the
А А
АА
C C
C
C
Fig. 12 Long-term tendencies of integral parameters. Dark arrows show the transition from the “epoch of high abundance” (1980s) to the “stage of the resources reduction” (1991-1995), and light arrows—the following transition to the “epoch of lowered productivity” (1996-2005). The extreme states of the system discussed before (talking about spatial variability, see Fig. 6) are marked by the letters A and C.
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
12
growth of small mesopelagic species share. It has
previously been shown that dynamic water exchange
and high water productivity correspond to point A,
while weak water exchange, abiotic conditions stability
and low primary production correspond to point C.
Similar directions of plurannual time vector and space
vector, which describes circum-continental trends, is
by no means a coincidence. It totally conforms to the
hypothesis [28-33] that large-scale biocoenotic
alterations in the northwestern Pacific are connected
with the ratio of the shelf and oceanic water landscape
areas. As a rule, their shares are inversely proportionate
to each other and change in reversed phase.
Accordingly, there are alternative groups of species,
some of which are favored by shelf landscapes
predomination, while the others are favored by oceanic
landscapes predomination. And not without reason,
upon the system transition from state A to state C, the
share of not only small mesopelagic fish and squid, but
also of salmon, which spend major part of marine
period of life far away from the shore, has increased
considerably.
The analysis of the integral characteristics changes
in the respective periods has shown that the spatial
and plurannual variability vectors are unidirectional
(Fig. 13). Thus, the reason for plurannual ecosystem
alteration during the analyzed period of time
(1979-2005) generally consists in the change of water
exchange regime, which caused the shift to the oceanic
water landscapes predomination over the shelf ones.
Such biotopic transformations caused the shift of
biocoenotic equilibrium to the correspondent group of
species predomination. The way it affected the system
integral characteristic complex has been analyzed
before.
As we can see at the outline scheme (Fig. 13), the
three vectors of space-time variability are differently
directed and perpendicular to each other. It creates the
points dispersion—depending on the system state,
position in space and time, the points density is
redistributed within the mount. At the same time,
12
3
Fig. 13 Three vectors of integral characteristics temporary variability. Directions of: 1—daily, 2—seasonal, 3—long-term changes.
neither point can leave its boundaries, because it is
drawn to the rigid interrelations carcass. This
“skeleton” is formed by small number of relatively
simple regularities. By no means are all of them purely
biological.
These are the metabiological fundamental
magnitude relation: S cannot be higher than N; if S is
constant, the lower limit of J decreases as N grows; if N
is constant, the lowest possible J will increase as S
grows, until S is equal to the given N.
The semibiological (particularly
hydrological-biogeographic, bio- and
physical-chemical, thermodynamic, ethological)
regularities are as follows:
Environment ecological capacity is directly
proportional to the water exchange intensity
(consequence of the fact that in the places of intense
circulation, the food capacity of the water increases and
the negative influence of density factor decreases—the
more dynamic the water exchange is, the more intense
are the substance flows: the inflow of the substances,
consumed by hydrobionts and the outflow of the
substances, excreted by them);
The concentration density is inversely
proportionate to the average individual weight of the
animals (consequence of the constancy of total
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
13
metabolism to substance and energy flows ratio);
The evenness is inversely proportionate to the
hydrobionts concentration density (consequence of the
fact that individuals, similar in species, size, biological
state, are inclined to form thick single-species
aggregations—schools, swarms, etc.).
4. Conclusions
The analyzed schemes allow to: (1) vividly prove the
postulate that the main principles of life organization in
the pelagial are common for the whole World Ocean,
but their displays are different in different parts of it,
and this community doesn’t mean the homogeneity of
all aquatic environment in macrofauna integral
characteristics, but helps to explain the generally
observed local differences and even predict most
possible values and ratios of the characteristics in some
point, on the basis of its spatial position; (2) underline
the interrelation and unity of the origin of two types of
spatial regularities: “horizontal”—geographical
(connected with geographic latitude and longitude) and
“vertical”—chorologic (connected with depth); (3)
connect the spatial variability with time plurannual
variability of the system according to the complex of its
integral characteristics, and both of them with such
ecosystem parameters as environmental conditions
stability and water exchange dynamics, which
determine the ecologic capacity of biotope, and,
consequently, biologic production and general
intensity of biogeochemical cycle.
One of the consequences of all the aforesaid is the
following: the reasons for large-scale plurannual
changes in pelagic cenosis abundance, composition
and structure are not in the rise of temperature, but in
water exchange regime shift.
References
[1] C.E. Shannon, A mathematical theory of communication, Bell Syst. Techn. J. 27 (1948) 379-423, 623-656.
[2] E.C. Pielou, The measurement of diversity in different types of biological collections, J. Theor. Biol. 13 (1966) 131-144.
[3] I.V. Volvenko, Species diversity of the northwest Pacific
pelagic macrofauna, Izv. TINRO 149 (2007) 21-63. [4] I.V. Volvenko, Species diversity of macrofauna biomass
in the pelagic northwest Pacific, Izv. TINRO 153 (2008) 27-48.
[5] I.V. Volvenko, Species richness of the northwest Pacific pelagic macrofauna, Izv. TINRO 153 (2008) 49-87.
[6] I.V. Volvenko, Species structure evenness of the northwest Pacific pelagic macrofauna: 1. Number equitability, Izv. TINRO 156 (2009) 3-26.
[7] I.V. Volvenko, Species structure evenness of the northwest Pacific pelagic macrofauna: 2. Biomass equitability, Izv. TINRO 156 (2009) 27-45.
[8] I.V. Volvenko, Abundance of the northwest Pacific pelagic macrofauna: 1. Number, Izv. TINRO 158 (2009) 3-39.
[9] I.V. Volvenko, Abundance of the northwest Pacific pelagic macrofauna: 2. Biomass, Izv. TINRO 158 (2009) 40-74.
[10] I.V. Volvenko, Average individual weight (size) of animals from pelagic macrofauna in the northwest Pacific, Izv. TINRO 158 (2009) 75-116.
[11] I.V. Volvenko, Interrelations between integral parameters of pelagic macrofauna in the northwest Pacific, Izv. TINRO 159 (2009) 3-34.
[12] I.V. Volvenko, General principles of spatial-temporal variability of integral parameters for pelagic macrofauna in the northwest Pacific, Izv. TINRO 159 (2009) 43-69.
[13] I.V. Volvenko, General patterns of spatiotemporal distribution of pelagic macrofauna integrative characteristics in the northwest Pacific, Bulletin of the Far Eastern Brunch of the Russian Academy of Sciences 3 (145) (2009) 23-31.
[14] I.V. Volvenko, Multidimensional space of the integrative characteristics: The invariance of its structures for the different biocenotic assemblages, Izv. TINRO 168 (2012) 3-19.
[15] V.G. Bogorov, The biological structure of the ocean, Dokl. AN SSSR 128 (4) (1959) 819-822.
[16] V.G. Bogorov, The biological productivity of the ocean and features of its geographical distribution, Voprosy Geografii 84 (1970) 80-102.
[17] V.G. Bogorov, L.A. Zenkevich, The biological structure of the ocean, in: Ecology of Aquatic Organisms, Nauka Press, Moscow, 1966, pp. 3-14.
[18] L.A. Zenkevich, The biological structure of the ocean, Zool. Zh. 27 (2) (1948) 113-124.
[19] V.G. Bogorov, Biomass of zooplankton and productive areas in the Pacific Ocean, geographical zonation of the ocean, in: Biology of the Pacific Ocean, Part 1, Plankton, Nauka Press, Moscow, 1967, pp. 221-227.
[20] O.I. Koblents-Mishke, V.V. Volkovinsky, Yu.G. Kabanova, Plankton primary production of the world
General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western Pacific and Biological Structure of Ocean
14
ocean, Scientific Committee on Oceanographic Research (SCOR) Symp. Sci. Explor. South Pacific, National Academy of Science, Washington, 1970, pp. 183-193.
[21] P.A. Moiseev, The Living Resources of the World Ocean, Pischevaya Promyshlennost Press, Moscow, 1969.
[22] P.A. Moiseev, Fishery Production of the World Ocean and Its Utilization, Oceanology: Ocean Biology[Online], Vol. 2. Biological Productivity of the Ocean, Nauka Press, Moscow, 1977, pp. 289-314. English translation, http://www.archive.org/stream/oceanologybiolog00nort/oceanologybiolog00nort_djvu.txt (accessed Dec. 14, 2011).
[23] V.P. Shuntov, Seabirds and Biological Structure of the Ocean, Dalnevostochnoe Kniznoye Izdatelstvo Press, Vladivostok, 1972.
[24] M.E. Vinogradov, Oceanology. Ocean biology. Vol. 1. Biological Structure of the Ocean, Nauka Press, Moscow, 1977.
[25] L.A. Zenkevich, Z.A. Filatova, G.M. Beliaev, T.A. Lukiyanova, I.A. Suetova, Quantitative distribution of the zoobenthos in the world ocean, Bull. Moip. Biol. Sect. 76 (3) (1971) 27-33.
[26] Z.N. Dontsova, Sergey Semenovich Neustruev, Nauka, Moscow, 1967.
[27] R.H. Whittaker, Communities and Ecosystems, Macmillan Publishing Co, New York, 1970.
[28] V.P. Shuntov, Biology of Far-Eastern Seas of Russia, Vol. 1, TINRO-Center Press, Vladivostok, 2001.
[29] V.P. Shuntov, The state of knowledge of long-term cyclic fluctuations of fish abundance in the far-eastern seas, Biol. Morya 3 (1986) 3-15. English translation: Soviet J. Mar. Biol. 12 (1987) 127-137.
[30] V.P. Shuntov, Reconstructions in the pelagic ecosystems of the Okhotsk Sea-The real fact, Rybnoye Khozyaistvo (Fisheries) 1 (1998) 25-27.
[31] V.P. Shuntov, I.V. Volvenko, A.F. Volkov, K.M. Gorbatenko, S.Yu. Shershenkov, A.N. Starovoitov, New data about condition of pelagic ecosystems of the Okhotsk and Japan Seas, Izv. TINRO 124 (1998) 139-177.
[32] I.V. Volvenko, Analysis of the rate of alternativeness in abundance dynamics of different species in the case of absent continuous long time series data: An example of the Okhotsk Sea nekton, Izv. TINRO 139 (2004) 78-90.
[33] I.V. Volvenko, E.A. Titiayeva, Dominance in nekton and macroplankton fauna of epipelagical water layer of the northern Okhotsk Sea, Izv. TINRO 126 (1999) 58-81.