1-General Patterns Spatial-temporary Distribution of the ...General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western

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.


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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


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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.


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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).


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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


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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.


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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).


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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


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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.


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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.


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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,

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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


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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.

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Documents

1-General Patterns Spatial-temporary Distribution of the ...General Patterns of Spatial-Temporary Distribution of the Integral Characteristics of Pelagic Macrofauna of the North-Western