TMP 814 F

Selforganizology
Vol. 1, No. 2, 1 September 2014
International Academy of Ecology and Environmental Sciences

Selforganizology
Volume 1, Number 2, 1 September 2014
Editor-in-Chief
WenJun Zhang
Sun Yat-sen University, China
International Academy of Ecology and Environmental Sciences, Hong Kong
E-mail: zhwj@mail.sysu.edu.cn, wjzhang@iaees.org
Editorial Board
Awf Al-Kassir (University of Extremadura, Spain)
Ramiz M. Aliguliyev (National Academy of Sciences, Azerbaijan)
Andre Bianconi (Sao Paulo State University (Unesp), Brazil)
Chris Cannings (University of Sheffield, UK)
Manuel De la Sen (University of Basque Country, Spain)
Alessandro Ferrarini (University of Parma, Italy)
Zeljko Kanovic (University of Novi Sad, Serbia)
Abdul Qadeer Khan (University of Azad Jammu & Kashmir, Pakistan)
Zdzislaw Kowalczuk (Gdansk University of Technology, Poland)
Lev V. Nedorezov (University of Nova Gorica, Slovenia)
Quanke Pan (Northeastern University, China) <br>
Sergio Valero Verdu (Universidad Miguel Hernandez de Elche, Spain)
Hafiz Abdul Wahab (Hazara University, Pakistan)
ZhiGuo Zhang (Sun Yat-sen University, China)
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Selforganizology, 2014, 1(2): 51-61
Article
Structure and dynamics of Lithocolletis ringoniella-Parasitoids food

web in apple orchards of Shaanxi, China
Xin Li, Yuyu Liu

College of Plant Protection, Northwest A & F University, Yangling 712100, Shaanxi, China
E-mail: lixin57@hotmail.com
Received 3 April 2014; Accepted 5 May 2014; Published online 1 September 2014
Abstract
The formation and evolution of food web is a self-organizing process. A food web, L. ringoniella-Parasitoids
food web, was proposed in present study. With the apple pest Lithocolletis ringoniella as the basic host, four
parasitoids, Apanteles theivorae, Sympiesis sericeicornis; Ageniaspis testaceipes, and Sympiesis Foerst are
included in the food web. In this food web, A. theivorae and A. testaceipes are primary parasitoids of L.
ringoniella. A. theivorae mainly parasitizes apodous larva of L. ringoniella while A. testaceipes only chooses L.
ringoniella egg to parasitize (egg-larva endoparasitization). S. Foerst and S. sericeicornis are facultative
hyper-parasitoids. They can parasitize not only the larvae and pupae of L. ringoniella, but also A. theivorae. S.
sericeicornis can be hyper-parasitized by S. Foerst. The occurrence mechanism and population dynamics of L.
ringoniella and parasitoids, and parasitization effect of parasitoids in apple orchards of Shaanxi, China, were
described in detail.
Keywords food web; Lithocolletis ringoniella; Apanteles theivorae; Sympiesis sericeicornis; Ageniaspis
testaceipes; Sympiesis Foerst; structure; apple; China.
Selforganizology
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EditorinChief: WenJun Zhang
1 Introduction
Lithocolletis ringoniella Mats. (Lepidoptera: Gracilariidae) is an important defoliator pest in the apple
orchards of northern China, Japan and Korea (Zhang and Zhao, 1996; Zhang et al., 2001). The larvae of L.
ringoniella usually live inside plant leaves and feed mesophyll, causing bubble-like cystics, reducing effective
leaf area for photosynthesis, and causing earlier defoliation (Sun et al., 2001; Zhou et al., 2002). It will not
only lead to the loss of organic nutrition but also affect the next year’s fruit quality and production. Since its
first serious outbreaks in 1992 and 1993, the pest has widely occurred in main apple-producing areas of
northern China (Zhao, 1995). Its occurrence and damage are still increasing year by year and has become a
predominant insect pest from the secondary pest over the past twenty years (Shi et al., 2003; Qiao and Hua,
2005).
Many control practices indicated that chemical insecticides were not quietly effective in the control of L.
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52 Selforganizology, 2014, 1(2): 51-61
ringoniella. Natural enemies are proved to be a major agent for natural equilibrium of L. ringoniella.
Dominant natural enemies of L. ringoniella in Shaanxi, China, include Ageniaspis testaceipes Raz, Apanteles
theivorae, Sympiesis sericeicornis Nees, etc (Sun et al., 1987). Some other insects, including ladybirds,
lacewings, etc., are also in the list of natural enemies. Parasitic rate of these natural enemies may reach
30%-50% in the apple orchards without or few chemical pesticide sprayings.
The present study aimed to propose a L. ringoniella-Parasitoids food web and describe the structure and
dynamics of the food web in apple orchards of Shaanxi, China, in order to understand self-organizing
mechanism of the food web and to provide some knowledge for the biological control of L. ringoniella.
2 Materials and Methods

2.1 Sampling L. ringoniella
Apples trees were sampled from mid-April to early December 2008 in Yangling, Shaanxi, China. The
sampling was conducted every 10 days. A total of 24 samplings were conducted. For each sampling, several
leaves with cystics caused by L. ringoniella were randomly sampled and taken to laboratory for recording the
number of larvae, pupae and puparium.
2.2 Sampling parasitoids of L. ringoniella
Apples trees were sampled from August to October 2007 and April to December 2008 in Yangling, Shaanxi,
China. The sampling was conducted every 10 days. Five sites, with one tree in each site, were selected in the
orchard. For a tree, four directions (East/South/West/North), three layers (upper/middle/lower) and two sides
(inside/outside; only for middle and lower layers), in total of 20 sampling units (i.e., positions), were sampled.
For each position, several leaves with cystics caused by L. ringoniella were randomly sampled and taken to
laboratory for recording species and number of L. ringoniella parasitoids. All samples were combined for
further analysis.
2.3 Hyperparasitism of parasitoids
In laboratory, parasitoids were dissected under microscopy to check hyperparasitism. Species and number of
hyperparasitization were recorded.
2.4 Data treatment
The emergence rate of L. ringoniella adults from pupae was calculated as
(puparium/larvae+pupae+puparium)*100%
And the parasitic rate of parasitoids was recorded as
(parasitized hosts/total hosts)*100%
3 Results and Analysis

3.1 Structure of food web
3.1.1 Reported parasitoids of L. ringoniella
According to a report from Japan, there were 20 species of parasitoids for L. ringoniella, among which some
are hyper-parasitoids (Han et al., 2004). The most occurred parasitoids included Apanteles theivorae (Genus:
Apanteles), Sympiesis sp., Sympiesis laevifrons Kamijo and Sympiesis sericeicornis Nees (Genus: Sympiesis),
Ageniaspis testaceipes Raz (Genus: Ageniaspis), Pnigalio sp. (Family: Eulophidae).
All of these parasitoids are mainly divided into two categories, i.e., the endoparasitoids that live and
mature inside host body, such as A. theivorae, A. testaceipes, and the ectoparasitoids that live and mature
outside host body, such as the species of genus Sympiesis, and P. sp.
In China, in total of 8 parasitoids belonging to 5 families of parasitoids of L. ringoniella have been
recorded, in which A. testaceipes, S. sericeicornis, and A. theivorae are dominant species (Hou, 1987, 1989;
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Selforganizology, 2014, 1(2): 51-61 53
Zhang, 1991). Occurrence of the three species is almost synchronous to that of L. ringoniella (Sun et al., 1987).
In Shaanxi, occurrence generations of A. testaceipes are almost the same as L. ringoniella and it has a stronger
time-synchronity with L. ringoniella. Major occurrence period of other parasitoids, S. sericeicornis, and A.
theivorae, is between May to August which corresponds to the first generation to the fourth generation of L.
ringoniella. They are seldom found after September (Zhang, 1990).
3.1.2 Description of dominant parasitoids of L. ringoniella in Shaanxi
In present study we found 8 species of parasitoids of L. ringoniella. Among these the dominant parasitoids are
A. testaceipes, S. sericeicornis, Sympiesis Foerst, and A. theivorae, which is similar to previous reports (Hou,
1987; Sun et al., 1987; Zhang, 1991; Chen et al., 2006). A. theivorae is an endoparasitoid (Askew and Shaw,
1986). Each host can only contain a parasitoid. S. sericeicornis and S. Foerst are ectoparasitoids and each host
can contain one parasitoid only. A. testaceipes is an endoparasitoid but a host may breed several or many
parasitoids. We found that in average 10 of A. testaceipes emerged from a body of L. ringoniella.
160
Luochuan
140 Yangling
Number of parasitoids
120
100
80
60
40
20
0
S.sericeicornisA.theivorae A.testaceipes
Fig. 1 Dominant parasitoids of L. ringoniella in apple orchards. In total of 500 cystics of L. ringoniella from different leaves
were examined (Yangling and Luochuan, Shaanxi, China, 2008).
From the population comparison in Fig. 1, we can find that in a sense S. sericeicornis is the most
dominant and stable parasitoids of L. ringoniella in Shaanxi.
(1) A. theivorae
A. theivorae (Apanteles, Microgastrinae, Braconidae) lays an egg inside the body of 1st to 3rd instar larva of L.
ringoniella, and the egg hatches as a larva, and the larva lives and matures inside the host’s body. The
parasitoid breaks away from matured larva of host, leaves the host larva died (Sun et al., 1987) and pupates
beside host body. With less than 10% only (in a few cases, greater than 10%), the parasitic rate of A. theivorae
is less than the other two parasitoids. A. theivorae is a primary parasitoid. Thus it is a beneficial parasitoid. A.
theivorae occurs 5 generations a year and the adult’s emergency dates are almost the same with that of L.
ringoniella.
In nature, the hyperparasitoids of A. theivorae include the species of Pteromalidae, Eurytomidae,
Elasmidae, and Ichneumonidae, et al. Parasitic rate of these hyperparasitoids is in average 27%-72%, and
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sometimes reaches 41%-98%.

Our experiment showed that A. theivorae demonstrates different emergence rates at different times in a day.
Emergence dynamics of A. theivorae are indicated in Table 1 and Table 2.
Table 1 Emergence rate of A. theivorae at different time intervals in a day (Yangling, Shaanxi,
China, 2008).
Time No. No. Total Female Male Sex
females males (%) (%) ratio
Before 6：00am 4.0 2.0 6.0 66.67 33.33 2.00
6：00am-8：00am 44.0 2.0 64.0 68.75 31.25 2.20
8：00am-10：00am 23.0 13.0 36.0 63.89 36.11 1.77
10：00am-12：00am 9.0 5.0 14.0 64.29 35.71 1.80
12：00am-14：00pm 9.0 6.0 15.0 60.00 40.00 1.50
14：00pm-16：00pm 5.0 5.0 10.0 50.00 50.00 1.00
16：00pm-18：00pm 2.0 1.0 3.0 66.67 33.33 2.00
18：00pm-20：00pm 2.0 1.0 3.0 66.67 33.33 2.00
20：00pm-22：00pm 5.0 1.0 6.0 83.33 16.67 5.00
22：00pm-24：00pm 4.0 2.0 6.0 66.67 33.33 2.00
Total 107.0 56.0 163.0 65.64 34.36 1.91
Table 2 Emergence rate of A. theivorae in different days (Yangling, Shaanxi, China,

2008).
Date No. No. Total Female Male (%)
(Day) females males (%)
1 5.00 0.00 5.00 4.67 0.00
2 8.00 1.00 9.00 7.48 1.79
3 10.00 2.00 12.00 9.35 3.57
4 13.00 3.00 16.00 12.15 5.36
5 14.00 15.00 29.00 13.08 26.79
6 14.00 11.00 25.00 13.08 19.64
7 17.00 12.00 29.00 15.89 21.43
8 16.00 7.00 23.00 14.95 12.50
9 7.00 5.00 12.00 6.54 8.93
10 3.00 0.00 3.00 2.80 0.00
11 0.00 0.00 0.00 0.00 0.00
Total 107.00 56.00 163.00 100.00 100.00
(2) S. sericeicornis and S. Foerst

S. sericeicornis or S. Foerst lays an egg (sometimes hyperparasitization is found) outside the body of the
higher instar larva or pupa of L. ringoniella; the egg hatches as a larva, and then sucks body fluid of the host.
The matured parasitoid finally leaves its host larva. They occur 5 generations in a year and the adult’s
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emergency dates are slightly later than or similar to that of L. ringoniella. S. sericeicornis and S. Foerst
parasitize not only the larvae and pupae of L. ringoniella but also A. theivorae. Moreover, S. Foerst can
parasitize S. sericeicornis.
The parasitization and hyperparasitization of S. sericeicornis and S. Foerst is indicated in Table 3.
Table 3 Parasitization and hyperparasitization of S. sericeicornis and S. Foerst (Yangling, Shaanxi, China, 2008).
Host of S. sericeicornis and S. Number Percentage Female Male Sex ratio
Foerst (%)
A. theivorae 1522 93.09 945 577 1.64
Pupa of L. ringoniella 15 0.92 14 1 14.00
Larva of L. ringoniella 98 5.99 90 8 11.25
Total 1635 100.00 959 586 1.64
From Table 3, we conclude that even though S. sericeicornis and S. Foerst are hyper-parasitoids of L.
ringoniella, the parasitization has been from their hyperparasitization to A. theivorae, which accounts for
93.09% of the total. Their parasitization to L. ringoniella pupae accounts for 0.92% only. In this sense, S.
sericeicornis and S. Foerst are not of significance in the biological control of L. ringoniella.
Table 4 Emergence statistics of A. testaceipes from 110 of L. ringoniella hosts (Yangling, Shaanxi, China, 2008).
No. hosts for
No. pupae of A. No. hosts for 1 No. hosts for 2 No. hosts for 3 No. hosts for 4
complete
testaceipes non-emergenced non-emergenced non-emergenced non-emergenced
emergence
4 0 0 0 0 2
5 0 0 1 0 0
6 1 0 0 0 0
7 0 0 0 0 1
8 3 1 3 1 4
9 9 1 4 2 2
10 17 1 2 4 0
11 12 5 2 1 2
12 11 0 2 1 0
13 9 1 0 0 0
14 2 1 0 0 0
15 2 0 0 0 0
Total 66 10 14 9 11
(3) A. testaceipes
A. testaceipes is an important parasitoid of L. ringoniella. A. testaceipes lays an egg into the body of L.
ringoniella larva and experiences a polyembryony. Larva of A. testaceipes cocoons and pupates inside the host
body. Each host may emerge up to 15 of A. testaceipes adults (Sun et al., 2006). It occurs 5 generations in a
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year and the emergence dates of adults are basically coincident with that of L. ringoniella. In Shaanxi, it has a
higher parasitic rate to L. ringoniella for the first generation and the fifth generation.
Table 4 shows the emergence statistics of A. testaceipes from 110 of L. ringoniella hosts.
3.1.3 Food web
From the past reports and our investigation, we may conclude that A. theivorae and A. testaceipes are primary
parasitoids of L. ringoniella. A. theivorae mainly parasitizes apodous larva of L. ringoniella while A.
testaceipes only chooses L. ringoniella egg to parasitize (egg-larva endoparasitization). S. Foerst and S.
sericeicornis are facultative hyper-parasitoids. They can parasitize not only the larvae and pupae of L.
ringoniella, but also A. theivorae. And S. sericeicornis can be hyper-parasitized by S. Foerst. Details of
parasitic mechanism are indicated in Table 5.
Table 5 Parasitic mechanism of four dominant parasitoids of L. ringoniella.

Host stage for
Parasitism Parasitism Parasitized
Parasitoid parasitoid’s
type level host stage
departure
Apanteles Konobiont endoparasitism Primary Apodous
Footed larva
theivorae Monoparasitism parasitism larva
Sympiesis Facultative
Idiobiont ectoparasitism Footed Footed larva,
sericeicor hyperparasitis
Monoparasitism larva, pupa pupa
nis m
Facultative
Sympiesis Idiobiont ectoparasitism Footed Footed larva,
hyperparasitis
Foerst Facultative polyparasitism larva, pupa pupa
m
Konobiont endoparasitism
Ageniaspis Primary
Polyembryony Egg Footed larva
testaceipes parasitism
monoparasitism
Based on known knowledge and our investigation, we have constructed a L. ringoniella-Parasitoids food
web in apple orchards of Shaanxi, China, as illustrated in Fig. 2.
Fig. 2 L. ringoniella-Parasitoids food web in apple orchards of Shaanxi, China, constructed from known knowledge and our
investigation.
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3.2 Dynamics of food web

3.2.1 Dynamics of L. ringoniella
The investigated field dynamics of L. ringoniella, with different development stages, is shown in Fig. 3.
100.00
Larva
90.00 Pupa
80.00 Adult
No. insects/No. cystics
70.00
60.00
50.00
40.00
30.00
20.00
10.00
0.00
12/03
10/09
11/09
11/21
11/28
10/29
5/06
5/17
5/30
6/12
6/18
6/30
7/09
7/18
7/28
8/11
8/20
8/30
9/08
9/18
9/28
4/18
4/29
10/1
9
Date
Fig. 3 Dynamics of L. ringoniella (Yangling, Shaanxi, China, 2008).
It can be concluded from Fig. 3 that in Yangling, Shaanxi, the larvae, pupae and adults of L. ringoniella
occurs firstly in late April to early May. The larva population of L. ringoniella discontinuously declines from
late April to early December. Pupa population has a peak in early and mid-June, and keeps at a very low level
from July to September. Pupa population starts to grow since early October.
Starting from the overwinter generation, the adult population tends to show a significant peak in June,
seconded by a weak peak during late September to late October.
3.2.2 Dynamics of four parasitoids
As an exception, A. testaceipes shows polyembryony in the reproduction. In average 10 individuals of A.
testaceipes can emerge from a host body. The dynamics of four parasitoids of L. ringoniella is shown in Fig. 4.
From Fig. 4, it can be found that the parasitic rate of A. testaceipes reaches as high as 90%; for S.
sericeicornis, A. theivorae and S. Foerst, it is 70%, 35% and 25% respectively. The dominancy rank of these
parasitoids is coincident with previous reports (Sun et al., 1987; Chen et al., 2003).
A. theivorae occurs mainly between May to September, corresponding to the first generation to fourth
generation of the host larvae, particularly between late May to early July, corresponding to the second
generation to third generation of the host larvae. This is similar to the reported (Zhang, 1990). Overall it
declines from late April to late December.
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100.00
90.00 A. theivorae
80.00 S. sericeicornis
A. testacesipes
No.parasitoids/No.
70.00
S.
60.00 Foerst
50.00
40.00
30.00
20.00
10.00
0.00
7/09
7/18
7/28
8/11
8/20
8/30
9/08
9/18
9/28
6/12
6/18
6/30
5/30
10/19
5/17
10/09
10/29
11/28
12/03
4/18
4/29
5/06
11/09
11/21
Date
Fig. 4 Dynamics of four parasitoids of L. ringoniella (Yangling, Shaanxi, 2008).
100.00
90.00
80.00
Parasitic rate
70.00
60.00
50.00
40.00
30.00
20.00
10.00
0.00
11/28
12/03
9/08
9/18
9/28
8/20
8/30
5/17
5/30
6/12
6/18
6/30
7/09
7/18
7/28
8/11
5/06
10/09
11/21
10/19
11/09
10/29
4/18
4/29
Date
Fig. 5 Dynamics of parasitization effect to L. ringoniella (Yangling, Shaanxi, 2008).
S. sericeicornis starts to occur since late May and increases to reach a maximum in late August to
mid-September, and keeps at a higher level thereafter. It thus mainly parasitizes the third generation, fourth
generation and overwinter generation of the host.
A. testaceipes has two population peaks in early September and late November. Overall its population
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discontinuously grows from late May to December.

Table 6 shows emergence dynamics of A. theivorae and S. sericeicornis, investigated in Yangling,
Shaanxi.
Table 6 Emergence dynamics of A. theivorae and S. sericeicornis (Yangling, Shaanxi, China, 2008).
Date No. A. theivorae S. sericeicornis No. Emergence Cocoon/cystics

(m/d) cocoons Number % Number % emergenced rate (%) (%)
4/18 0 0 0 0 0 0 0.00 0.00

4/29 0 0 0 0 0 0 0.00 0.00
5/06 1 1 100.00 0 0 1 100.00 33.33
5/17 1 1 100.00 0 0 1 100.00 7.69
5/30 1 0 0.00 1 100.0 1 100.00 3.45
6/12 27 23 88.46 3 11.54 26 96.30 12.86
6/18 129 106 86.89 16 13.11 122 94.57 15.62
6/30 296 137 59.31 94 40.69 231 78.04 20.96
7/09 619 186 41.06 267 58.94 453 73.18 41.18
7/18 122 51 64.56 28 35.44 79 64.75 16.46
7/28 399 60 20.00 240 80.00 300 75.19 25.73
8/11 74 6 9.38 58 90.63 64 86.49 21.20
8/20 29 4 16.67 20 83.33 24 82.76 21.64
8/30 55 7 15.91 37 84.09 44 80.00 22.54
9/08 4 1 33.33 2 66.67 3 75.00 6.90
9/18 9 1 14.29 6 85.71 7 77.78 11.39
9/28 20 2 14.29 12 85.71 14 70.00 12.90
10/09 43 1 3.70 26 96.30 27 62.79 26.54
10/19 51 1 3.23 30 96.77 31 60.78 21.79
10/29 46 1 3.70 26 96.30 27 58.70 19.91
11/09 205 10 7.35 126 92.65 136 66.34 50.00
11/21 138 9 9.00 91 91.00 100 72.46 56.33
11/28 41 0 0.00 22 100.0 22 53.66 47.13
12/03 60 0 0.00 30 100.0 30 50.00 49.59
3.2.3 Dynamics of parasitization effect

Dynamics of parasitization effect of four parasitoids (A. theivorae, S. sericeicornis, A. testaceipes, S. Foerst) is
illustrated in Fig. 5.
From Fig. 5 we know that parasitization starts to occur since late April; overall it grows and
asymptotically reaches a maximum of 89% in late April to early September. Thereafter the parasitization keeps
at a higher level. It is obvious that the parasitization effect of parasitoids to L. ringoniella is much significant.
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4 Discussion
The conclusions in present study are drawn from comprehensive and complete analysis on previous reports
and our investigation. Therefore, our conclusions are reliable. More experiments and investigations to
overwinter generations of L. ringoniella and parasitoids are needed, which will help to further understand the
dynamics and overwinter mechanism of these insects.
We constructed real food web and described its structure and dynamics. Further studies can be conducted
on modeling of population and community dynamics (Ivanchikov and Nedorezov, 2011, 2012; Elsadany, 2012;
Nedorezov, 2012; Zhang, 2012a, 2012b), topological analysis of food web (Dorman, 2011; Kuang and Zhang,
2011; Zhang, 2011), self-organizing process (Zhang, 2012a, 2012b, 2013), etc.
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Normal University of Science and Technology, 9: 16
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Article
Two-dimensional ordered cluster analysis of component groups in

self-organization
WenJun Zhang1,2, YanHong Qi3, ZhiGuo Zhang4

1 2
School of Life Sciences, Sun Yat-sen University, Guangzhou, China; International Academy of Ecology and Environmental
Sciences, Hong Kong
3
Libraries of Sun Yat-sen University, Sun Yat-sen University, Guangzhou, China
4
Department of Computer Science, Sun Yat-sen University, Guangzhou, China
Received 23 May 2014; Accepted 28 June 2014; Published online 1 September 2014
Abstract
An algorithm for two-dimensional cluster analysis of component groups, originally from Zhang et al., (2004),
was introduced in this study. The algorithm composes of three procedures, i.e., calculation of distance
measures, randomization statistic test, and ordered clustering of components.
Keywords two-dimensional cluster analysis; components; groups.
Selforganizology
1 Introduction
Ecological processes at the landscape level are self-organizing processes. Geographicaly, a landscape is
composed of many mosaics (components, as called in self-organization) occupied by various organisms. For
most organisms, mosaic structure determines their feeding, survival and prosperity (Bell et al., 1991; Zhang et
al., 2004). As a membrane or barrier or channel, the boundary between two mosaics affects the movement and
migration of organisms (Forman, 1995; Zhang and Schoenly, 1999; Qi, 2002; Qi and Zhang, 2002; Zhang,
2012, 2013; Zhang et al., 2013). Relationship between mosaic boundaries and biotic/abiotic factors has been
confirmed in many studies (Gillison and Brewer, 1985; Hansen et al., 1988; Holland et al., 1991). It was found
that homogeneous and continuous mosaics with less natural barriers were conducive to the spread of
organisms. The number and shape of mosaics determines ecosystems at the landscape level. In this study, an
algorithm for two-dimensional cluster analysis of component groups, originally from Zhang et al., (2004), was
introduced to address ordered classification of mosaics.
2 Method
Assume there are m mosaic indices and n sampling sites (mosaics, or components). Now we have the n×m
matrix (yij), and the algorithm is as follows:
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2.1 Choose distance measures

Distance measures are used to measure the similarity between mosaics (components). A larger distance value
means a lower similarity between two mosaics. We chose four categories, a total of 14 types of distance
measures here (Qi, 2002; Zhang, 2007; Zhang et al., 2004; Zhang, 2012; Zhang and Fang, 1982; Krebs, 1989).
(1) Minkowski distance measures. They include Euclidean distance, Manhattan distance, and Chebyshov
distance:
zij=(∑mk=1(yik - yjk)2/m)1/2
zij= ∑mk=1|yik - yjk|/m
zij= max k |yik - yjk| i,j=1,2,…,n
(2) Correlation based measures. They include Pearson correlation and angular cosine:
zij= 1-∑mk=1 ((yik - yib)(yjk- yjb) )/(∑mk=1 (yik - yib)2 ∑mk=1 (yjk - yjb)2)1/2
zij= 1-∑mk=1 (yik yjk）/(∑mk=1 yik2 ∑mk=1 yjk2)1/2 i,j=1,2,…,n
The two categories of indices are used to the indices with real values.
(3) Nominal distance measures. They include linkage coefficient, and three co-linkage coefficients based
distance measures:
zij=1-(x2/(x2+n..))1/2
zij=1-(x2/(n.. max(p-1,q-1)))1/2
zij=1-(x2/(n.. min(p-1,q-1)))1/2
zij=1-(x2/(n.. ((p-1)(q-1))1/2))1/2
where
x2= n..(∑pi=1∑qj=1nij2/(ni. n.j)-1)

n.. =∑pi=1 ni. , ni. =∑qj=1 nij , n.j =∑pi=1 nij
where there are p nominal values, i.e., t1, t2,…, tp, for sampling site i and q nominal values, i.e., r1, r2,…, rq, for
sampling site j. Assume that nkl is the number of sampling site i takes value tk and sampling site j takes value rl,
k = 1, 2, . . . , p; l = 1, 2, . . . ,q.
(4) Boolean distance measures, including point correlation, quadratic correlation, two angular cosine
functions, and Jaccard coefficient based distance measures:
zij=1-(ad-bc)/((a+b)(c+d)(a+c)(b+d))1/2
zij=1-sin((a+d-(b+c))/(a+b+c+d)*3.1415926/2)
zij=1-(a*a/((a+b)(a+c)))1/2
zij=1-(a*a*d*d/((a+b)(a+c)(b+d)(c+d)))1/2
zij=(b+c)/(b+c+d) i,j=1,2,…,n
where both sampling site i and sampling site j take values 0 or 1. a is the number of both sampling site i and
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sampling site j take value 0, b is the number of sampling site i takes 0 and sampling site j takes 1, c is the
number of sampling site i takes 1 and sampling site j takes 0, and d is the number of both sampling site i and
sampling site j take value 1.
2.2 Randomization test
If min yij <0, let yij = yij - min yij , i=1,2,…,n; j=1,2,…,m. zij is the decimal number of yij, and dij=10Zij. Let
yij= yij max j dij i=1,2,…,n
Through these transformations all of the values in sampling data become integers which are equivalent to
numbers of individuals. The randomization test used in present study is based on the idea that, if no difference
exists, then the distribution of individuals in sampling sites i and j will be a result of allocating the mixed
sampling site values at random into two sampling site of size equal to those of the original sampling site
(Solow, 1993; Manly, 1997; Zhang and Schoenly, 2001; Zhang et al., 2004; Zhang, 2007). The randomization
test procedure is described as the follows. Assume the two sampling sites to be tested are i and j, i=1, 2, …,
n−1; j>i. The sij=∑mk=1 yik+∑mk=1 yjk individuals of the combined sampling site are randomly reallocated into
two randomized sampling sites with ∑mk=1 yik and ∑mk=1 yjk labeled individuals. Calculate the expected
absolute distance (similarity) between the two randomized sampling sites and compare whether it is not less
than the absolute distance (similarity) between the true sampling sites i and j. Repeat the simulation many
times, calculate the number of the expected are not less than the absolute distance (similarity) between the
sampling site i and j, take the percentage as the p value. The p value is used to make statistical test. The
threshold p value for test may be defined as 0.05, 0.01, etc. If the calculated p value is less than (in the case of
similarity, larger than) p threshold, then the difference between sampling sites i and j is statistically significant.
2.3 Ordered regionalization of mosaics (components)
This procedure aims to conduct ordered regionalization of mosaics according to their similarity (Fang, 1979;
Zhang, 1993). Mosaics with significant difference belong to different groups (clusters). In the ordered cluster
analysis, the mosaics in the same group are geographically adjacent, and the lines between any two mosaics
that belong to different groups are not intersected.
Assume the two-dimensional coordinate of each sampling site (mosaic) is ( yi11, yi12), i=1,2,…,n. And for
sampling sites i and j, i=1,2,...,n-1; j>i , calculate
g1(i,j)={k, i | yk12>( yj12- yi12)( yk11- yi11)/( yj11- yi11)+ yi12}

g2(i,j)={k, j | yk12<( yj12- yi12)( yk11- yi11)/( yj11- yi11)+ yi12}
G1(i,j)={k, j | yk12>( yj12- yi12)( yk11- yi11)/( yj11- yi11)+ yi12}
G2(i,j)={k, i | yk12<( yj12- yi12)( yk11- yi11)/( yj11- yi11)+ yi12}
Thus there will be n(n-1) possible classifications. In each classification, for the sampling site i (i1≤i≤i2-1) in
sub-group 1 and the sampling site j (i2≤j≤i3-1) in sub-group 2, calculate the p value, pij, in the randomization
test, and choose the classification that meets the condition:
min pbar=(∑i2-1i=i1∑i3-1j=i2 pij)/(( i2- i1)( i3- i2))
Suppose there have been k groups, obtain two sub-groups for each group, and choose a group to make
sub-classification. If k=n, then stop the procedure. Therefore a cluster graph with different statistical
significance levels can be achieved.
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If all values of max min pbar are larger than the given threshold p value, then the landscape is continuous
and homogeneous. Or else there are several mosaics with significant differences.
The algorithm is implemented as a Java program, TwoDimBoundDetector, based on JDK 1.1.8, in which
several classes and an HTML file is included (http://www.iaees.org/publications/software/index.asp) (Fig. 1
and 2). In data file, the first two columns are 2-dimensional coordinates of sampling sites and the remaining
columns are for every index.
Fig. 1 Applet window of the algorithm-Input.
Fig. 2 Applet window of the algorithm-Output.
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3 Applications
3.1 Two-dimensional distribution of grass families in Zhuhai
Sampling survey was conducted on May 22, 2008 in a grass community of Zhuhai, China (Fig. 3). In total of
100 samples were taken along a linear transect. Each sample had a size of 1m × 1m. Grass families and cover
(%) in each sample were recorded. Totally 16 grass families were found and treated as the indices for cluster
analysis.
1 11 21 31 41 51 61 71 81 91
2 12 22 32 42 52 62 72 82 92
3 13 23 33 43 53 63 73 83 93
4 14 24 34 44 54 64 74 84 94
5 15 25 35 45 55 65 75 85 95
6 16 26 36 46 56 66 76 86 96
7 17 27 37 47 57 67 77 87 97
8 18 28 38 48 58 68 78 88 98
9 19 29 39 49 59 69 79 89 99
10 20 30 40 50 60 70 80 90 100
Fig. 3 Geographical distribution of sampling sites in a meadow of Zhuhai.
We chose Euclidean distance, and randomize 100 times; the threshold p value is 0.05. The classification
results of 100 samples were as follows:
P_test value=0.0017<=0.05, statistically significant

(1 ) (2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37
38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72
73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 )
(1 ) (2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38
39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73
74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (20 )
(1 ) (2 ) (3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37
38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72
73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (20 )
(1 ) (2 ) (3 ) (4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36
37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71
72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (20 )
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(1 ) (2 ) (3 ) (4 5 6 7 9 10 11 12 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36
37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71
72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 5 6 7 9 10 11 12 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36
37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72
73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 5 6 7 9 10 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38
39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74
75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (11 12 ) (60 ) (8 )
(20 )
(1 ) (2 ) (3 ) (4 5 6 7 9 10 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38
39 40 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (41 ) (11 12 ) (60 ) (8 )
(20 )
(1 ) (2 ) (3 ) (4 5 6 7 9 10 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 33 34 35 36 37 38 39
40 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76
77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (32 ) (41 ) (11 12 ) (60 )
(8 ) (20 )
(1 ) (2 ) (3 ) (4 5 6 7 9 10 13 14 15 16 17 18 19 21 22 23 24 25 26 27 28 29 30 31 33 34 35 36 37 38 39
40 42 43 44 45 46 47 48 49 50 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77
78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (51 ) (32 ) (41 ) (11 12 )
(60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 6 7 9 10 14 15 16 17 18 19 23 24 25 26 27 28 29 30 33 34 35 36 37 38
39 40 42 43 44 45 46 47 48 49 50 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76
77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (51 ) (32 ) (41 ) (11 12 )
(60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 6 9 10 14 15 16 17 18 19 23 24 25 26 27 28 29 30 33 34 35 36 37 38 39
40 42 43 44 45 46 47 48 49 50 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77
78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (7 ) (51 ) (32 ) (41 ) (11
12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 6 9 10 14 15 16 17 18 19 23 24 25 26 27 28 29 30 33 34 35 36 37 38 39
40 42 43 44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78
79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (50 ) (7 ) (51 ) (32 ) (41 )
(11 12 ) (60 ) (8 ) (20 )
P_test value=0.0593>0.05, not statistically significant
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 14 15 16 17 18 19 23 24 25 26 27 28 29 30 33 34 35 36 37 38 39
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40 42 43 44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78
79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 )
(11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 14 15 16 17 18 19 23 24 25 26 27 28 29 30 33 34 35 36 37 38 39
40 42 43 44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78
79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 95 96 97 98 99 100 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 )
(41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 14 15 16 17 18 19 24 25 26 27 28 29 30 33 34 35 36 37 38 39 40
43 44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80
81 82 83 84 85 86 87 88 89 90 91 92 93 95 96 97 98 99 100 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 )
(32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 14 15 16 17 18 19 24 25 26 27 28 29 30 33 34 35 36 37 38 39 40
43 44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80
81 82 83 84 85 86 87 88 89 90 91 92 93 96 97 98 99 100 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 )
(32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 26 27 28 29 30 33 34 35 36 37 38 39
40 43 44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79
80 81 82 83 84 85 86 87 88 89 90 91 92 93 96 97 98 99 100 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 )
(51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 26 28 29 30 33 34 35 36 37 38 39 40
43 44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80
81 82 83 84 85 86 87 88 89 90 91 92 93 96 97 98 99 100 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 )
(51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 28 29 30 33 34 35 36 37 38 39 40 43
44 45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81
82 83 84 85 86 87 88 89 90 91 92 93 96 97 98 99 100 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 )
(7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82
83 84 85 86 87 88 89 90 91 92 93 96 97 98 99 100 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 )
(50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82
83 84 85 86 87 88 89 90 91 92 93 97 98 99 100 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 )
(50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
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(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82
83 84 85 86 87 88 89 90 91 92 93 98 99 100 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 )
(6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82
83 84 85 86 87 88 89 90 91 92 93 99 100 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 )
(6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82
83 84 85 86 87 88 89 90 91 92 93 100 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 )
(94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82
83 84 85 86 87 88 90 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23
42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 59 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 87 88
90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 )
(23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 61 62 63 64 65 66 67 68 71 72 73 74 75 76 77 87 ) (59 69 70 78 79 80
88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 )
(23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 61 62 63 64 65 66 67 68 71 72 73 75 76 77 87 ) (74 ) (59 69 70 78 79
80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 )
(95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 49 52 53 54 55 56 57 58 61 62 63 64 65 67 68 71 72 73 75 76 77 87 ) (66 ) (74 ) (59 69 70
78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 )
(95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 52 53 54 55 56 57 61 62 63 64 65 71 72 73 75 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69
70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 )
(27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
IAEES www.iaees.org

(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 52 53 54 55 56 57 61 62 63 64 71 72 73 75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59
69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 )
(27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 46 47 48 52 53 54 55 56 57 61 62 63 64 71 72 73 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 )
(59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 )
(26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 47 48 52 53 54 55 56 57 61 62 63 64 71 72 73 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 )
(74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 )
(28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 )
(20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 47 48 53 54 55 56 57 63 64 72 73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 )
(74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 )
(28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 )
(20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 24 25 29 30 33 34 35 36 37 38 39 40 43 44
45 47 48 53 54 55 56 57 63 64 72 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 )
(66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 )
(96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 )
(8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 29 30 33 34 35 36 37 38 39 40
43 44 45 47 48 53 54 55 56 57 63 64 72 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77
87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 )
(97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 )
(60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 29 33 34 35 36 37 38 39 40 43
44 45 47 48 53 54 55 56 57 63 64 72 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76
77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 )
(97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 )
(60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 29 33 34 35 36 37 38 39 40 43
44 45 47 48 53 54 55 56 63 64 72 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68
76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 )
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(98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 )
(11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 29 33 34 35 36 37 38 39 40 43
44 45 48 53 54 55 56 63 64 72 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58
67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 )
(98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 )
(11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 29 33 34 35 36 38 39 40 43 44
45 48 53 54 55 56 63 64 72 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49
58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 )
(99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 )
(41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 35 36 38 39 40 43 44 45
48 53 54 55 56 63 64 72 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 )
(49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 )
(99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 )
(41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 35 36 38 39 40 43 44 45
48 53 54 55 56 63 72 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 )
(65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 )
(89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 )
(32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35
36 38 39 40 45 48 55 56 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 )
(65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 )
(89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 )
(32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35
36 38 40 45 48 55 56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 )
(75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92
93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 )
(51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35
36 38 40 45 48 55 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 )
(75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92
93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 )
(51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
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(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35
36 38 45 48 55 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 )
(46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86
91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 )
(7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35
36 38 45 55 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 )
(46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86
91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 )
(7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35
36 45 55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62
71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84
85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 )
(50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35
36 45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52 61
62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82 83
84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 )
(6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35 )
(36 45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 ) (52
61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 ) (81 82
83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 ) (94 )
(6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 63 72 ) (35 )
(36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 ) (73 )
(52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88 90 )
(81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 ) (23 42 )
(94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 33 34 43 44 53 54 72 ) (63 )
(35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 )
(73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88
90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 )
(23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 34 44 53 54 72 ) (33 43 ) (63 )
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(35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 ) (30 )
(73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79 80 88
90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 ) (95 )
(23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 34 53 54 72 ) (44 ) (33 43 )
(63 ) (35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 ) (57 )
(30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70 78 79
80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 )
(95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 34 53 72 ) (54 ) (44 ) (33
43 ) (63 ) (35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 )
(57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70
78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 )
(95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 25 34 72 ) (53 ) (54 ) (44 )
(33 43 ) (63 ) (35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 )
(57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70
78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 )
(95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 34 72 ) (25 ) (53 ) (54 ) (44 )
(33 43 ) (63 ) (35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 ) (47 )
(57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 ) (59 69 70
78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 ) (26 ) (27 )
(95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 ) (34 72 ) (25 ) (53 ) (54 )
(44 ) (33 43 ) (63 ) (35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 )
(47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 )
(59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 )
(26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 ) (34 ) (72 ) (25 ) (53 ) (54 )
(44 ) (33 43 ) (63 ) (35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 ) (37 )
(47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 ) (74 )
(59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 ) (28 )
(26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 ) (20 )
(1 ) (2 ) (3 ) (4 13 21 22 31 ) (5 9 10 15 16 17 18 19 ) (14 ) (24 ) (34 ) (72 ) (25 ) (53 ) (54 )
(44 ) (33 ) (43 ) (63 ) (35 ) (36 ) (45 ) (55 ) (38 ) (48 ) (40 ) (56 ) (39 ) (64 ) (29 )
(37 ) (47 ) (57 ) (30 ) (73 ) (52 61 62 71 ) (46 ) (75 ) (65 ) (49 58 67 68 76 77 87 ) (66 )
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(74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 )

(28 ) (26 ) (27 ) (95 ) (23 42 ) (94 ) (6 ) (50 ) (7 ) (51 ) (32 ) (41 ) (11 12 ) (60 ) (8 )
(20 )
....
Fig. 4 indicates 2-dimensional classification for p=0.0482 (<=0.05, statistically significant).
Fig. 4 Classification for p=0.0482 (<=0.05, statistically significant).
3.2 Two-dimensional vegetation distribution in Ngari

Data on vegetation types and environmental indicators for Ngari, Tibet, was chosen for the case application
(Chang, 1991). A total of 15 sampling sites (they were sampled based on vegetation types as low mountain
desert, low mountain steppe desert, low mountain desert steppe, middle mountain desert, middle mountain
grassland desert, middle mountain desert grassland, upper-middle mountain desert grassland, upper-middle
mountain grassland, sub-alpine grasslands, sub-alpine meadow grassland, alpine grassland desert, alpine desert
grassland, alpine grasslands, alpine meadow grassland and alpine meadow, with numbers 1-15), and 8 indices
of mosaic characteristics (mean annual temperature, temperature of the hottest month, the coldest temperature,
annual precipitation, biological temperature, transpiration rate, surface soil organic matter content and pH
value) were used. Latitude and longitude, i.e., geographical coordinates, were used as the two-dimensional
coordinates (Fig. 5; Table 1).
We chose Euclidean distance, and randomize 100 times; the threshold p value is 0.05. The classification
results were as follows:
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Fig. 5 Geographical distribution of sampling sites in Ngari.
Table 1 Data on vegetation types and environmental indicators for Ngari, Tibet (Zhang et al., 1991).
Latitude Longitude Mean Temper. The Annual Biologica Transpiration Surface soil PH
(o) (o) annual of the coldest precipi. l temper. rate organic
temper. hottest temper. matter
month content
78.917 31.867 5.3 18.3 -5 123.6 7.7 2.2 0.9 8
78.783 31.8 7.3 20.6 -1 161.3 9.9 1.3 1.1 7.9
78.917 31.833 3.4 16 -8.2 97.8 6 2.8 1.9 7.8
79.317 33.6 -2.5 11 -14.8 48.3 3 2.9 0.5 8.3
79.433 32.867 -3 10.7 -14.1 68.7 3 2.5 0.7 8
80.083 32.333 -1.2 11.1 -13.3 92 3.2 2.3 1 7.9
80.917 31.817 -1.4 10.3 -13.4 120.6 2.9 1.6 1.5 7.8
80.667 31.383 -1.9 9.2 -13.6 123.2 2.5 1.3 2.4 7.5
80.6 32.3 -3 8.8 -14.6 112.3 2.3 1.2 2.6 7.5
80.767 31.133 -2.7 8 -13.8 142.2 2.2 0.6 4.1 6.9
80.45 34.683 -9 4.5 -18.8 81.2 0.8 0.4 0.4 8.4
81.667 34.833 -8.1 5.7 -19.1 84.1 0.9 1.1 0.9 8.1
80.933 34.3 -7.7 5.6 -18.3 87 1 0.8 2.4 7.6
80.533 33.833 -7.6 5.2 -17.5 101.3 1.1 0 3.2 7.2
80.533 32.667 -6.5 5.2 -16.1 129.5 1.3 -0.5 6.5 6.4
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We chose Pearson correlation coefficient, and randomize 100 times; the threshold p value is 0.05. The
results were as follows:

(1 2 3 5 6 7 8 9 10 11 12 13 14 15 ) (4 )
(1 2 3 5 6 7 8 9 11 12 13 14 15 ) (10 ) (4 )
(1 2 3 5 6 7 9 11 12 13 14 15 ) (8 ) (10 ) (4 )
(1 2 3 5 6 9 11 12 13 14 15 ) (7 ) (8 ) (10 ) (4 )
(1 2 3 5 6 11 12 13 14 15 ) (9 ) (7 ) (8 ) (10 ) (4 )
(1 2 3 5 6 11 12 13 14 ) (15 ) (9 ) (7 ) (8 ) (10 ) (4 )
(1 2 3 5 11 12 13 14 ) (6 ) (15 ) (9 ) (7 ) (8 ) (10 ) (4 )
Between-sampling site p values matrix is listed as the following:
1 0.23 0.38 0 0 0 0 0 0 0 0 0 0 0 0
0.23 1 0 0 0 0 0 0 0 0 0 0 0 0 0
0.38 0 1 0 0 0.24 0.09 0.05 0.02 0 0 0 0 0 0
0 0 0 1 0.04 0 0 0 0 0 0 0 0 0 0
0 0 0 0.04 1 0.22 0 0 0 0 0.13 0.26 0.13 0 0
0 0 0.24 0 0.22 1 0.24 0.1 0.42 0 0.02 0.04 0.07 0.05 0
0 0 0.09 0 0 0.24 1 1 0.9 0.34 0 0 0 0.02 0.01
0 0 0.05 0 0 0.1 1 1 0.93 0.78 0 0 0 0.04 0.1
0 0 0.02 0 0 0.42 0.9 0.93 1 0.23 0 0 0.01 0.23 0.11
0 0 0 0 0 0 0.34 0.78 0.23 1 0 0 0 0 0.2
0 0 0 0 0.13 0.02 0 0 0 0 1 1 0.95 0.08 0
0 0 0 0 0.26 0.04 0 0 0 0 1 1 0.97 0.14 0
0 0 0 0 0.13 0.07 0 0 0.01 0 0.95 0.97 1 0.56 0
0 0 0 0 0 0.05 0.02 0.04 0.23 0 0.08 0.14 0.56 1 0.07
0 0 0 0 0 0 0.01 0.1 0.11 0.2 0 0 0 0.07 1
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Zhang WJ, Schoenly KG. 1999. IRRI Biodiversity Software Series. III BOUNDARY: a program for detecting
boundaries in ecological landscapes. IRRI Technical Bulletin No. 3. International Rice Research Institute,
Manila, Philippines
Zhang WJ, Schoenly KG. 2001. A randomization test and software to compare ecological communities.
International Rice Research Notes, 26(2): 48-49
Zhang YT, Fang KT. 1982. Introduction to Multivariate Statistics. 393-401, Science Press, Beijing, China
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Article
Diversity and aggregation patterns of plant species in a grass

community
Ran Li1, DanTing Chen1, GongWei Liang1, Wei Huang1, HanQing Li1, AiJuan Li1, PengHui Huang1,
BiNi Chen1, Bing Chen1, Liang Chen1, LinXing Chen1, ShaoJing Chen1, QinSon Wang1, HeCon Wang1,
Jing Wang1, CiHui Wu1, YuanHui Xu1, WenJun Zhang1,2
1
School of Life Sciences, Sun Yat-sen University, Guangzhou, China;
2
International Academy of Ecology and Environmental Sciences, Hong Kong
Received 21 June 2014; Accepted 23 July 2014; Published online 1 September 2014
Abstract
Both composition and aggregation patterns of species in a community are the outcome of community
self-organizing. In this paper we conducted analysis on species diversity and aggregation patterns of plant
species in a grass community, Zhuhai, China. According to the sampling survey, in total of 47 plant species,
belonging to 16 families, were found. Compositae had 10 species (21.3%), seconded by Gramineae (9 species,
19.1%), Leguminosae (6 species, 12.8%), Cyperaceae (4 species, 8.5%), and Malvaceae (3 species, 6.4%). The
results revealed that the means of aggregation indices Iδ, I and m*/m were 21.71, 15.71 and 19.89 respectively
and thus individuals of most of plant species strongly followed aggregative distribution. Iwao analysis
indicated that both individuals of all species and clumps of all individuals of all species followed aggregative
distribution. Taylor’s power law indicated that individuals of all species followed aggregative distribution and
aggregation intensity strengthened as the increase of mean density. We held that the strong aggregation
intensity of a species has been resulted from the strong adaptation ability to the environment, the strong
interspecific competition ability and the earlier establishment of the species. Fitting goodness of the mean, I, Iδ,
m*/m with probability distributions demonstrated that the mean (density), I, Iδ, and m*/m over all species
followed Weibull distribution rather than normal distribution. Lophatherum gracile, Paederia scandens (Lour.)
Merr., Eleusine indica, and Alternanthera philoxeroides (Mart.) Griseb. were mostly aggregative, and Oxalis
sp., Eleocharis plantagineiformis, Vernonia cinerea (L.) Less., and Sapium sebiferum (L.) Roxb, were mostly
uniform in the spatial distribution. Importance values (IV) showed that Cynodon dactylon was the most
important species, seconded by Desmodium triflorum (L.) DC., Cajanus scarabaeoides (L.) Benth., Paspalum
scrobiculatum L., and Rhynchelytrum repens. Oxalis sp., Eleocharis plantagineiformis, and Vernonia cinerea
(L.) Less. were the least important species in the community. Summed dominance ratio (SDR2) revealed that
Cynodon dactylon and Desmodium triflorum (L.) DC. were the most dominant species in the community,
followed by Rhynchelytrum repens, Paspalum scrobiculatum L., and Cajanus scarabaeoides (L.) Benth.
Keywords grass community; aggregation pattern; importance value; summed dominance ratio.
Selforganizology
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1 Introduction
Community succession is a self-organizing process over time and space (Clements, 1916.). Both composition
and aggregation patterns of species in a community are the outcome of community self-organizing, and the
former in turn affect the community succession. It is important to understand the structure and aggregation
patterns of species in the community (Liu and Zhang, 2011). It also plays the fundamental role in the
maintainence of biodiversity (Altieri, 1994; IRRI, 1998; Stang et al., 2007; Zhang, 2012; Jayakumar et al.,
2011; Ahmad et al., 2013).
In present study, we will conduct analysis on species diversity and aggregation patterns of plant species in
a grass community, Zhuhai, China, using various indices and methods, in order to further understand the
species composition and spatial structure of the grass community.

2.1 Sampling survey
Sampling survey was conducted on May 12, 2008 in a grass community of Zhuhai, China (Fig. 1). In total of
100 samples were taken (10 by 10 samples; Fig. 2). Each sample had a size of 1m × 1m. Grass species,
frequency (ratio of species presence), cover (area covered/total area) and density (individuals per sample) in
each sample were recorded.
Fig. 1 The grassland for sampling survey.
2.2 Methods
2.2.1 Aggregation patterns
For the detection of aggregation patterns of plant species, the following three indices were used
Index of clumping: I= s2 / m
Index of dispersion: Iδ= n Σxj(xj-1) / (Σxj(Σxj-1))
Index of patchiness (Iwao, 1968): m* / m= 1+ (s2 - m) / m2
where m, s2: mean and variance of number of individuals in a sample, n: number of samples (here n=100), xj:
density of plant individuals in the sample j, j=1,2,…,n. I>1, aggregative distribution; I<1, uniform distribution;
I≈1, random distribution. Iδ>1, aggregative distribution; Iδ<1, uniform distribution; Iδ≈1, random distribution;
m*/m >1, aggregative distribution; m*/m <1, uniform distribution; m*/m≈1, random distribution.
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A1 B1 C1 D1 E1 F1 G1 H1 I1 J1
A2 B2 C2 D2 E2 F2 G2 H2 I2 J2
A3 B3 C3 D3 E3 F3 G3 H3 I3 J3
A4 B4 C4 D4 E4 F4 G4 H4 I4 J4
A5 B5 C5 D5 E5 F5 G5 H5 I5 J5
A6 B6 C6 D6 E6 F6 G6 H6 I6 J6
A7 B7 C7 D7 E7 F7 G7 H7 I7 J7
A8 B8 C8 D8 E8 F8 G8 H8 I8 J8
A9 B9 C9 D9 E9 F9 G9 H9 I9 J9
A10 B10 C10 D10 E10 F10 G10 H10 I10 J10
Fig. 2 Sampling design.
Iwao regression (Iwao, 1968) and Taylor’s power law are represented by
m* = a+ b m
and log(s2) = log (a)+ b log (m)
in Iwao regression, b >1, aggregative distribution; b <1, uniform distribution; b ≈1, random distribution. In
Taylor’s power law, log (a) > 0 and b≥1, aggregative distribution; log (a) = 0 and b ≈ 1, random distribution;
log (a) < 0 and b < 1, uniform distribution.
Negative binomial distribution is expressed as
pr= (k+r-1)!pr / (r!(k-1)!Qk+r) r>0
where p= (s2-m) / m, Q=1+p, k= m2 / (s2-m), pr: the probability that a sample contains r individuals, m: mean, s2:
variance.
2.2.2 Importance and dominance of species
The importance value (IV) of a plant species is
IV = (relative cover + relative frequency + relative density) / 3
where
Relative cover=area covered / total area covered by all species
Relative frequency= frequency of the species / total frequencies of all species
Relative density= density of the species / total density of all species
The summed dominance ratio for two factors (SDR2) is expressed as
SDR2= (density ratio + relative frequency) / 2 × 100%
2.2.3 Diversity of community

Diversity was measure by two indices
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Simpson index: Sp=1 - Σxj(xj-1) / (N(N-1))

Shannon-wiener index: H′= -Σ(Pj ln Pj )
where S: total number of plant species, Pj= xj / N, N: total number of individuals.

2.2.4 One-dimensional cluster analysis
One-dimensional cluster method was used to cluster species into groups without change their natural order (Qi,
2005; Zhang, 2012), for example, the order of importance values from the large to the small.
3 Results
According to the sampling survey, in total of 47 plant species, belonging to 16 families, were found.
Compositae has 10 species (21.3%), seconded by Gramineae (9 species, 19.1%), Leguminosae (6 species,
12.8%), Cyperaceae (4 species, 8.5%), Malvaceae (3 species, 6.4%). Convolvulaceae, Amaranthaceae,
Oxalidaceae and Euphorbiaceae has 2 species (4.3%) respectively. The remaining 7 families account for
14.9% of total species.
3.1 Aggregation patterns
Aggregation patterns of 47 species are indicated in the Table 1. It can be found from Table 1 that most of the
plant species follow aggregative distribution, and a little species follow uniform distribution (or random
distribution). Lophatherum gracile, Paederia scandens (Lour.) Merr., Eleusine indica, and Alternanthera
philoxeroides (Mart.) Griseb. are mostly aggregative, and Oxalis sp., Eleocharis plantagineiformis, Vernonia
cinerea (L.) Less., and Sapium sebiferum (L.) Roxb, are mostly uniform in the spatial distribution.
Table 1 Aggregation patterns of 47 species.

ID Plant species Mean I Iδ m*/m Mean of Spatial distri.
I, Iδ & pattern
m*/m
1 Lophatherum gracile 0.58 54.06 93.16 92.483 79.901 Aggregative distri.
2 Paederia scandens (Lour.) Merr. 0.06 6 100 84.333 63.444 Aggregative distri.
3 Eleusine indica 0.04 4 100 76.000 60.000 Aggregative distri.
Alternanthera philoxeroides (Mart.)
4 0.03 3 100 67.667 56.889 Aggregative distri.
Griseb.
5 Ischaemum aristatum L. 5.77 121.3 18.99 21.849 54.046 Aggregative distri.
6 Eragrostis pilosa (L.) Beauv 1.02 43.43 42.59 42.598 42.873 Aggregative distri.
7 Rhus chinensis Mill. 0.11 6.225 52.72 48.500 35.815 Aggregative distri.
8 Artemisia indica Willd． 0.39 15.95 39.95 39.333 31.744 Aggregative distri.
9 Panicum repens L. 2.43 45.75 17.05 19.416 27.405 Aggregative distri.
10 Verbena officinalis L. 0.31 10.99 33.91 33.226 26.042 Aggregative distri.
11 Euphorbia hirta L. 0.08 3.707 39.24 34.838 25.928 Aggregative distri.
12 Lespedeza bicolor 0.18 6.664 33.93 32.467 24.354 Aggregative distri.
13 Paspalum scrobiculatum L. 9.46 57.99 7.523 7.024 24.179 Aggregative distri.
14 Kyllinga brevifolia 0.91 21.12 23.13 23.110 22.453 Aggregative distri.
15 Chrysopogon aciculatus 1.18 24.3 20.72 20.746 21.922 Aggregative distri.
16 Bidens pilosa L. 2.39 31.25 13.58 13.657 19.496 Aggregative distri.
17 Aster subulatus Michx. 0.99 16.82 16.99 16.980 16.930 Aggregative distri.
18 Oxalis corniculata L. 0.15 4.226 23.81 22.507 16.848 Aggregative distri.
19 Moghania philippinensis (Merr. et 0.09 2.939 25 22.544 16.828 Aggregative distri.
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Rolfe) Li
20 Cajanus scarabaeoides (L.) Benth. 9.07 37.28 5.072 5.000 15.784 Aggregative distri.
21 Desmodium triflorum (L.) DC. 11.21 37.54 4.223 4.260 15.341 Aggregative distri.
22 Centella asiatica (L.) Urb 0.46 8.67 17.79 17.674 14.711 Aggregative distri.
23 Vetiveria zizanioides (L.) Vach 0.22 4.461 17.32 16.732 12.838 Aggregative distri.
24 Rhynchelytrum repens 4.78 25.41 6.455 6.107 12.657 Aggregative distri.
25 Ageratum conyzoides 3.35 22.53 7.499 7.427 12.485 Aggregative distri.
26 Erigeron acer L. 0.43 6.12 13.07 12.907 10.699 Aggregative distri.
27 Cynodon dactylon 15.87 25.93 2.556 2.571 10.352 Aggregative distri.
28 Wedelia triloba 0.75 8.63 11.21 11.173 10.338 Aggregative distri.
29 Pteris vittata L. 0.41 5.622 12.41 12.273 10.102 Aggregative distri.
30 Cayerus exaltatus Retz. 0.33 4.717 12.51 12.264 9.830 Aggregative distri.
31 Sesbania cannabina (Retz.) Poir. 0.39 5.278 12.13 11.969 9.792 Aggregative distri.
32 Bidens bipinnata (L.) 0.64 7.201 9.896 10.689 9.262 Aggregative distri.
33 Sida acuta Burm. F 0.71 7.406 10.06 10.023 9.163 Aggregative distri.
34 Ipomoea triloba 0.22 3.173 11.23 10.877 8.427 Aggregative distri.
35 Cyperus globosus All. 1.97 11.85 6.397 6.508 8.252 Aggregative distri.
36 Alternanthera sessilis (L.) DC 0.54 5.291 9.012 8.946 7.750 Aggregative distri.
37 Sida. rhombifolia L. 1.76 8.92 5.478 5.500 6.633 Aggregative distri.
38 Emilia sonchifolia (L.) DC. 0.16 1.919 11.11 6.744 6.591 Aggregative distri.
39 Mimosa pudica 0.19 2.2 7.612 7.316 5.709 Aggregative distri.
40 Syngonium podophyllum 0.75 4.744 6.222 5.992 5.653 Aggregative distri.
41 Pharbitis purpurea L. 0.61 4.236 6.335 6.305 5.625 Aggregative distri.
42 Ixeris polycephala Cass 0.18 2.061 7.19 6.894 5.382 Aggregative distri.
43 Urena lobata 0.64 3.741 5.32 5.283 4.781 Aggregative distri.
44 Oxalis sp. 0.01 1 0 1.000 0.667 Uniform distri.
45 Eleocharis plantagineiformis 0.01 1 0 1.000 0.667 Uniform distri.
46 Vernonia cinerea (L.) Less. 0.01 1 0 1.000 0.667 Uniform distri.
47 Sapium sebiferum (L.) Roxb 0.01 1 0 1.000 0.667 Uniform distri.
Mean=number of individuals/sample.
The means of Iδ , I and m*/m are 21.71, 15.71 and 19.89 respectively, and the variances of Iδ , I and m*/m
are 700.63, 469.01, 480.09 respectively. There is a strong correlation between Iδ and m*/m (R2=0.969). The
correlations between I and Iδ (R2=0.002) and m*/m (R2=0.012) are not significant.
Iwao regression of 47 species is: m* = 8.638+ 4.49 m, R2=0.396, p<0.00001. This means that both
individuals of all species and clumps of individuals of all species follow aggregative distribution.
Taylor’s power law: log(s2) = 2.529 + 1.542 log (m), R2=0.956, p<0.00001. It means that individuals of
all species follow aggregative distribution, and aggregation intensity will strengthen with the increase of the
mean density. Logically, it is obvious that increased aggregation intensity leads to the increase of the mean
density. We thus hold that the strong aggregation intensity of a species has been resulted from the strong
adaptation ability to the environment, the strong competition ability and the earlier establishment of the
species.
Fitting goodness of mean, I, Iδ, m*/m with normal distribution and Weibull distribution, calculated by
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Matlab functions, are illustrated in Fig. 3. Linear relationship between probability and data means that the
mean and three indices follow Weibull distribution.
Fig. 3 Fitting goodness of the mean, I, Iδ, m*/m with normal distribution and Weibull distribution. Linear relationship between
probability and data means that the mean and three indices follow Weibull distribution rather than normal distribution.
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One-dimensional cluster tree, calculated from the mean of I, Iδ and m*/m in Table 1, is indicated in Fig. 4.
As an example, we give the spatial distribution (individuals per sample) of Cynodon dactylon as the
following
0 0 2 3 0 5 0 0 0 0
0 0 0 11 0 17 0 0 6 18
0 39 22 24 22 10 5 0 11 16
9 26 34 37 25 26 22 19 19 18
8 0 3 27 26 34 9 44 10 8
0 0 0 1 6 54 35 10 7 10
18 13 14 17 20 93 82 45 0 107
35 32 19 4 9 3 16 20 20 3
69 50 43 18 0 22 35 30 0 0
0 12 0 0 0 0 0 0 0 0
For Cynodon dactylon, m= 15.9, s2= 407.373. Thus p= (s2-m) / m= 24.669, Q= 1+ p =25.669, k= 1.372,
and
pr= (1.372+r-1)! 24.669r/(r!( 1.372-1)! 25.669381.371545+r)
χ2=925<χ0.052, therefore Cynodon dactylon follows negative binomial distribution in the community.
Fig. 4 One-dimensional cluster tree of species based on their similarity in aggregation degree (the mean of I, Iδ and m*/m) in
Table 1. Species for ID number 1 to 47 are listed in Table 1.
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3.2 Importance values of plant species

Importance values of 47 species are indicated in the Table 2.
Table 2 Importance values of 47 species.

ID Plant species Relative Relative Relative Importance
cover frequency density value (IV)
1 Cynodon dactylon 0.31 0.68 0.1944 0.3948
2 Desmodium triflorum (L.) DC. 0.1381 0.49 0.1373 0.2552
3 Cajanus scarabaeoides (L.) Benth. 0.0569 0.41 0.1111 0.1927
4 Paspalum scrobiculatum L. 0.0925 0.35 0.1061 0.1829
5 Rhynchelytrum repens 0.0423 0.44 0.0544 0.1789
6 Ageratum conyzoides 0.0444 0.29 0.0403 0.1249
7 Sida rhombifolia L. 0.0276 0.3 0.0216 0.1164
8 Bidens pilosa L. 0.0186 0.26 0.0268 0.1018
9 Urena lobata 0.0081 0.25 0.0078 0.0887
10 Syngonium podophyllum 0.0092 0.24 0.0088 0.086
11 Ischaemum aristatum L. 0.046 0.13 0.0812 0.0858
12 Cyperus globosus All. 0.0322 0.16 0.0245 0.0722
13 Pharbitis purpurea L. 0.005 0.2 0.0075 0.0708
14 Aster subulatus Michx. 0.0072 0.18 0.0121 0.0664
15 Bidens bipinnata (L.) 0.0083 0.17 0.0086 0.0623
16 Chrysopogon aciculatus 0.0114 0.16 0.0145 0.0619
17 Panicum repens L. 0.0345 0.11 0.0339 0.0595
18 Alternanthera sessilis (L.) DC 0.0091 0.16 0.0066 0.0586
19 Sida acuta Burm. F 0.0104 0.13 0.0087 0.0497
20 Wedelia triloba 0.008 0.12 0.0092 0.0457
21 Sesbania cannabina (Retz.) Poir. 0.0042 0.11 0.0048 0.0397
22 Cayerus exaltatus Retz. 0.0019 0.11 0.004 0.0387
23 Mimosa pudica 0.0118 0.1 0.0023 0.038
24 Ixeris polycephala Cass 0.0035 0.1 0.0022 0.0352
25 Erigeron acer L. 0.0056 0.09 0.0053 0.0336
26 Ipomoea triloba 0.0031 0.09 0.0027 0.0319
27 Centella asiatica (L.) Urb 0.0046 0.08 0.0056 0.0301
28 Pteris vittata L. 0.0037 0.08 0.005 0.0296
29 Kyllinga brevifolia 0.0051 0.07 0.0111 0.0288
30 Verbena officinalis L. 0.0064 0.06 0.0038 0.0234
31 Emilia sonchifolia (L.) DC. 0.0023 0.06 0.001 0.0211
32 Artemisia indica Willd． 0.0026 0.05 0.0048 0.0191
33 Vetiveria zizanioides (L.) Vach 0.0022 0.05 0.0027 0.0183
34 Lespedeza bicolor 0.0019 0.05 0.0022 0.018
35 Eragrostis pilosa (L.) Beauv 0.0072 0.03 0.0125 0.0166
36 Moghania philippinensis (Merr. et Rolfe) Li 0.0034 0.04 0.0011 0.0148
37 Oxalis corniculata L. 0.0004 0.04 0.0018 0.0141
38 Lophatherum gracile 0.0039 0.03 0.0071 0.0137
39 Rhus chinensis Mill. 0.0027 0.03 0.0013 0.0114
40 Euphorbia hirta L. 0.0004 0.03 0.001 0.0105
41 Paederia scandens (Lour.) Merr. 0.0014 0.01 0.0007 0.004
42 Sapium sebiferum (L.) Roxb 0.0009 0.01 0.0001 0.0037
43 Eleusine indica 0.0003 0.01 0.0005 0.0036
44 Alternanthera philoxeroides (Mart.) Griseb. 0.0003 0.01 0.0004 0.0035
45 Oxalis sp. 0.0001 0.01 0.0001 0.0034
46 Eleocharis plantagineiformis 0.0001 0.01 0.0001 0.0034
47 Vernonia cinerea (L.) Less. 0.0001 0.01 0.0001 0.0034
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We may find from Table 2 that Cynodon dactylon is relatively the most important species, seconded by
Desmodium triflorum (L.) DC., Cajanus scarabaeoides (L.) Benth., Paspalum scrobiculatum L., and
Rhynchelytrum repens. Oxalis corniculata L., Eleocharis plantagineiformis, and Vernonia cinerea (L.) Less.
are the least important species in the community.
One-dimensional cluster tree, calculated from importance values in Table 2, is indicated in Fig. 5.
Fig. 5 One-dimensional cluster tree of species based on their similarity in species importance in Table 2. Species for ID number 1
to 47 are listed in Table 2.
Table 3 indicates that Cynodon dactylon and Desmodium triflorum (L.) DC. are the most dominant
species in the community, followed by Rhynchelytrum repens, Paspalum scrobiculatum L., and Cajanus
scarabaeoides (L.) Benth.
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3.3 Diversity of plant species

Simpson index and Shannon-wiener index of the community is 0.903 and 2.788 respectively. Thus the
community is relatively diverse.
Table 3 Summed dominance ratio for two factors (SDR2) for 47 species.
Plant species SDR2(%) Plant species SDR2(%)

Cynodon dactylon 100.00 Centella asiatica (L.) Urb 22.50
Desmodium triflorum (L.) DC. 80.00 Chrysopogon aciculatus 22.50
Rhynchelytrum repens 65.00 Kyllinga brevifolia 17.50
Paspalum scrobiculatum L. 65.00 Sida acuta Burm. F 17.50
Cajanus scarabaeoides (L.) Benth. 62.50 Artemisia indica Willd. 15.00
Bidens pilosa L. 52.50 Ageratum conyzoides 15.00
Sida rhombifolia L. 50.00 Vetiveria zizanioides (L.) Vach 15.00
Urena lobata 47.50 Moghania philippinensis (Merr. et Rolfe) Li 15.00
Alternanthera sessilis (L.) DC 47.50 Lespedeza bicolor 15.00
Pharbitis purpurea (L.) 47.50 Cyperus globosus All. 15.00
Oxalis corniculata L. 47.50 Euphorbia hirta L. 15.00
Syngonium podophyllum 47.50 Rhus chinensis Mill. 15.00
Bidens bipinnata (L.) 42.50 Pteris vittata L. 12.50
Ixeris polycephala Cass 40.00 Verbena officinalis L. 10.00
Aster subulatus Michx. 37.50 Eragrostis pilosa (L.) Beauv 7.50
Ischaemum aristatum L. 37.50 Lophatherum gracile 7.50
Sesbania cannabina (Retz.) Poir. 30.00 Vernonia cinerea (L.) Less. 5.00
Wedelia triloba 27.50 Eleusine indica 5.00
Emilia sonchifolia (L.)DC. 25.00 Eleocharis plantagineiformis 5.00
Mimosa pudica 25.00 Sapium sebiferum (L.) Roxb. 5.00
Cayerus exaltatus Retz. 25.00 Alternanthera philoxeroides (Mart.) Griseb. 5.00
Ipomoea triloba 25.00 Oxalis sp. 5.00
Erigeron acer L. 22.50 Paederia scandens (Lour.) Merr. 5.00
Panicum repens L. 22.50
4 Discussion and Main Conclusion

Individuals of most of plant species follow aggregative distribution. In addition, clumps of all individuals of
most species follow aggregative distribution. The plant density, the aggregation indices I, Iδ, and m*/m over all
species follow Weibull distribution, rather than normal distribution. We guess that most of the indices over
multiple species used in ecological studies might follow Weibull distribution. This should be further validated
in the future studies.
Acknowledgment
We would like to thank all students of Department of Ecology, Sun Yat-sen University, for their field sampling
in Zhuai. Ran Li, Dating Chen, and WenJun Zhang are the first authors of this article.
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Article
Interspecific associations and community structure: A local survey

and analysis in a grass community
WenJun Zhang1,2
1 2
School of Life Sciences, Sun Yat-sen University, Guangzhou, China; International Academy of Ecology and Environmental
Sciences, Hong Kong
Received 18 July 2014; Accepted 23 August 2014; Published online 1 September 2014
Abstract
Interspecific associations in the plant community may help to understand the self-organizing assembly and
succession of the community. In present study, Pearson correlation, net correlation, Spearman rank correlation,
and point correlation were used to detect the interspecific (inter-family) associations of grass species (families)
using the sampling data collected in a grass community of Zhuhai, China. We found that most associations
between grass species (families) were positive associations. The competition/interference/niche separation
between grass species (families) was not significant. A lot of pairs of grass species and families with
statistically significant interspecific (inter-family) associations based on four correlation measures were
discovered. Cluster trees for grass species/families were obtained by using cluster analysis. Relationship
among positive/negative associations, interspecific relationship and community succession/stability/robustness
was discussed. I held that species with significant positive or negative associations are generally keystone
species in the community. Although both negative and positive associations occur in the community
succession, the adaptation and selection will finally result in the successful coexistence of the species with
significant positive associations in the climax community. As the advance of community succession, the
significant positive associations increase and maximize in climax community, and the significant negative
associations increase to a maximum and then decline into climax community. Dominance of significant
positive associations in the climax community means the relative stablility and equilibrium of the community.
No significant associations usually account for the majority of possible interspecific associations at each phase
of community succession. They guarantee the robustness of community. They are candidates of keystone
species. Lose of some existing keystone species might be filled with some species previously with no
significant associations. In addition, a Java program, associCoeff, re-writed from my earlier work, was
introduced. A large number of data were thus given also.
Keywords interspecific associations; community analysis; grass; Pearson correlation; net correlation;
Spearman rank correlation; point correlation; cluster analysis; Zhuhai.
Selforganizology
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1 Introduction
Interspecific association refers to between-species interconnectedness on the spatial distribution, which is
usually resulted from differences in the environment and habitat (Greig-Smith, 1983; Kuželová and Chytry,
2004; Meyer, 2006; Waterman and Roth, 2007; Chu et al., 2008; Menzel et al., 2008; Haugaasen and Peres,
2009; Nansen et al., 2009; Desbiez et al., 2010; Oliveira and Dietz, 2011; Ladygina and Rineau, 2012; Lan et
al., 2012; Lin et al., 2012; Gonzalez-Moreno et al., 2013; Zhang et al., 2013). It is the representation of
interactions of species-species and species-environment. Interspecific association is the basis for assembly and
succession of biological communities (Zhang, 2012c). It is important to understand the structure, function and
classification of communities. It also plays the fundamental role in the maintainence of biodiversity (Altieri,
1994; IRRI, 1998; Stang et al., 2007; Zhang, 2012a; Jayakumar et al., 2011; Ahmad et al., 2013). So far a lot
of research on interspecific association have been published. Kuželová and Chytry (2004) reviewed the
difference of interspecific associations in phytosociological data sets between local and regional scale.
Waterman and Roth (2007) compared the benefit and cost of interspecific associations of Cape ground
squirrels with two mongoose species. Desbiez et al. (2010) determined the interspecific association between an
ungulate and a carnivore or a primate. Oliveira and Dietz (2011) studied the interspecific association of two
Brazilian Atlantic forest primates in cabruca agroforest. Lan et al. (2012) combined the spatial distribution in
the interspecific associations of tree species in a tropical seasonal rain forest.
There are many measures for interspecific association, including χ2 test, variance ratio method, coupling
coefficient, co-occurrence percentage, point correlation, the similarity percentage for distribution, Spearman
rank correlation, Pearson correlation, etc (Schoenly and Zhang, 1999; Zhang, 2007, 2011, 2012a, 2012b).
Cluster analysis is based on the similarity between communities/species. It is widely used in the classification
of communities and other objects (Qi, 2003; Zhang, 2012a).
In present study, Pearson correlation, net correlation, Spearman rank correlation, point correlation, and
cluster analysis were used to detect and compare the interspecific (inter-family) associations of grass species
(families) collected in a grass community of Zhuahai, in order to further understand the structure and
succession of grass community.

2.1 Sampling survey
Sampling survey was conducted on April 22, 2007 in a grass community of Zhuhai, China (Fig. 1). In total of
50 samples were taken along a linear transect. Each sample had a size of 1m × 1m, and had an interval of 1m
with the adjacent samples. Grass species and cover (%) in each sample were recorded.
2.2 Interspecific (inter-family) associations
Positive association refers to that two species (families) tend to spatially occur together and negative
association means that two species (families) tend to occur exclusively (Schoenly and Zhang, 1999; Zhang,
2007, 2011, 2012a, 2012b).
Pearson correlation between species (families) i and j is
n n n
rij=∑((aik-aibarr)(ajk-ajbar))/[∑(aik-aibar)2∑(ajk-ajbar)2]1/2
k=1 k=1 k=1
where, -1≤rij≤1, aik and ajk are k-th sample of sampling set of species (families) i and j respectively, aibar and
ajbar are means of aik and ajk respectively, n is the number of samples. The t-test values of Pearson correlation is
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t=rij/[(1-rij2)/(n-2)]1/2
where m is the number of species (families). If t>tα, then the interspecific (inter-family) correlation (association)
is statistically significant; rij>0, positive correlation (association) (mutulism, parasitism, or they require the
similar environmental conditions (niche overlap)); rij<0, negative correlation (association) (competition,
interference, or they require the distinct environmental conditions (niche separation)).
Pearson correlation (association) is a linear association (Zhang, 2011, 2012a, 2012b).
Fig. 1 A profile of the grass community.
Net correlation is an extension of Pearson correlation. Net correlation between species (families) i and j is
calculated as (Zhang, 2012a)
Rij=-rij/(rii*rjj)1/2
where -1≤Rij≤1, and rij is the element in inverse matrix of Pearson correlation matrix.
The t-test values of net correlation is
t=Rij/[(1-Rij2)/(n-m)]1/2
where m is the number of species (families), and n is the number of samples. If t>tα, then the interspecific
(inter-family) correlation (association) is statistically significant.
Spearman rank correlation is (Spearman, 1904; Schoenly and Zhang, 1999; Zhang, 2011, 2012a, 2012b)
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rij=1-6*∑d2/[n(n2-1)]
where -1≤rij≤1, d=r(i)-r(j), and r(i) and r(j) are rank of an element in the sampling set of species (families) i
and j, from the smaller to the larger values in n elements. The above t-test can be used in the significance test
of Spearman rank correlation. If t>tα, then the interspecific (inter-family) correlation (association) is
statistically significant; rij>0, positive correlation (association); rij<0, negative correlation (association).
Point correlation is defined as (Zhang, 2007)
dij=(ad-bc)/((a+b)(c+d)(a+c)(b+d))1/2
where -1≤dij≤1, both species (family) i and species (family) j take values 0 or 1. a is number of samples that
both species (family) i and species (family) j take value 0, b is number of samples that species (family) i takes
0 and species (family) j takes 1, c is number of samples that species (family) i takes 1 and species (family) j
takes 0, and d is number of samples that both species (family) i and species (family) j take value 1. The χ2-test
value of point correlation is
χ2=n(ad-bc)2/[(a+b)(c+d)(a+c)(b+d)].
If χ2>χ2α(1) (χ20.01(1)=6.64, χ20.05(1)=3.84), then interspecific (inter-family) association is statistically

significant; dij>0, positive association; dij<0, negative association.
I have re-writed the program in Zhang (2012a) for easier use. The re-writed Java program, based on JDK
1.1.8, in which several classes and an HTML file is included
(http://www.iaees.org/publications/software/index.asp; associCoeff.html; Fig. 2). In sampling data file, the
first row is sample ID numbers and the first column is taxon ID numbers.
2.3 Cluster analysis
Fussy cluster analysis (Qi, 2003) was used to classify grass families and species. In the algorithm, cosine
coefficient, which is an unstandardized form of Pearson correlation, was chosen as the similarity measure
between families (species).
Fig. 2 Applet window of the program.
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3 Results
According to the sampling survey, in total of 48 grass species, belonging to 17 families, were recorded in the
meadow, as indicated in Table 1 and 2.
Fig. 3 indicates that the sampling is complete for creating a fuller species/families list, i.e., most of taxa
have been collected in the sampling survey.
Table 1 Grass families found in the survey.

1 Euphorbiaceae 10 Connaraceae
2 Leguminosae 11 Umbelliferae
3 Pteridaceae 12 Cyperaceae
4 Lygodiaceae 13 Lycopodiaceae
5 Gramineae 14 Amaranthaceae
6 Apocynaceae 15 Convolvulaceae
7 Malvaceae 16 Commelinaceae
8 Asteraceae 17 Oxalidaceae
9 Onagraceae
Table 2 Grass species found in the survey.
1 Euphorbia hirta 17 Lophatherum gracile 33 Pterocypsela indica
2 Sapium sebiferum 18 Panicum repens 34 Vernonia cinerea

3 Atylosia scarabaeoides 19 Paspalum scrobiculatum 35 Wedelia triloba
4 Desmodium gangeticum 20 Rhynchelytrum repens 36 Jussiaea linifolia
5 Desmodium triflorum 21 Catharanthus roseus 37 Pteroloma triquetrum
6 Sesbania cannabina 22 Sida rhombifolia 38 Centella asiatica
7 Pteris vittata 23 Urena sp. 39 Cayerus exaltatus
8 Lygodium japonicum 24 Urena lobata 40 Cyperus compressus
9 Axonopus compressus 25 Ageratum conyzoides 41 Kyllinga brevifolia
10 Chrysopogon aciculatus 26 Aster subulatus 42 Lycopodium cernuum
11 Cymbopogon citratus 27 Bidens bipinnata 43 Alternanthera philoxeroides
12 Cynodon dactylon 28 Bidens pilosa 44 Alternanthera sessilis
13 Echinochloa crusgalli 29 Emilia sonchifolia 45 Ipomoea triloba
14 Eragrostis minor 30 Erigeron acer 46 Pharbitis purpurea
15 Eragrostis pilosa 31 Eupatorium chinense 47 Zebrina pendula
16 Ischaemum aristatum 32 Mikania micrantha 48 Oxalis corniculata
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Fig. 3 Illustration of sampling completeness for creating a fuller species/families list. 100 times of randomizations were
conducted for each sample size. Error bars are confidential intervals with 95% siginificance level (p=0.05).
3.1 Associations measured by Pearson correlation

Inter-family Pearson correlation coefficients are shown in Table 3.
Table 3 Inter-family Pearson correlation coefficients.

Euphorbiaceae Leguminosae Pteridaceae Lygodiaceae Gramineae Apocynaceae Malvaceae Asteraceae Onagraceae
Euphorbiaceae 1 0.393 0.067 -0.032 0.016 -0.03 0.308 0.114 0.555
Leguminosae 0.393 1 0.019 -0.078 -0.273 -0.076 0.422 0.248 0.724
Pteridaceae 0.067 0.019 1 -0.021 0.048 -0.02 -0.039 -0.059 -0.021
Lygodiaceae -0.032 -0.078 -0.021 1 0.191 -0.021 -0.041 -0.106 -0.023
Gramineae 0.016 -0.273 0.048 0.191 1 -0.118 -0.093 -0.308 0.039
Apocynaceae -0.03 -0.076 -0.02 -0.021 -0.118 1 -0.039 0.063 -0.021
Malvaceae 0.308 0.422 -0.039 -0.041 -0.093 -0.039 1 0.423 0.573
Asteraceae 0.114 0.248 -0.059 -0.106 -0.308 0.063 0.423 1 -0.008
Onagraceae 0.555 0.724 -0.021 -0.023 0.039 -0.021 0.573 -0.008 1
Connaraceae -0.03 -0.076 -0.02 -0.021 -0.105 -0.02 -0.039 -0.09 -0.021
Umbelliferae -0.067 -0.124 -0.044 -0.048 0.151 -0.044 0.138 0.053 0.012
Cyperaceae -0.054 -0.136 -0.036 -0.039 0.354 -0.036 -0.069 -0.087 -0.039
Lycopodiaceae -0.03 -0.029 -0.02 0.056 0.037 -0.02 -0.039 -0.107 -0.021
Amaranthaceae 0.784 -0.033 -0.024 -0.026 0.036 -0.024 -0.023 0.124 -0.026
Convolvulaceae 0.378 0.485 -0.044 -0.047 0.054 -0.044 0.396 -0.093 0.723
Commelinaceae -0.053 -0.133 -0.035 -0.038 0.242 -0.035 -0.068 -0.009 -0.038
Oxalidaceae -0.042 -0.102 -0.028 -0.03 0.056 -0.028 -0.041 -0.055 -0.03
n=50.
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Table 3 Inter-family Pearson correlation coefficients (Continue).

Connaraceae Umbelliferae Cyperaceae Lycopodiaceae Amaranthaceae Convolvulaceae Commelinaceae Oxalidaceae
Euphorbiaceae -0.03 -0.067 -0.054 -0.03 0.784 0.378 -0.053 -0.042
Leguminosae -0.076 -0.124 -0.136 -0.029 -0.033 0.485 -0.133 -0.102
Pteridaceae -0.02 -0.044 -0.036 -0.02 -0.024 -0.044 -0.035 -0.028
Lygodiaceae -0.021 -0.048 -0.039 0.056 -0.026 -0.047 -0.038 -0.03
Gramineae -0.105 0.151 0.354 0.037 0.036 0.054 0.242 0.056
Apocynaceae -0.02 -0.044 -0.036 -0.02 -0.024 -0.044 -0.035 -0.028
Malvaceae -0.039 0.138 -0.069 -0.039 -0.023 0.396 -0.068 -0.041
Asteraceae -0.09 0.053 -0.087 -0.107 0.124 -0.093 -0.009 -0.055
Onagraceae -0.021 0.012 -0.039 -0.021 -0.026 0.723 -0.038 -0.03
Connaraceae 1 -0.044 -0.036 -0.02 -0.024 -0.044 -0.035 -0.028
Umbelliferae -0.044 1 -0.08 -0.044 -0.023 0.113 -0.01 0.166
Cyperaceae -0.036 -0.08 1 -0.036 -0.044 -0.03 0.291 -0.05
Lycopodiaceae -0.02 -0.044 -0.036 1 -0.024 -0.044 -0.035 -0.028
Amaranthaceae -0.024 -0.023 -0.044 -0.024 1 -0.053 -0.043 -0.034
Convolvulaceae -0.044 0.113 -0.03 -0.044 -0.053 1 -0.077 0.001
Commelinaceae -0.035 -0.01 0.291 -0.035 -0.043 -0.077 1 0.531
Oxalidaceae -0.028 0.166 -0.05 -0.028 -0.034 0.001 0.531 1
n=50.
At the 99 % confidence level, no significant negative association is found between grass families. Family
pairs with positive associations (mutualism, commensalism, proto-cooperation, complementary
resource-partitioning, or they require the similar environmental conditions (nich overlap)) include
Euphorbiaceae-: Leguminosae (0.393), Onagraceae (0.555), Amaranthaceae (0.784), Convolvulaceae

(0.378);
Leguminosae-Convolvulaceae (0.485);
Malvaceae-: Asteraceae (0.423); Onagraceae (0.573), Convolvulaceae (0.396);
Onagraceae-Convolvulaceae (0.723).
Interspecific Pearson correlation coefficients can be found in Table 4.
Table 4 Interspecfic Pearson correlation coefficients.

A. D. D. S. L. A. C. C. C.
E. hirta S. sebiferum P. vittata
scarabaeoides gangeticum triflorum cannabina japonicum compressus aciculatus citratus dactylon
E. hirta 1 -0.023 0.723 -0.023 -0.056 -0.024 0.144 -0.025 0.02 -0.034 -0.023 -0.035
S. sebiferum -0.023 1 -0.07 -0.02 0.503 -0.021 -0.02 -0.021 0.137 -0.03 -0.02 -0.03
A. scarabaeoides 0.723 -0.07 1 -0.07 -0.144 -0.072 0.024 -0.075 -0.182 -0.1 -0.069 0.141
D. gangeticum -0.023 -0.02 -0.07 1 -0.049 -0.021 -0.02 0.056 0.056 -0.03 -0.02 -0.03
D. triflorum -0.056 0.503 -0.144 -0.049 1 -0.051 -0.049 -0.052 0.065 -0.072 -0.049 -0.074
S. cannabina -0.024 -0.021 -0.072 -0.021 -0.051 1 -0.021 -0.023 -0.064 -0.031 -0.021 -0.032
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P. vittata 0.144 -0.02 0.024 -0.02 -0.049 -0.021 1 -0.021 0.343 -0.03 -0.02 -0.03
L. japonicum -0.025 -0.021 -0.075 0.056 -0.052 -0.023 -0.021 1 -0.081 -0.032 -0.021 -0.033
A. compressus 0.02 0.137 -0.182 0.056 0.065 -0.064 0.343 -0.081 1 -0.108 0.098 -0.128
C. aciculatus -0.034 -0.03 -0.1 -0.03 -0.072 -0.031 -0.03 -0.032 -0.108 1 0.003 -0.045
C. citratus -0.023 -0.02 -0.069 -0.02 -0.049 -0.021 -0.02 -0.021 0.098 0.003 1 -0.03
C. dactylon -0.035 -0.03 0.141 -0.03 -0.074 -0.032 -0.03 -0.033 -0.128 -0.045 -0.03 1
E. crusgalli -0.033 -0.029 -0.1 -0.029 -0.07 -0.03 -0.029 -0.031 0 -0.042 -0.029 -0.044
E. minor -0.07 -0.06 -0.181 0.767 0.124 -0.063 -0.06 -0.001 0.013 -0.089 -0.06 -0.091
E. pilosa -0.04 -0.034 -0.104 -0.034 -0.083 -0.036 -0.034 -0.037 -0.134 -0.051 -0.034 -0.052
I. aristatum -0.062 -0.054 -0.17 -0.054 0.026 -0.056 -0.054 0.59 -0.159 -0.079 -0.054 -0.081
L. gracile -0.03 -0.026 0.031 -0.026 -0.064 -0.028 -0.026 -0.028 -0.113 -0.039 -0.026 -0.04
P. repens -0.066 0.012 -0.212 -0.107 -0.074 0.125 -0.082 -0.105 -0.13 -0.079 0.029 -0.158
P. scrobiculatum -0.041 -0.061 -0.012 -0.01 0.221 -0.064 0.118 -0.062 0.105 -0.091 -0.061 -0.055
R. repens 0.581 -0.058 0.665 -0.058 -0.007 -0.06 -0.058 -0.062 -0.21 -0.085 -0.058 0.074
C. roseus -0.023 -0.02 -0.07 -0.02 -0.049 -0.021 -0.02 -0.021 0 -0.03 -0.02 -0.03
S. rhombifolia -0.028 -0.024 0.079 -0.024 -0.059 -0.026 -0.024 -0.026 -0.105 -0.036 -0.024 -0.019
U. sp. -0.023 -0.02 0.002 -0.02 -0.049 -0.021 -0.02 -0.021 -0.086 -0.03 -0.02 -0.03
U. lobata 0.918 0.011 0.652 -0.028 -0.047 -0.03 -0.028 -0.031 -0.066 -0.042 -0.028 -0.043
A. conyzoides -0.071 -0.062 0.173 -0.062 -0.149 -0.065 -0.062 -0.066 -0.197 -0.091 -0.062 0.053
A. subulatus -0.029 -0.025 -0.084 -0.025 -0.06 0.029 -0.025 -0.027 -0.106 -0.037 -0.025 -0.038
B. bipinnata -0.043 -0.037 0.183 -0.037 -0.09 -0.039 -0.037 -0.04 -0.148 -0.055 -0.037 -0.014
B. pilosa 0.247 0.482 0.272 -0.033 0.198 -0.035 -0.033 -0.036 0.064 -0.049 -0.033 -0.051
E. sonchifolia -0.12 -0.096 -0.12 -0.062 -0.183 -0.109 -0.104 -0.024 -0.142 0.039 -0.104 0.228
E. acer 0.071 0.333 -0.023 -0.046 0.093 0.458 0.714 -0.05 0.25 -0.068 -0.046 -0.07
E. chinense -0.055 -0.059 -0.002 -0.047 -0.139 0.05 0.023 -0.037 0.109 -0.087 -0.059 -0.075
M. micrantha -0.032 -0.028 -0.096 -0.028 -0.067 -0.029 -0.028 -0.03 0.066 -0.041 -0.028 -0.042
P. indica -0.023 -0.02 -0.052 -0.02 -0.049 -0.021 -0.02 -0.021 -0.086 -0.03 -0.02 -0.03
V. cinerea -0.056 0.567 -0.066 -0.049 0.215 -0.051 -0.049 -0.053 -0.008 -0.072 -0.049 -0.051
W. triloba -0.049 0.526 -0.148 -0.042 0.204 -0.018 -0.042 -0.046 0.319 -0.054 0.031 -0.065
J. linifolia 0.983 -0.021 0.721 -0.021 -0.052 -0.022 -0.021 -0.023 -0.042 -0.032 -0.021 -0.033
P. triquetrum -0.023 -0.02 -0.07 -0.02 -0.049 -0.021 -0.02 -0.021 0.085 -0.03 -0.02 -0.03
C. asiatica -0.051 -0.044 -0.117 -0.044 -0.042 -0.047 -0.044 -0.048 -0.183 0.033 -0.044 -0.067
C. exaltatus -0.023 -0.02 -0.07 -0.02 -0.049 -0.021 -0.02 -0.021 -0.086 -0.03 -0.02 -0.03
C. compressus -0.023 -0.02 -0.07 -0.02 -0.049 -0.021 -0.02 -0.021 0.152 -0.013 -0.02 -0.03
K. brevifolia -0.032 -0.028 -0.097 -0.028 -0.067 -0.029 -0.028 -0.03 -0.113 -0.041 -0.028 -0.042
L. cernuum -0.023 -0.02 -0.07 0.999 -0.049 -0.021 -0.02 0.056 0.056 -0.03 -0.02 -0.03
A. philoxeroides -0.023 -0.02 -0.024 -0.02 0.464 -0.021 -0.02 -0.021 -0.064 -0.03 -0.02 -0.03
A. sessilis -0.023 0.999 -0.07 -0.02 0.503 -0.021 -0.02 -0.021 0.137 -0.03 -0.02 -0.03
I. triloba -0.023 -0.02 -0.052 -0.02 -0.049 -0.021 -0.02 -0.021 -0.086 -0.03 -0.02 -0.03
P. purpurea 0.743 -0.043 0.515 -0.043 -0.104 -0.045 -0.043 -0.046 -0.132 0.26 -0.043 0.207
Z. pendula -0.041 -0.035 -0.123 -0.035 -0.085 -0.037 -0.035 -0.038 -0.121 -0.052 -0.035 -0.054
O. corniculata -0.032 -0.028 -0.095 -0.028 -0.068 -0.029 -0.028 -0.03 -0.089 0.005 -0.028 -0.042
n=50.
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Table 4 Interspecfic Pearson correlation coefficients (Continue).

U.
E. crusgalli E. minor E. pilosa I. aristatum L. gracile P. repens P. scrobiculatum R. repens C. roseus S. rhombifolia U. sp.
lobata
E. hirta -0.033 -0.07 -0.04 -0.062 -0.03 -0.066 -0.041 0.581 -0.023 -0.028 -0.023 0.918
S. sebiferum -0.029 -0.06 -0.034 -0.054 -0.026 0.012 -0.061 -0.058 -0.02 -0.024 -0.02 0.011
A. scarabaeoides -0.1 -0.181 -0.104 -0.17 0.031 -0.212 -0.012 0.665 -0.07 0.079 0.002 0.652
D. gangeticum -0.029 0.767 -0.034 -0.054 -0.026 -0.107 -0.01 -0.058 -0.02 -0.024 -0.02 -0.028
D. triflorum -0.07 0.124 -0.083 0.026 -0.064 -0.074 0.221 -0.007 -0.049 -0.059 -0.049 -0.047
S. cannabina -0.03 -0.063 -0.036 -0.056 -0.028 0.125 -0.064 -0.06 -0.021 -0.026 -0.021 -0.-03
P. vittata -0.029 -0.06 -0.034 -0.054 -0.026 -0.082 0.118 -0.058 -0.02 -0.024 -0.02 -0.028
L. japonicum -0.031 -0.001 -0.037 0.59 -0.028 -0.105 -0.062 -0.062 -0.021 -0.026 -0.021 -0.031
A. compressus 0 0.013 -0.134 -0.159 -0.113 -0.13 0.105 -0.21 0 -0.105 -0.086 -0.066
C. aciculatus -0.042 -0.089 -0.051 -0.079 -0.039 -0.079 -0.091 -0.085 -0.03 -0.036 -0.03 -0.042
C. citratus -0.029 -0.06 -0.034 -0.054 -0.026 0.029 -0.061 -0.058 -0.02 -0.024 -0.02 -0.028
C. dactylon -0.044 -0.091 -0.052 -0.081 -0.04 -0.158 -0.055 0.074 -0.03 -0.019 -0.03 -0.043
E. crusgalli 1 -0.086 0.063 0.163 -0.038 0.181 -0.088 -0.082 -0.029 -0.035 -0.029 -0.041
E. minor -0.086 1 -0.103 -0.161 -0.079 -0.305 -0.046 0.054 -0.06 -0.073 -0.06 -0.085
E. pilosa 0.063 -0.103 1 0.054 -0.045 0.168 0.07 -0.099 -0.034 0.15 -0.034 -0.036
I. aristatum 0.163 -0.161 0.054 1 -0.07 0.119 -0.114 -0.153 -0.054 -0.056 -0.054 -0.067
L. gracile -0.038 -0.079 -0.045 -0.07 1 -0.125 -0.081 0.03 -0.026 -0.032 0.927 -0.037
P. repens 0.181 -0.305 0.168 0.119 -0.125 1 -0.187 -0.223 -0.033 -0.065 -0.107 0.068
P. scrobiculatum -0.088 -0.046 0.07 -0.114 -0.081 -0.187 1 -0.118 0.041 -0.025 -0.061 -0.079
R. repens -0.082 0.054 -0.099 -0.153 0.03 -0.223 -0.118 1 -0.058 -0.056 0.055 0.535
C. roseus -0.029 -0.06 -0.034 -0.054 -0.026 -0.033 0.041 -0.058 1 -0.024 -0.02 -0.028
S. rhombifolia -0.035 -0.073 0.15 -0.056 -0.032 -0.065 -0.025 -0.056 -0.024 1 -0.024 -0.035
U. sp. -0.029 -0.06 -0.034 -0.054 0.927 -0.107 -0.061 0.055 -0.02 -0.024 1 -0.028
U. lobata -0.041 -0.085 -0.036 -0.067 -0.037 0.068 -0.079 0.535 -0.028 -0.035 -0.028 1
A. conyzoides 0.19 -0.184 0.166 -0.058 0.131 -0.067 -0.109 -0.126 0.198 0.657 -0.062 -0.079
A. subulatus -0.035 -0.074 0.043 -0.052 -0.032 0.339 -0.048 -0.071 -0.025 -0.008 -0.025 0.33
B. bipinnata -0.053 -0.111 -0.064 -0.099 0.906 -0.143 -0.114 0 -0.037 -0.037 0.813 -0.053
B. pilosa -0.048 -0.099 -0.057 -0.089 -0.044 -0.052 -0.102 0.095 -0.033 -0.041 -0.033 0.245
E. sonchifolia 0.152 -0.237 0.175 0.332 -0.136 0.168 0.138 -0.206 0.401 -0.045 -0.104 -0.072
E. acer -0.066 -0.138 -0.079 -0.123 -0.061 0.13 0.008 -0.115 -0.046 -0.056 -0.046 -0.051
E. chinense -0.085 -0.11 -0.088 -0.114 -0.078 0.193 0.046 0.045 -0.059 -0.073 -0.059 0.015
M. micrantha 0.717 -0.084 0.092 -0.012 -0.037 0.11 -0.067 -0.08 -0.028 -0.022 -0.028 -0.039
P. indica -0.029 -0.06 0.757 -0.015 -0.026 0.005 0.144 -0.058 -0.02 0.223 -0.02 -0.028
V. cinerea -0.07 -0.146 0.395 -0.06 -0.064 0.024 0.162 -0.118 0.567 0.015 -0.049 -0.021
W. triloba -0.061 -0.112 -0.073 -0.114 -0.056 0.074 -0.13 -0.122 -0.042 -0.052 -0.042 -0.038
J. linifolia -0.031 -0.064 -0.037 -0.057 -0.028 -0.029 -0.066 0.592 -0.021 -0.026 -0.021 0.955
P. triquetrum -0.029 -0.023 -0.034 -0.054 -0.026 -0.107 0.247 -0.058 -0.02 -0.024 -0.02 -0.028
C. asiatica -0.064 -0.133 0.286 0.004 -0.058 0.364 0.019 -0.108 -0.044 0.04 -0.044 0.265
C. exaltatus 0.699 -0.06 0.123 0.27 -0.026 0.224 -0.061 -0.058 -0.02 -0.024 -0.02 -0.028
C. compressus -0.029 -0.06 -0.034 -0.054 -0.026 0.104 -0.061 -0.058 -0.02 -0.024 -0.02 -0.028
K. brevifolia -0.04 -0.07 -0.048 0.121 -0.036 0.342 -0.085 -0.08 -0.028 -0.034 -0.028 -0.039
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L. cernuum -0.029 0.767 -0.034 -0.054 -0.026 -0.107 -0.01 -0.058 -0.02 -0.024 -0.02 -0.028
A. philoxeroides -0.029 -0.06 -0.034 0.257 -0.026 0.122 -0.061 -0.058 -0.02 -0.024 -0.02 -0.028
A. sessilis -0.029 -0.06 -0.034 -0.054 -0.026 0.012 -0.061 -0.058 -0.02 -0.024 -0.02 0.011
I. triloba -0.029 -0.06 0.757 -0.015 -0.026 0.005 0.144 -0.058 -0.02 0.223 -0.02 -0.028
P. purpurea -0.062 -0.126 0.21 -0.089 -0.056 -0.079 -0.05 0.427 -0.043 0.03 -0.043 0.702
Z. pendula 0.331 -0.106 0.466 0.232 -0.046 0.287 -0.027 -0.101 -0.035 -0.043 -0.035 -0.05
O. corniculata -0.04 -0.084 0.049 0.15 -0.037 0.065 0.09 -0.08 -0.028 -0.034 -0.028 -0.018
n=50.

M.
A. conyzoides A. subulatus B. bipinnata B. pilosa E. sonchifolia E. acer E. chinense P. indica V. cinerea W. triloba J. linifolia
micrantha
E. hirta -0.071 -0.029 -0.043 0.247 -0.12 0.071 -0.055 -0.032 -0.023 -0.056 -0.049 0.983
S. sebiferum -0.062 -0.025 -0.037 0.482 -0.096 0.333 -0.059 -0.028 -0.02 0.567 0.526 -0.021
A. scarabaeoides 0.173 -0.084 0.183 0.272 -0.12 -0.023 -0.002 -0.096 -0.052 -0.066 -0.148 0.721
D. gangeticum -0.062 -0.025 -0.037 -0.033 -0.062 -0.046 -0.047 -0.028 -0.02 -0.049 -0.042 -0.021
D. triflorum -0.149 -0.06 -0.09 0.198 -0.183 0.093 -0.139 -0.067 -0.049 0.215 0.204 -0.052
S. cannabina -0.065 0.029 -0.039 -0.035 -0.109 0.458 0.05 -0.029 -0.021 -0.051 -0.018 -0.022
P. vittata -0.062 -0.025 -0.037 -0.033 -0.104 0.714 0.023 -0.028 -0.02 -0.049 -0.042 -0.021
L. japonicum -0.066 -0.027 -0.04 -0.036 -0.024 -0.05 -0.037 -0.03 -0.021 -0.053 -0.046 -0.023
A. compressus -0.197 -0.106 -0.148 0.064 -0.142 0.25 0.109 0.066 -0.086 -0.008 0.319 -0.042
C. aciculatus -0.091 -0.037 -0.055 -0.049 0.039 -0.068 -0.087 -0.041 -0.03 -0.072 -0.054 -0.032
C. citratus -0.062 -0.025 -0.037 -0.033 -0.104 -0.046 -0.059 -0.028 -0.02 -0.049 0.031 -0.021
C. dactylon 0.053 -0.038 -0.014 -0.051 0.228 -0.07 -0.075 -0.042 -0.03 -0.051 -0.065 -0.033
E. crusgalli 0.19 -0.035 -0.053 -0.048 0.152 -0.066 -0.085 0.717 -0.029 -0.07 -0.061 -0.031
E. minor -0.184 -0.074 -0.111 -0.099 -0.237 -0.138 -0.11 -0.084 -0.06 -0.146 -0.112 -0.064
E. pilosa 0.166 0.043 -0.064 -0.057 0.175 -0.079 -0.088 0.092 0.757 0.395 -0.073 -0.037
I. aristatum -0.058 -0.052 -0.099 -0.089 0.332 -0.123 -0.114 -0.012 -0.015 -0.06 -0.114 -0.057
L. gracile 0.131 -0.032 0.906 -0.044 -0.136 -0.061 -0.078 -0.037 -0.026 -0.064 -0.056 -0.028
P. repens -0.067 0.339 -0.143 -0.052 0.168 0.13 0.193 0.11 0.005 0.024 0.074 -0.029
P. scrobiculatum -0.109 -0.048 -0.114 -0.102 0.138 0.008 0.046 -0.067 0.144 0.162 -0.13 -0.066
R. repens -0.126 -0.071 0 0.095 -0.206 -0.115 0.045 -0.08 -0.058 -0.118 -0.122 0.592
C. roseus 0.198 -0.025 -0.037 -0.033 0.401 -0.046 -0.059 -0.028 -0.02 0.567 -0.042 -0.021
S. rhombifolia 0.657 -0.008 -0.037 -0.041 -0.045 -0.056 -0.073 -0.022 0.223 0.015 -0.052 -0.026
U. sp. -0.062 -0.025 0.813 -0.033 -0.104 -0.046 -0.059 -0.028 -0.02 -0.049 -0.042 -0.021
U. lobata -0.079 0.33 -0.053 0.245 -0.072 -0.051 0.015 -0.039 -0.028 -0.021 -0.038 0.955
A. conyzoides 1 -0.051 0.229 -0.102 0.084 -0.141 -0.16 0.144 0.103 0.136 -0.13 -0.066
A. subulatus -0.051 1 -0.046 -0.041 0.072 -0.029 0.204 -0.03 0.067 -0.004 -0.052 0.045
B. bipinnata 0.229 -0.046 1 -0.062 -0.094 -0.086 -0.11 -0.052 -0.037 -0.09 -0.079 -0.04
B. pilosa -0.102 -0.041 -0.062 1 -0.168 0.332 -0.022 -0.046 -0.033 0.23 0.216 0.253
E. sonchifolia 0.084 0.072 -0.094 -0.168 1 -0.213 0.065 0.218 -0.104 0.313 -0.215 -0.097
E. acer -0.141 -0.029 -0.086 0.332 -0.213 1 0.082 -0.064 -0.046 0.117 0.123 -0.049
E. chinense -0.16 0.204 -0.11 -0.022 0.065 0.082 1 -0.07 -0.059 -0.08 -0.105 -0.045
M. micrantha 0.144 -0.03 -0.052 -0.046 0.218 -0.064 -0.07 1 0.022 -0.007 -0.059 -0.03
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P. indica 0.103 0.067 -0.037 -0.033 -0.104 -0.046 -0.059 0.022 1 0.259 -0.042 -0.021
V. cinerea 0.136 -0.004 -0.09 0.23 0.313 0.117 -0.08 -0.007 0.259 1 0.24 -0.052
W. triloba -0.13 -0.052 -0.079 0.216 -0.215 0.123 -0.105 -0.059 -0.042 0.24 1 -0.046
J. linifolia -0.066 0.045 -0.04 0.253 -0.097 -0.049 -0.045 -0.03 -0.021 -0.052 -0.046 1
P. triquetrum -0.062 -0.025 -0.037 -0.033 0.089 -0.046 -0.034 -0.028 -0.02 -0.049 -0.042 -0.021
C. asiatica -0.026 0.855 -0.082 -0.074 0.055 -0.056 0.215 -0.036 0.344 0.126 -0.094 0.012
C. exaltatus 0.139 -0.025 -0.037 -0.033 0.064 -0.046 -0.059 0.022 -0.02 -0.049 -0.042 -0.021
C. compressus -0.062 -0.025 -0.037 -0.033 -0.104 -0.021 -0.001 -0.028 -0.02 -0.049 0.328 -0.021
K. brevifolia -0.085 -0.034 -0.052 -0.046 0.001 -0.064 -0.083 -0.039 -0.028 -0.068 0.251 -0.03
L. cernuum -0.062 -0.025 -0.037 -0.033 -0.062 -0.046 -0.047 -0.028 -0.02 -0.049 -0.042 -0.021
A. philoxeroides -0.062 -0.025 -0.037 -0.033 -0.02 -0.046 -0.059 -0.028 -0.02 -0.049 -0.042 -0.021
A. sessilis -0.062 -0.025 -0.037 0.482 -0.096 0.333 -0.059 -0.028 -0.02 0.567 0.526 -0.021
I. triloba 0.103 0.067 -0.037 -0.033 -0.104 -0.046 -0.059 0.022 1 0.259 -0.042 -0.021
P. purpurea -0.064 -0.012 -0.08 0.155 -0.074 -0.099 -0.115 -0.043 0.3 0.033 -0.011 0.754
Z. pendula 0.129 -0.043 -0.065 -0.059 0.392 -0.081 -0.084 0.109 -0.035 0.129 -0.075 -0.038
O. corniculata 0.001 -0.01 -0.052 -0.047 0.171 -0.064 -0.022 0.022 -0.028 0.039 -0.059 -0.03
n=50.

C. C. K. L. A. A. P. Z. O.
P. triquetrum C. asiatica I. triloba
exaltatus compressus brevifolia cernuum philoxeroides sessilis purpurea pendula corniculata
E. hirta -0.023 -0.051 -0.023 -0.023 -0.032 -0.023 -0.023 -0.023 -0.023 0.743 -0.041 -0.032
S. sebiferum -0.02 -0.044 -0.02 -0.02 -0.028 -0.02 -0.02 0.999 -0.02 -0.043 -0.035 -0.028
A. scarabaeoides -0.07 -0.117 -0.07 -0.07 -0.097 -0.07 -0.024 -0.07 -0.052 0.515 -0.123 -0.095
D. gangeticum -0.02 -0.044 -0.02 -0.02 -0.028 0.999 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
D. triflorum -0.049 -0.042 -0.049 -0.049 -0.067 -0.049 0.464 0.503 -0.049 -0.104 -0.085 -0.068
S. cannabina -0.021 -0.047 -0.021 -0.021 -0.029 -0.021 -0.021 -0.021 -0.021 -0.045 -0.037 -0.029
P. vittata -0.02 -0.044 -0.02 -0.02 -0.028 -0.02 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
L. japonicum -0.021 -0.048 -0.021 -0.021 -0.03 0.056 -0.021 -0.021 -0.021 -0.046 -0.038 -0.03
A. compressus 0.085 -0.183 -0.086 0.152 -0.113 0.056 -0.064 0.137 -0.086 -0.132 -0.121 -0.089
C. aciculatus -0.03 0.033 -0.03 -0.013 -0.041 -0.03 -0.03 -0.03 -0.03 0.26 -0.052 0.005
C. citratus -0.02 -0.044 -0.02 -0.02 -0.028 -0.02 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
C. dactylon -0.03 -0.067 -0.03 -0.03 -0.042 -0.03 -0.03 -0.03 -0.03 0.207 -0.054 -0.042
E. crusgalli -0.029 -0.064 0.699 -0.029 -0.04 -0.029 -0.029 -0.029 -0.029 -0.062 0.331 -0.04
E. minor -0.023 -0.133 -0.06 -0.06 -0.07 0.767 -0.06 -0.06 -0.06 -0.126 -0.106 -0.084
E. pilosa -0.034 0.286 0.123 -0.034 -0.048 -0.034 -0.034 -0.034 0.757 0.21 0.466 0.049
I. aristatum -0.054 0.004 0.27 -0.054 0.121 -0.054 0.257 -0.054 -0.015 -0.089 0.232 0.15
L. gracile -0.026 -0.058 -0.026 -0.026 -0.036 -0.026 -0.026 -0.026 -0.026 -0.056 -0.046 -0.037
P. repens -0.107 0.364 0.224 0.104 0.342 -0.107 0.122 0.012 0.005 -0.079 0.287 0.065
P. scrobiculatum 0.247 0.019 -0.061 -0.061 -0.085 -0.01 -0.061 -0.061 0.144 -0.05 -0.027 0.09
R. repens -0.058 -0.108 -0.058 -0.058 -0.08 -0.058 -0.058 -0.058 -0.058 0.427 -0.101 -0.08
C. roseus -0.02 -0.044 -0.02 -0.02 -0.028 -0.02 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
S. rhombifolia -0.024 0.04 -0.024 -0.024 -0.034 -0.024 -0.024 -0.024 0.223 0.03 -0.043 -0.034
U. sp. -0.02 -0.044 -0.02 -0.02 -0.028 -0.02 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
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U. lobata -0.028 0.265 -0.028 -0.028 -0.039 -0.028 -0.028 0.011 -0.028 0.702 -0.05 -0.018
A. conyzoides -0.062 -0.026 0.139 -0.062 -0.085 -0.062 -0.062 -0.062 0.103 -0.064 0.129 0.001
A. subulatus -0.025 0.855 -0.025 -0.025 -0.034 -0.025 -0.025 -0.025 0.067 -0.012 -0.043 -0.01
B. bipinnata -0.037 -0.082 -0.037 -0.037 -0.052 -0.037 -0.037 -0.037 -0.037 -0.08 -0.065 -0.052
B. pilosa -0.033 -0.074 -0.033 -0.033 -0.046 -0.033 -0.033 0.482 -0.033 0.155 -0.059 -0.047
E. sonchifolia 0.089 0.055 0.064 -0.104 0.001 -0.062 -0.02 -0.096 -0.104 -0.074 0.392 0.171
E. acer -0.046 -0.056 -0.046 -0.021 -0.064 -0.046 -0.046 0.333 -0.046 -0.099 -0.081 -0.064
E. chinense -0.034 0.215 -0.059 -0.001 -0.083 -0.047 -0.059 -0.059 -0.059 -0.115 -0.084 -0.022
M. micrantha -0.028 -0.036 0.022 -0.028 -0.039 -0.028 -0.028 -0.028 0.022 -0.043 0.109 0.022
P. indica -0.02 0.344 -0.02 -0.02 -0.028 -0.02 -0.02 -0.02 1 0.3 -0.035 -0.028
V. cinerea -0.049 0.126 -0.049 -0.049 -0.068 -0.049 -0.049 0.567 0.259 0.033 0.129 0.039
W. triloba -0.042 -0.094 -0.042 0.328 0.251 -0.042 -0.042 0.526 -0.042 -0.011 -0.075 -0.059
J. linifolia -0.021 0.012 -0.021 -0.021 -0.03 -0.021 -0.021 -0.021 -0.021 0.754 -0.038 -0.03
P. triquetrum 1 -0.044 -0.02 -0.02 -0.028 -0.02 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
C. asiatica -0.044 1 -0.044 -0.044 -0.062 -0.044 0.084 -0.044 0.344 0.079 -0.01 0.166
C. exaltatus -0.02 -0.044 1 -0.02 -0.028 -0.02 -0.02 -0.02 -0.02 -0.043 0.499 -0.028
C. compressus -0.02 -0.044 -0.02 1 -0.028 -0.02 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
K. brevifolia -0.028 -0.062 -0.028 -0.028 1 -0.028 -0.028 -0.028 -0.028 0.005 -0.049 -0.039
L. cernuum -0.02 -0.044 -0.02 -0.02 -0.028 1 -0.02 -0.02 -0.02 -0.043 -0.035 -0.028
A. philoxeroides -0.02 0.084 -0.02 -0.02 -0.028 -0.02 1 -0.02 -0.02 -0.043 -0.035 -0.028
A. sessilis -0.02 -0.044 -0.02 -0.02 -0.028 -0.02 -0.02 1 -0.02 -0.043 -0.035 -0.028
I. triloba -0.02 0.344 -0.02 -0.02 -0.028 -0.02 -0.02 -0.02 1 0.3 -0.035 -0.028
P. purpurea -0.043 0.079 -0.043 -0.043 0.005 -0.043 -0.043 -0.043 0.3 1 -0.076 0.004
Z. pendula -0.035 -0.01 0.499 -0.035 -0.049 -0.035 -0.035 -0.035 -0.035 -0.076 1 0.531
O. corniculata -0.028 0.166 -0.028 -0.028 -0.039 -0.028 -0.028 -0.028 -0.028 0.004 0.531 1
n=50.
It can be found from Table 4 that at the 99 % confidence level, all significant grass species pairs are
positive associated. They are
E. hirta-: A. scarabaeoides (0.723), R. repens (0.581), U. lobata (0.918), J. linifolia (0.983);

S. sebiferum-: D. triflorum (0.503), B. pilosa (0.482), V. cinerea (0.567), W. triloba (0.526), A. sessilis
(0.999);
A. scarabaeoides-: R. repens (0.665), U. lobata (0.652), J. linifolia (0.721), P. purpurea (0.515);
D. gangeticum-: E. minor (0.767), L. cernuum (0.999);
D. triflorum-: A. philoxeroides (0.464), A. sessilis (0.503);
S. cannabina-E. acer (0.458)
P. vittata-E. acer (0.714)
L. japonicum-I. aristatum (0.59)
E. crusgalli-: M.micrantha (0.717), C. exaltatus (0.699);
E. minor-L. cernuum (0.767)
E. pilosa-: P. indica (0.757), V. cinerea (0.395), I. triloba (0.757), Z. pendula (0.466);
L. gracile-: U. sp. (0.927), B. bipinnata (0.906);
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P. repens-C. asiatica (0.364)

R. repens-: U. lobata (0.535), J. linifolia (0.592), P. purpurea (0.427);
C. roseus-: E. sonchifolia (0.401), V. cinerea (0.567);
S. rhombifolia-A. conyzoides (0.657);
U. sp.-B. bipinnata (0.813);
U. lobata-: J. linifolia (0.955), P. purpurea (0.702);
A. subulatus-C. asiatica (0.855);
B. pilosa-A. sessilis (0.482);
E. sonchifolia-Z. pendula (0.392)
V. cinerea-A. sessilis (0.567);
W. triloba-A. sessilis (0.526);
J. linifolia-P. purpurea (0.754);
C. exaltatus-Z. pendula (0.499);
Z. pendula-O. corniculata (0.531)
3.2 Associations measured by net correlation coefficient

Inter-family net correlation coefficients are indicated in Table 5.
Table 5 Inter-family net correlation coefficients.

Euphorbiaceae 1 -0.195 0.491 0.071 -0.213 -0.051 -0.098 0.19 0.822
Leguminosae -0.195 1 0.164 0.002 -0.347 -0.216 -0.259 0.395 0.541
Pteridaceae 0.491 0.164 1 -0.08 0.186 0.023 0.067 -0.151 -0.426
Lygodiaceae 0.071 0.002 -0.08 1 0.217 -0.002 0.028 -0.046 -0.055
Gramineae -0.213 -0.347 0.186 0.217 1 -0.149 -0.089 -0.081 0.311
Apocynaceae -0.051 -0.216 0.023 -0.002 -0.149 1 -0.125 0.137 0.142
Malvaceae -0.098 -0.259 0.067 0.028 -0.089 -0.125 1 0.553 0.373
Asteraceae 0.19 0.395 -0.151 -0.046 -0.081 0.137 0.553 1 -0.364
Onagraceae 0.822 0.541 -0.426 -0.055 0.311 0.142 0.373 -0.364 1
Connaraceae -0.009 -0.131 -0.01 -0.015 -0.151 -0.059 -0.01 -0.066 0.063
Umbelliferae -0.259 -0.194 0.091 -0.07 0.111 -0.059 0.067 0.148 0.207
Cyperaceae 0.086 -0.003 -0.097 -0.12 0.298 -0.002 0.016 -0.015 -0.085
Lycopodiaceae 0.044 0.022 -0.055 0.022 0.039 -0.011 0.037 -0.104 -0.041
Amaranthaceae 0.972 0.168 -0.482 -0.08 0.227 0.037 0.07 -0.142 -0.786
Convolvulaceae 0.12 -0.001 -0.101 -0.063 0.016 -0.031 0.014 -0.121 0.171
Commelinaceae 0.001 -0.069 -0.023 -0.058 0.16 -0.026 -0.073 0.14 0.057
Oxalidaceae 0.063 -0.017 -0.037 -0.005 -0.072 -0.013 0.01 -0.094 -0.06
n=50.
Table 5 Inter-family net correlation coefficients (Continue).

Euphorbiaceae -0.009 -0.259 0.086 0.044 0.972 0.12 0.001 0.063
Leguminosae -0.131 -0.194 -0.003 0.022 0.168 -0.001 -0.069 -0.017
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Pteridaceae -0.01 0.091 -0.097 -0.055 -0.482 -0.101 -0.023 -0.037
Lygodiaceae -0.015 -0.07 -0.12 0.022 -0.08 -0.063 -0.058 -0.005
Gramineae -0.151 0.111 0.298 0.039 0.227 0.016 0.16 -0.072
Apocynaceae -0.059 -0.059 -0.002 -0.011 0.037 -0.031 -0.026 -0.013
Malvaceae -0.01 0.067 0.016 0.037 0.07 0.014 -0.073 0.01
Asteraceae -0.066 0.148 -0.015 -0.104 -0.142 -0.121 0.14 -0.094
Onagraceae 0.063 0.207 -0.085 -0.041 -0.786 0.171 0.057 -0.06
Connaraceae 1 -0.031 -0.01 -0.033 0.004 -0.057 0 -0.031
Umbelliferae -0.031 1 -0.085 -0.026 0.239 0.151 -0.128 0.193
Cyperaceae -0.01 -0.085 1 -0.056 -0.099 0.008 0.28 -0.22
Lycopodiaceae -0.033 -0.026 -0.056 1 -0.047 -0.058 -0.016 -0.019
Amaranthaceae 0.004 0.239 -0.099 -0.047 1 -0.126 -0.016 -0.061
Convolvulaceae -0.057 0.151 0.008 -0.058 -0.126 1 -0.094 0.039
Commelinaceae 0 -0.128 0.28 -0.016 -0.016 -0.094 1 0.576
Oxalidaceae -0.031 0.193 -0.22 -0.019 -0.061 0.039 0.576 1
n=50.
Table 5 showed that there are significant direct negative associations (competition, interference, or they
require the distinct environmental conditions) for family pairs Pteridaceae-Amaranthaceae (-0.48), and
Onagraceae-Amaranthaceae (-0.79). Family pairs with significant direct positive associations include:
Euphorbiaceae-Pteridaceae (0.491), Euphorbiaceae-Onagraceae (0.822), Leguminosae-Onagraceae (0.541),
Malvaceae-Asteraceae (0.553), and Commelinaceae-Oxalidaceae (0.576).
Interspecific net correlation coefficients are shown in Table 6.
Table 6 Interspecific net correlation coefficients.

A. D. D. S. P. L. A. C. C. C.
E. hirta S. sebiferum
scarabaeoides gangeticum triflorum cannabina vittata japonicum compressus aciculatus citratus dactylon
E. hirta 1 0 0.072 0 -0.022 0.857 0.999 0.566 -0.141 0.529 0.855 -0.673
S. sebiferum 0 1 0 -0.011 0 0 0 0 0 0 0 0
A. scarabaeoides 0.072 0 1 0 -0.166 -0.513 -0.082 0.247 -0.351 0.335 0.241 0.685
D. gangeticum 0 -0.011 0 1 0 0 0 0 0 0 0 0
D. triflorum -0.022 0 -0.166 0 1 0.016 0.022 -0.188 0.083 0.188 0.05 0.017
S. cannabina 0.857 0 -0.513 0 0.016 1 -0.863 -0.308 -0.049 -0.221 -0.558 0.953
P. vittata 0.999 0 -0.082 0 0.022 -0.863 1 -0.561 0.139 -0.522 -0.85 0.681
L. japonicum 0.566 0 0.247 0 -0.188 -0.308 -0.561 1 0.005 -0.41 -0.671 0.174
A. compressus -0.141 0 -0.351 0 0.083 -0.049 0.139 0.005 1 -0.034 0.094 0.165
C. aciculatus 0.529 0 0.335 0 0.188 -0.221 -0.522 -0.41 -0.034 1 -0.735 0.017
C. citratus 0.855 0 0.241 0 0.05 -0.558 -0.85 -0.671 0.094 -0.735 1 0.345
C. dactylon -0.673 0 0.685 0 0.017 0.953 0.681 0.174 0.165 0.017 0.345 1
E. crusgalli -0.651 0 -0.588 0 -0.029 0.202 0.642 0.671 -0.23 0.744 0.872 0.044
E. minor 0.099 0 -0.555 0 0.303 -0.336 -0.104 0.11 -0.536 -0.171 -0.031 0.396
E. pilosa -0.653 0 0.702 0 -0.011 0.947 0.66 0.125 0.18 0.031 0.322 -0.996
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I. aristatum -0.526 0 -0.398 0 0.261 0.21 0.52 0.883 -0.235 0.301 0.626 -0.065
L. gracile -0.78 0 0.465 0 -0.094 0.955 0.785 0.271 0.131 0.209 0.519 -0.943
P. repens -0.61 0 -0.632 0 -0.012 0.146 0.601 0.658 -0.266 0.716 0.842 0.1
P. scrobiculatum 0.078 0 -0.082 0 0.672 -0.183 -0.081 0.296 -0.234 -0.269 -0.058 0.17
R. repens -0.06 0 0.8 0 0.274 0.396 0.068 -0.253 -0.017 -0.417 -0.253 -0.528
C. roseus -0.691 0 0.56 0 0.075 0.911 0.697 0.361 0.165 0.027 0.408 -0.945
S. rhombifolia -0.654 0 0.672 0 0.004 0.938 0.661 0.184 0.197 0.016 0.334 -0.988
U. sp. 0.651 0 -0.669 0 0.015 -0.937 -0.659 -0.166 -0.201 -0.026 -0.333 0.986
U. lobata 0.805 0 -0.57 0 0.015 -0.993 -0.811 -0.265 -0.101 -0.183 -0.505 0.977
A. conyzoides 0.695 0 -0.627 0 0.024 -0.951 -0.702 -0.221 -0.206 -0.086 -0.396 0.984
A. subulatus -0.916 0 0.412 0 -0.019 0.99 0.92 0.389 0.014 0.32 0.659 -0.911
B. bipinnata -0.268 0 0.899 0 0.153 0.696 0.278 -0.07 0.248 -0.352 -0.099 -0.853
B. pilosa -0.657 0 -0.559 0 -0.002 0.212 0.648 0.67 -0.232 0.751 0.87 0.031
E. sonchifolia 0.625 0 0.268 0 -0.228 -0.344 -0.62 -0.882 0.036 -0.442 -0.743 0.204
E. acer 0.631 0 0.602 0 0.027 -0.174 -0.622 -0.668 0.221 -0.765 -0.864 -0.073
E. chinense 0.458 0 0.637 0 0.113 -0.012 -0.449 -0.438 0.247 -0.794 -0.727 -0.235
M. micrantha 0.623 0 0.612 0 0.032 -0.164 -0.614 -0.658 0.238 -0.744 -0.856 -0.084
P. indica 0 0 0 0 0 0 0 0 0 0 0 0
V. cinerea 0 0 -0.038 0 0.031 0 0 -0.806 0.223 -0.76 0 0.882
W. triloba -0.75 0 0.618 0 -0.003 0.979 0.757 0.213 0.119 0.087 0.425 -0.988
J. linifolia 0.936 0 0.234 0 0.023 -0.624 -0.933 -0.643 0.258 -0.638 -0.91 0.374
P. triquetrum -0.438 0 -0.434 0 -0.081 0.17 0.433 0.526 -0.176 0.33 0.535 -0.036
C. asiatica -0.635 0 -0.501 0 -0.024 0.225 0.627 0.636 -0.2 0.833 0.837 0.006
C. exaltatus -0.43 0 0.824 0 -0.004 0.833 0.44 -0.072 0.234 -0.178 0.057 -0.948
C. compressus 0.942 0 -0.286 0 0.021 -0.942 -0.944 -0.486 -0.011 -0.42 -0.763 0.839
K. brevifolia 0.632 0 0.607 0 0.023 -0.172 -0.623 -0.677 0.223 -0.741 -0.86 -0.078
L. cernuum 0 0.011 0 0.999 0 0 0 0 0 0 0 0
A. philoxeroides 0.633 0 0.6 0 0.131 -0.188 -0.624 -0.687 0.188 -0.78 -0.864 -0.056
A. sessilis 0 0.999 0 0.011 0 0 0 0 0 0 0 0
I. triloba 0 0 0 0 0 0 0 0 0 0 0 0
P. purpurea 0.634 0 -0.716 0 0.008 -0.939 -0.642 -0.109 -0.188 0.002 -0.297 0.995
Z. pendula 0.642 0 -0.71 0 0.011 -0.943 -0.65 -0.113 -0.183 -0.02 -0.308 0.995
O. corniculata -0.642 0 0.709 0 -0.011 0.943 0.65 0.114 0.183 0.02 0.309 -0.995
n=50.
Table 6 Interspecific net correlation coefficients (Continue).

E. I. L. P.
E. crusgalli E. minor P. scrobiculatum R. repens C. roseus S. rhombifolia U. sp. U. lobata
pilosa aristatum gracile repens
E. hirta -0.651 0.099 -0.653 -0.526 -0.78 -0.61 0.078 -0.06 -0.691 -0.654 0.651 0.805
S. sebiferum 0 0 0 0 0 0 0 0 0 0 0 0
A. scarabaeoides -0.588 -0.555 0.702 -0.398 0.465 -0.632 -0.082 0.8 0.56 0.672 -0.669 -0.57
D. gangeticum 0 0 0 0 0 0 0 0 0 0 0 0
D. triflorum -0.029 0.303 -0.011 0.261 -0.094 -0.012 0.672 0.274 0.075 0.004 0.015 0.015
S. cannabina 0.202 -0.336 0.947 0.21 0.955 0.146 -0.183 0.396 0.911 0.938 -0.937 -0.993
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P. vittata 0.642 -0.104 0.66 0.52 0.785 0.601 -0.081 0.068 0.697 0.661 -0.659 -0.811
L. japonicum 0.671 0.11 0.125 0.883 0.271 0.658 0.296 -0.253 0.361 0.184 -0.166 -0.265
A. compressus -0.23 -0.536 0.18 -0.235 0.131 -0.266 -0.234 -0.017 0.165 0.197 -0.201 -0.101
C. aciculatus 0.744 -0.171 0.031 0.301 0.209 0.716 -0.269 -0.417 0.027 0.016 -0.026 -0.183
C. citratus 0.872 -0.031 0.322 0.626 0.519 0.842 -0.058 -0.253 0.408 0.334 -0.333 -0.505
C. dactylon 0.044 0.396 -0.996 -0.065 -0.943 0.1 0.17 -0.528 -0.945 -0.988 0.986 0.977
E. crusgalli 1 -0.275 0.074 -0.722 -0.161 -0.993 -0.104 0.469 -0.05 0.06 -0.063 0.135
E. minor -0.275 1 0.431 -0.428 0.373 -0.341 -0.579 0.189 0.325 0.431 -0.446 -0.367
E. pilosa 0.074 0.431 1 -0.004 -0.94 0.132 0.22 -0.534 -0.926 -0.985 0.985 0.974
I. aristatum -0.722 -0.428 -0.004 1 -0.155 -0.734 -0.502 0.243 -0.268 -0.06 0.037 0.159
L. gracile -0.161 0.373 -0.94 -0.155 1 -0.102 0.277 -0.307 -0.919 -0.95 0.96 0.965
P. repens -0.993 -0.341 0.132 -0.734 -0.102 1 -0.151 0.482 0 0.118 -0.123 0.077
P. scrobiculatum -0.104 -0.579 0.22 -0.502 0.277 -0.151 1 -0.179 0.058 0.196 -0.224 -0.196
R. repens 0.469 0.189 -0.534 0.243 -0.307 0.482 -0.179 1 -0.406 -0.493 0.486 0.435
C. roseus -0.05 0.325 -0.926 -0.268 -0.919 0 0.058 -0.406 1 -0.958 0.949 0.927
S. rhombifolia 0.06 0.431 -0.985 -0.06 -0.95 0.118 0.196 -0.493 -0.958 1 0.994 0.963
U. sp. -0.063 -0.446 0.985 0.037 0.96 -0.123 -0.224 0.486 0.949 0.994 1 -0.962
U. lobata 0.135 -0.367 0.974 0.159 0.965 0.077 -0.196 0.435 0.927 0.963 -0.962 1
A. conyzoides 0.003 -0.449 0.981 0.086 0.968 -0.057 -0.231 0.436 0.959 0.994 -0.994 -0.972
A. subulatus -0.333 0.285 -0.901 -0.303 -0.942 -0.279 0.161 -0.316 -0.883 -0.894 0.892 0.975
B. bipinnata 0.471 0.426 -0.853 0.173 -0.671 0.515 0.036 -0.734 -0.798 -0.858 0.848 0.746
B. pilosa -0.989 -0.282 0.061 -0.727 -0.162 -0.99 -0.136 0.431 -0.06 0.05 -0.055 0.147
E. sonchifolia 0.736 0.128 0.147 0.891 0.308 0.727 0.349 -0.269 0.419 0.215 -0.194 -0.3
E. acer 0.993 0.269 -0.101 0.716 0.134 0.993 0.106 -0.488 0.026 -0.087 0.09 -0.107
E. chinense 0.875 0.167 -0.241 0.451 -0.013 0.877 -0.058 -0.524 -0.193 -0.244 0.238 0.052
M. micrantha 0.999 0.288 -0.113 0.713 0.123 0.994 0.107 -0.487 0.01 -0.1 0.103 -0.095
P. indica 0 0 0 0 0 0 0 0 0 0 0 0
V. cinerea 0.948 -0.101 0.852 0.76 0 0.895 -0.093 0.035 0.914 0.854 -0.851 0
W. triloba -0.049 0.351 -0.985 -0.117 -0.953 0.005 0.174 -0.489 -0.931 -0.972 0.971 0.991
J. linifolia 0.84 0.075 0.347 0.65 0.532 0.817 0.004 -0.171 0.428 0.355 -0.352 -0.547
P. triquetrum -0.624 -0.323 0 -0.625 -0.165 -0.635 -0.092 0.307 -0.16 -0.03 0.021 0.13
C. asiatica -0.923 -0.177 0.026 -0.655 -0.183 -0.917 -0.06 0.427 -0.08 0.018 -0.021 0.169
C. exaltatus 0.35 0.482 -0.96 0.2 -0.837 0.403 0.238 -0.633 -0.854 -0.942 0.943 0.877
C. compressus 0.48 -0.226 0.824 0.409 0.903 0.429 -0.13 0.206 0.834 0.822 -0.82 -0.923
K. brevifolia 0.992 0.284 -0.109 0.73 0.127 0.994 0.117 -0.487 0.021 -0.093 0.097 -0.102
L. cernuum 0 0 0 0 0 0 0 0 0 0 0 0
A. philoxeroides 0.977 0.208 -0.082 0.715 0.159 0.974 0.035 -0.518 0.049 -0.064 0.066 -0.123
A. sessilis 0 0 0 0 0 0 0 0 0 0 0 0
I. triloba 0 0 0 0 0 0 0 0 0 0 0 0
P. purpurea -0.102 -0.436 0.999 -0.012 0.933 -0.16 -0.219 0.545 0.924 0.985 -0.985 -0.968
Z. pendula -0.089 -0.435 0.999 -0.007 0.936 -0.147 -0.222 0.541 0.923 0.984 -0.985 -0.971
O. corniculata 0.088 0.434 -0.999 0.006 -0.936 0.146 0.222 -0.54 -0.923 -0.985 0.985 0.971
n=50.
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A. E. M. P. V. W.
A. conyzoides B. bipinnata B. pilosa E. acer E. chinense J. linifolia
subulatus sonchifolia micrantha indica cinerea triloba
E. hirta 0.695 -0.916 -0.268 -0.657 0.625 0.631 0.458 0.623 0 0 -0.75 0.936
S. sebiferum 0 0 0 0 0 0 0 0 0 0 0 0
A. scarabaeoides -0.627 0.412 0.899 -0.559 0.268 0.602 0.637 0.612 0 -0.038 0.618 0.234
D. gangeticum 0 0 0 0 0 0 0 0 0 0 0 0
D. triflorum 0.024 -0.019 0.153 -0.002 -0.228 0.027 0.113 0.032 0 0.031 -0.003 0.023
S. cannabina -0.951 0.99 0.696 0.212 -0.344 -0.174 -0.012 -0.164 0 0 0.979 -0.624
P. vittata -0.702 0.92 0.278 0.648 -0.62 -0.622 -0.449 -0.614 0 0 0.757 -0.933
L. japonicum -0.221 0.389 -0.07 0.67 -0.882 -0.668 -0.438 -0.658 0 -0.806 0.213 -0.643
A. compressus -0.206 0.014 0.248 -0.232 0.036 0.221 0.247 0.238 0 0.223 0.119 0.258
C. aciculatus -0.086 0.32 -0.352 0.751 -0.442 -0.765 -0.794 -0.744 0 -0.76 0.087 -0.638
C. citratus -0.396 0.659 -0.099 0.87 -0.743 -0.864 -0.727 -0.856 0 0 0.425 -0.91
C. dactylon 0.984 -0.911 -0.853 0.031 0.204 -0.073 -0.235 -0.084 0 0.882 -0.988 0.374
E. crusgalli 0.003 -0.333 0.471 -0.989 0.736 0.993 0.875 0.999 0 0.948 -0.049 0.84
E. minor -0.449 0.285 0.426 -0.282 0.128 0.269 0.167 0.288 0 -0.101 0.351 0.075
E. pilosa 0.981 -0.901 -0.853 0.061 0.147 -0.101 -0.241 -0.113 0 0.852 -0.985 0.347
I. aristatum 0.086 -0.303 0.173 -0.727 0.891 0.716 0.451 0.713 0 0.76 -0.117 0.65
L. gracile 0.968 -0.942 -0.671 -0.162 0.308 0.134 -0.013 0.123 0 0 -0.953 0.532
P. repens -0.057 -0.279 0.515 -0.99 0.727 0.993 0.877 0.994 0 0.895 0.005 0.817
P. scrobiculatum -0.231 0.161 0.036 -0.136 0.349 0.106 -0.058 0.107 0 -0.093 0.174 0.004
R. repens 0.436 -0.316 -0.734 0.431 -0.269 -0.488 -0.524 -0.487 0 0.035 -0.489 -0.171
C. roseus 0.959 -0.883 -0.798 -0.06 0.419 0.026 -0.193 0.01 0 0.914 -0.931 0.428
S. rhombifolia 0.994 -0.894 -0.858 0.05 0.215 -0.087 -0.244 -0.1 0 0.854 -0.972 0.355
U. sp. -0.994 0.892 0.848 -0.055 -0.194 0.09 0.238 0.103 0 -0.851 0.971 -0.352
U. lobata -0.972 0.975 0.746 0.147 -0.3 -0.107 0.052 -0.095 0 0 0.991 -0.547
A. conyzoides 1 0.915 0.818 0.01 -0.255 0.022 0.177 0.036 0 -0.914 0.974 -0.408
A. subulatus 0.915 1 -0.604 -0.343 0.435 0.307 0.135 0.296 0 0 -0.949 0.719
B. bipinnata 0.818 -0.604 1 0.45 -0.063 -0.494 -0.62 -0.506 0 0.309 -0.794 -0.06
B. pilosa 0.01 -0.343 0.45 1 0.741 0.995 0.867 0.988 0 0.955 -0.062 0.841
E. sonchifolia -0.255 0.435 -0.063 0.741 1 -0.738 -0.48 -0.721 0 -0.888 0.239 -0.706
E. acer 0.022 0.307 -0.494 0.995 -0.738 1 -0.884 -0.993 0 -0.92 0.019 -0.827
E. chinense 0.177 0.135 -0.62 0.867 -0.48 -0.884 1 -0.884 0 -0.685 -0.143 -0.678
M. micrantha 0.036 0.296 -0.506 0.988 -0.721 -0.993 -0.884 1 0 -0.91 0.01 -0.823
P. indica 0 0 0 0 0 0 0 0 1 0 0 0
V. cinerea -0.914 0 0.309 0.955 -0.888 -0.92 -0.685 -0.91 0 1 0.995 0
W. triloba 0.974 -0.949 -0.794 -0.062 0.239 0.019 -0.143 0.01 0 0.995 1 0.475
J. linifolia -0.408 0.719 -0.06 0.841 -0.706 -0.827 -0.678 -0.823 0 0 0.475 1
P. triquetrum 0.063 -0.251 0.238 -0.62 0.587 0.619 0.444 0.619 0 0.633 -0.092 0.543
C. asiatica 0.043 -0.349 0.407 -0.942 0.705 0.943 0.855 0.921 0 0.919 -0.077 0.797
C. exaltatus 0.92 -0.753 -0.933 0.335 -0.07 -0.374 -0.471 -0.387 0 0.534 -0.911 0.09
C. compressus -0.853 0.974 0.482 0.487 -0.54 -0.454 -0.269 -0.446 0 0 0.889 -0.786
K. brevifolia 0.029 0.304 -0.497 0.988 -0.737 -0.993 -0.88 -0.993 0 -0.924 0.019 -0.833
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L. cernuum 0 0 0 0 0 0 0 0 0 0 0 0
A. philoxeroides -0.002 0.318 -0.482 0.974 -0.74 -0.983 -0.876 -0.977 0 -0.921 0.034 -0.821
A. sessilis 0 0 0 0 0 0 0 0 0 0 0 0
I. triloba 0 0 0 0 0 0 0 0 -0.999 0 0 0
P. purpurea -0.979 0.89 0.866 -0.09 -0.129 0.13 0.268 0.141 0 -0.824 0.982 -0.324
Z. pendula -0.979 0.895 0.859 -0.076 -0.134 0.117 0.254 0.128 0 -0.836 0.983 -0.334
O. corniculata 0.98 -0.895 -0.859 0.075 0.135 -0.116 -0.253 -0.128 0 0.837 -0.983 0.334
n=50.

P. C. C. K. A. P. Z. O.
C. exaltatus L. cernuum A. sessilis I. triloba
triquetrum asiatica compressus brevifolia philoxeroides purpurea pendula corniculata
E. hirta -0.438 -0.635 -0.43 0.942 0.632 0 0.633 0 0 0.634 0.642 -0.642
S. sebiferum 0 0 0 0 0 0.011 0 0.999 0 0 0 0
A. scarabaeoides -0.434 -0.501 0.824 -0.286 0.607 0 0.6 0 0 -0.716 -0.71 0.709
D. gangeticum 0 0 0 0 0 0.999 0 0.011 0 0 0 0
D. triflorum -0.081 -0.024 -0.004 0.021 0.023 0 0.131 0 0 0.008 0.011 -0.011
S. cannabina 0.17 0.225 0.833 -0.942 -0.172 0 -0.188 0 0 -0.939 -0.943 0.943
P. vittata 0.433 0.627 0.44 -0.944 -0.623 0 -0.624 0 0 -0.642 -0.65 0.65
L. japonicum 0.526 0.636 -0.072 -0.486 -0.677 0 -0.687 0 0 -0.109 -0.113 0.114
A. compressus -0.176 -0.2 0.234 -0.011 0.223 0 0.188 0 0 -0.188 -0.183 0.183
C. aciculatus 0.33 0.833 -0.178 -0.42 -0.741 0 -0.78 0 0 0.002 -0.02 0.02
C. citratus 0.535 0.837 0.057 -0.763 -0.86 0 -0.864 0 0 -0.297 -0.308 0.309
C. dactylon -0.036 0.006 -0.948 0.839 -0.078 0 -0.056 0 0 0.995 0.995 -0.995
E. crusgalli -0.624 -0.923 0.35 0.48 0.992 0 0.977 0 0 -0.102 -0.089 0.088
E. minor -0.323 -0.177 0.482 -0.226 0.284 0 0.208 0 0 -0.436 -0.435 0.434
E. pilosa 0 0.026 -0.96 0.824 -0.109 0 -0.082 0 0 0.999 0.999 -0.999
I. aristatum -0.625 -0.655 0.2 0.409 0.73 0 0.715 0 0 -0.012 -0.007 0.006
L. gracile -0.165 -0.183 -0.837 0.903 0.127 0 0.159 0 0 0.933 0.936 -0.936
P. repens -0.635 -0.917 0.403 0.429 0.994 0 0.974 0 0 -0.16 -0.147 0.146
P. scrobiculatum -0.092 -0.06 0.238 -0.13 0.117 0 0.035 0 0 -0.219 -0.222 0.222
R. repens 0.307 0.427 -0.633 0.206 -0.487 0 -0.518 0 0 0.545 0.541 -0.54
C. roseus -0.16 -0.08 -0.854 0.834 0.021 0 0.049 0 0 0.924 0.923 -0.923
S. rhombifolia -0.03 0.018 -0.942 0.822 -0.093 0 -0.064 0 0 0.985 0.984 -0.985
U. sp. 0.021 -0.021 0.943 -0.82 0.097 0 0.066 0 0 -0.985 -0.985 0.985
U. lobata 0.13 0.169 0.877 -0.923 -0.102 0 -0.123 0 0 -0.968 -0.971 0.971
A. conyzoides 0.063 0.043 0.92 -0.853 0.029 0 -0.002 0 0 -0.979 -0.979 0.98
A. subulatus -0.251 -0.349 -0.753 0.974 0.304 0 0.318 0 0 0.89 0.895 -0.895
B. bipinnata 0.238 0.407 -0.933 0.482 -0.497 0 -0.482 0 0 0.866 0.859 -0.859
B. pilosa -0.62 -0.942 0.335 0.487 0.988 0 0.974 0 0 -0.09 -0.076 0.075
E. sonchifolia 0.587 0.705 -0.07 -0.54 -0.737 0 -0.74 0 0 -0.129 -0.134 0.135
E. acer 0.619 0.943 -0.374 -0.454 -0.993 0 -0.983 0 0 0.13 0.117 -0.116
E. chinense 0.444 0.855 -0.471 -0.269 -0.88 0 -0.876 0 0 0.268 0.254 -0.253
M. micrantha 0.619 0.921 -0.387 -0.446 -0.993 0 -0.977 0 0 0.141 0.128 -0.128
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P. indica 0 0 0 0 0 0 0 0 -0.999 0 0 0
V. cinerea 0.633 0.919 0.534 0 -0.924 0 -0.921 0 0 -0.824 -0.836 0.837
W. triloba -0.092 -0.077 -0.911 0.889 0.019 0 0.034 0 0 0.982 0.983 -0.983
J. linifolia 0.543 0.797 0.09 -0.786 -0.833 0 -0.821 0 0 -0.324 -0.334 0.334
P. triquetrum 1 -0.559 0.175 0.343 0.623 0 0.627 0 0 -0.015 -0.009 0.008
C. asiatica -0.559 1 0.284 0.475 0.922 0 0.944 0 0 -0.058 -0.041 0.04
C. exaltatus 0.175 0.284 1 0.639 -0.38 0 -0.352 0 0 0.966 0.964 -0.964
C. compressus 0.343 0.475 0.639 1 -0.453 0 -0.463 0 0 -0.809 -0.816 0.816
K. brevifolia 0.623 0.922 -0.38 -0.453 1 0 -0.978 0 0 0.136 0.124 -0.123
L. cernuum 0 0 0 0 0 1 0 -0.011 0 0 0 0
A. philoxeroides 0.627 0.944 -0.352 -0.463 -0.978 0 1 0 0 0.111 0.098 -0.097
A. sessilis 0 0 0 0 0 -0.011 0 1 0 0 0 0
I. triloba 0 0 0 0 0 0 0 0 1 0 0 0
P. purpurea -0.015 -0.058 0.966 -0.809 0.136 0 0.111 0 0 1 -0.999 0.999
Z. pendula -0.009 -0.041 0.964 -0.816 0.124 0 0.098 0 0 -0.999 1 0.999
O. corniculata 0.008 0.04 -0.964 0.816 -0.123 0 -0.097 0 0 0.999 0.999 1
n=50.
The significant species pairs, with 99% significant level, are listed as bellow
E. hirta-P. vittata, E. hirta-J. linifolia, E. hirta-C. compressus;

S. sebiferum-A. sessilis;
D. gangeticum-L. cernuum;
S. cannabina-C. dactylon, S. cannabina-E. pilosa, S. cannabina-L. gracile, S. cannabina-S. rhombifolia, S.
cannabina-U. sp., S. cannabina-U. lobata, S. cannabina-A. conyzoides, S. cannabina-A. subulatus, S.
cannabina-W. triloba, S. cannabina-C. compressus, S. cannabina-P. purpurea, S. cannabina-Z. pendula, S.
cannabina-O. corniculata;
P. vittata-J. linifolia, P. vittata-C. compressus;
C. dactylon-E. pilosa, C. dactylon-L. gracile, C. dactylon-C. roseus, C. dactylon-S. rhombifolia, C.
dactylon-U. sp., C. dactylon-U. lobata, C. dactylon-A. conyzoides, C. dactylon-W. triloba, C. dactylon-C.
exaltatus, C. dactylon-P. purpurea, C. dactylon-Z. pendula, C. dactylon-O. corniculata;
E. crusgalli-P. repens, E. crusgalli-B. pilosa, E. crusgalli-E. acer, E. crusgalli-M. micrantha, E.
crusgalli-V. cinerea, E. crusgalli-K. brevifolia, E. crusgalli-A. philoxeroides;
E. pilosa-L. gracile, E. pilosa-S. rhombifolia, E. pilosa-U. sp., E. pilosa-U. lobata, E. pilosa-A.
conyzoides, E. pilosa-W. triloba, E. pilosa-C. exaltatus, E. pilosa-P. purpurea, E. pilosa-Z. pendula, E.
pilosa-O. corniculata;
L. gracile-S. rhombifolia, L. gracile-U. sp., L. gracile-U. lobata, L. gracile-A. conyzoides, L. gracile-A.
subulatus, L. gracile-W. triloba, L. gracile-P. purpurea, L. gracile-Z. pendula, L. gracile-O. corniculata;
P. repens-B. pilosa, P. repens-E. acer, P. repens-M. micrantha, P. repens-K. brevifolia, P. repens-A.
philoxeroides;
C. roseus-S. rhombifolia, C. roseus-U. sp., C. roseus-A. conyzoides;
S. rhombifolia-U. sp., S. rhombifolia-U. lobata, S. rhombifolia-A. conyzoides, S. rhombifolia-W. triloba, S.
rhombifolia-C. exaltatus, S. rhombifolia-P. purpurea, S. rhombifolia-Z. pendula, S. rhombifolia-O.
corniculata;
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U. sp.-U. lobata, U. sp.-A. conyzoides, U. sp.-W. triloba, U. sp.-C. exaltatus, U. sp.-P. purpurea, U. sp.-Z.
pendula, U. sp.-O. corniculata;
U. lobata-A. conyzoides, U. lobata-A. subulatus, U. lobata-W. triloba, U. lobata-P. purpurea, U. lobata-Z.
pendula, U. lobata-O. corniculata;
A. conyzoides-W. triloba, A. conyzoides-P. purpurea, A. conyzoides-Z. pendula, A. conyzoides-O.
corniculata;
A. subulatus-W. triloba, A. subulatus-C. compressus;
B. bipinnata-C. exaltatus;
B. pilosa-E. acer, B. pilosa-M. micrantha, B. pilosa-V. cinerea, B. pilosa-C. asiatica, B. pilosa-K.
brevifolia, B. pilosa-A. philoxeroides;
E. acer-M. micrantha, E. acer-C. asiatica, E. acer-K. brevifolia, E. acer-A. philoxeroides;
M. micrantha-K. brevifolia, M. micrantha-A. philoxeroides;
P. indica-I. triloba;
V. cinerea-W. triloba;
W. triloba-P. purpurea, W. triloba-Z. pendula, W. triloba-O. corniculata;
C. asiatica-A. philoxeroides;
C. exaltatus-P. purpurea, C. exaltatus-Z. pendula, C. exaltatus-O. corniculata;
K. brevifolia-A. philoxeroides;
P. purpurea-Z. pendula, P. purpurea-O. corniculata;
Z. pendula-O. corniculata.
3.3 Associations measured by Spearman rank correlation coefficient

Inter-family Spearman rank correlation coefficients are listed in Table 9.
Table 9 Inter-family Spearman rank correlation coefficients.

Euphorbiaceae 1 0.56 0.939 0.85 0.451 0.882 0.827 0.467 0.911
Leguminosae 0.56 1 0.54 0.466 -0.329 0.472 0.475 0.343 0.505
Pteridaceae 0.939 0.54 1 0.91 0.491 0.939 0.78 0.46 0.91
Lygodiaceae 0.85 0.466 0.91 1 0.503 0.91 0.743 0.391 0.879
Gramineae 0.451 -0.329 0.491 0.503 1 0.445 0.26 -0.159 0.487
Apocynaceae 0.882 0.472 0.939 0.91 0.445 1 0.78 0.501 0.91
Malvaceae 0.827 0.475 0.78 0.743 0.26 0.78 1 0.567 0.87
Asteraceae 0.467 0.343 0.46 0.391 -0.159 0.501 0.567 1 0.495
Onagraceae 0.911 0.505 0.91 0.879 0.487 0.91 0.87 0.495 1
Connaraceae 0.882 0.472 0.939 0.91 0.45 0.939 0.78 0.453 0.91
Umbelliferae 0.708 0.317 0.78 0.743 0.468 0.78 0.766 0.378 0.809
Cyperaceae 0.79 0.287 0.855 0.822 0.517 0.855 0.673 0.401 0.822
Lycopodiaceae 0.882 0.53 0.939 0.969 0.486 0.939 0.78 0.439 0.91
Amaranthaceae 0.914 0.524 0.91 0.879 0.484 0.91 0.797 0.451 0.879
Convolvulaceae 0.775 0.424 0.78 0.743 0.376 0.78 0.76 0.317 0.812
Commelinaceae 0.819 0.347 0.882 0.85 0.519 0.882 0.708 0.465 0.85
Oxalidaceae 0.79 0.357 0.855 0.822 0.469 0.855 0.731 0.395 0.822
n=50.
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Table 9 Inter-family Spearman rank correlation coefficients (Continue).

Euphorbiaceae 0.882 0.708 0.79 0.882 0.914 0.775 0.819 0.79
Leguminosae 0.472 0.317 0.287 0.53 0.524 0.424 0.347 0.357
Pteridaceae 0.939 0.78 0.855 0.939 0.91 0.78 0.882 0.855
Lygodiaceae 0.91 0.743 0.822 0.969 0.879 0.743 0.85 0.822
Gramineae 0.45 0.468 0.517 0.486 0.484 0.376 0.519 0.469
Apocynaceae 0.939 0.78 0.855 0.939 0.91 0.78 0.882 0.855
Malvaceae 0.78 0.766 0.673 0.78 0.797 0.76 0.708 0.731
Asteraceae 0.453 0.378 0.401 0.439 0.451 0.317 0.465 0.395
Onagraceae 0.91 0.809 0.822 0.91 0.879 0.812 0.85 0.822
Connaraceae 1 0.78 0.855 0.939 0.91 0.78 0.882 0.855
Umbelliferae 0.78 1 0.673 0.78 0.797 0.701 0.762 0.797
Cyperaceae 0.855 0.673 1 0.855 0.822 0.73 0.851 0.759
Lycopodiaceae 0.939 0.78 0.855 1 0.91 0.78 0.882 0.855
Amaranthaceae 0.91 0.797 0.822 0.91 1 0.743 0.85 0.822
Convolvulaceae 0.78 0.701 0.73 0.78 0.743 1 0.708 0.79
Commelinaceae 0.882 0.762 0.851 0.882 0.85 0.708 1 0.913
Oxalidaceae 0.855 0.797 0.759 0.855 0.822 0.79 0.913 1
n=50.
Table 9 indicates that at the 99 % confidence level, Most of family pairs show positive associations. All
family pairs are associated positively and significantly, other than the following family pairs
Leguminosae-: Gramineae, Asteraceae, Umbelliferae, Cyperaceae, Commelinaceae, Oxalidaceae;

Gramineae-: Malvaceae, Asteraceae;
Asteraceae-Convolvulaceae.
Interspecific Spearman rank correlation coefficients are shown in Table 10.
Table 10 Interspecfic Spearman rank correlation coefficients.

S. A. D. D. S. A. C. C.
E. hirta P. vittata L. japonicum C. dactylon
sebiferum scarabaeoides gangeticum triflorum cannabina compressus aciculatus citratus
E. hirta 1 0.91 0.612 0.91 0.768 0.879 0.969 0.879 0.559 0.822 0.91 0.768
S. sebiferum 0.91 1 0.521 0.939 0.87 0.91 0.939 0.91 0.563 0.855 0.939 0.804
A. scarabaeoides 0.612 0.521 1 0.521 0.351 0.501 0.581 0.465 -0.123 0.435 0.556 0.698
D. gangeticum 0.91 0.939 0.521 1 0.804 0.91 0.939 0.969 0.549 0.855 0.939 0.804
D. triflorum 0.768 0.87 0.351 0.804 1 0.768 0.804 0.768 0.457 0.701 0.804 0.638
S. cannabina 0.879 0.91 0.501 0.91 0.768 1 0.91 0.879 0.513 0.822 0.91 0.768
P. vittata 0.969 0.939 0.581 0.939 0.804 0.91 1 0.91 0.573 0.855 0.939 0.804
L. japonicum 0.879 0.91 0.465 0.969 0.768 0.879 0.91 1 0.487 0.822 0.91 0.768
A. compressus 0.559 0.563 -0.123 0.549 0.457 0.513 0.573 0.487 1 0.485 0.556 0.268
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C. aciculatus 0.822 0.855 0.435 0.855 0.701 0.822 0.855 0.822 0.485 1 0.913 0.701
C. citratus 0.91 0.939 0.556 0.939 0.804 0.91 0.939 0.91 0.556 0.913 1 0.804
C. dactylon 0.768 0.804 0.698 0.804 0.638 0.768 0.804 0.768 0.268 0.701 0.804 1
E. crusgalli 0.879 0.91 0.465 0.91 0.768 0.879 0.91 0.879 0.492 0.822 0.91 0.768
E. minor 0.595 0.64 -0.001 0.716 0.667 0.595 0.64 0.669 0.208 0.508 0.64 0.426
E. pilosa 0.822 0.855 0.452 0.855 0.701 0.822 0.855 0.822 0.345 0.759 0.855 0.701
I. aristatum 0.653 0.694 0.153 0.694 0.566 0.653 0.694 0.725 0.239 0.573 0.694 0.498
L. gracile 0.879 0.91 0.585 0.91 0.768 0.879 0.91 0.879 0.426 0.822 0.91 0.768
P. repens 0.456 0.509 0.011 0.446 0.293 0.505 0.478 0.402 0.141 0.473 0.515 0.223
P. scrobiculatum 0.672 0.657 0.275 0.704 0.583 0.613 0.717 0.659 0.46 0.529 0.657 0.52
R. repens 0.638 0.608 0.618 0.608 0.496 0.561 0.608 0.561 -0.184 0.469 0.608 0.659
C. roseus 0.91 0.939 0.521 0.939 0.804 0.91 0.939 0.91 0.542 0.855 0.939 0.804
S. rhombifolia 0.879 0.91 0.581 0.91 0.768 0.879 0.91 0.879 0.426 0.822 0.91 0.824
U. sp. 0.91 0.939 0.576 0.939 0.804 0.91 0.939 0.91 0.486 0.855 0.939 0.804
U. lobata 0.886 0.911 0.493 0.855 0.763 0.822 0.855 0.822 0.431 0.759 0.855 0.701
A. conyzoides 0.613 0.657 0.413 0.657 0.449 0.613 0.657 0.613 0.125 0.529 0.657 0.706
A. subulatus 0.822 0.855 0.483 0.855 0.701 0.879 0.855 0.822 0.308 0.759 0.855 0.701
B. bipinnata 0.768 0.804 0.677 0.804 0.638 0.768 0.804 0.768 0.237 0.701 0.804 0.864
B. pilosa 0.856 0.891 0.572 0.829 0.737 0.794 0.829 0.794 0.492 0.729 0.829 0.669
E. sonchifolia 0.418 0.509 -0.075 0.516 0.354 0.418 0.48 0.501 0.21 0.411 0.48 0.465
E. acer 0.809 0.842 0.468 0.78 0.676 0.809 0.847 0.743 0.495 0.723 0.78 0.608
E. chinense 0.532 0.532 0.227 0.575 0.311 0.544 0.588 0.568 0.201 0.422 0.532 0.378
M. micrantha 0.768 0.804 0.298 0.804 0.638 0.768 0.804 0.768 0.406 0.701 0.804 0.638
P. indica 0.91 0.939 0.569 0.939 0.804 0.91 0.939 0.91 0.486 0.855 0.939 0.804
V. cinerea 0.768 0.869 0.419 0.804 0.709 0.768 0.804 0.768 0.407 0.701 0.804 0.694
W. triloba 0.794 0.891 0.337 0.829 0.737 0.853 0.829 0.794 0.562 0.838 0.884 0.669
J. linifolia 0.942 0.91 0.553 0.91 0.768 0.879 0.91 0.879 0.475 0.822 0.91 0.768
P. triquetrum 0.91 0.939 0.521 0.939 0.804 0.91 0.939 0.91 0.552 0.855 0.939 0.804
C. asiatica 0.743 0.78 0.409 0.78 0.665 0.743 0.78 0.743 0.211 0.732 0.78 0.608
C. exaltatus 0.91 0.939 0.521 0.939 0.804 0.91 0.939 0.91 0.486 0.855 0.939 0.804
C. compressus 0.91 0.939 0.521 0.939 0.804 0.91 0.939 0.91 0.568 0.911 0.939 0.804
K. brevifolia 0.879 0.91 0.465 0.91 0.768 0.879 0.91 0.879 0.455 0.822 0.91 0.768
L. cernuum 0.91 0.939 0.521 1 0.804 0.91 0.939 0.969 0.549 0.855 0.939 0.804
A. philoxeroides 0.91 0.939 0.574 0.939 0.866 0.91 0.939 0.91 0.522 0.855 0.939 0.804
A. sessilis 0.91 1 0.521 0.939 0.87 0.91 0.939 0.91 0.563 0.855 0.939 0.804
I. triloba 0.91 0.939 0.569 0.939 0.804 0.91 0.939 0.91 0.486 0.855 0.939 0.804
P. purpurea 0.812 0.78 0.501 0.78 0.608 0.795 0.78 0.743 0.305 0.739 0.78 0.674
Z. pendula 0.85 0.882 0.411 0.882 0.734 0.85 0.882 0.85 0.445 0.79 0.882 0.734
O. corniculata 0.822 0.855 0.444 0.855 0.701 0.822 0.855 0.822 0.417 0.816 0.855 0.701
n=50.
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Table 10 Interspecfic Spearman rank correlation coefficients (Continue).

I. L. P.
E. crusgalli E. minor E. pilosa P. repens R. repens C. roseus S. rhombifolia U. sp. U. lobata
aristatum gracile scrobiculatum
E. hirta 0.879 0.595 0.822 0.653 0.879 0.456 0.672 0.638 0.91 0.879 0.91 0.886
S. sebiferum 0.91 0.64 0.855 0.694 0.91 0.509 0.657 0.608 0.939 0.91 0.939 0.911
A. scarabaeoides 0.465 -0.001 0.452 0.153 0.585 0.011 0.275 0.618 0.521 0.581 0.576 0.493
D. gangeticum 0.91 0.716 0.855 0.694 0.91 0.446 0.704 0.608 0.939 0.91 0.939 0.855
D. triflorum 0.768 0.667 0.701 0.566 0.768 0.293 0.583 0.496 0.804 0.768 0.804 0.763
S. cannabina 0.879 0.595 0.822 0.653 0.879 0.505 0.613 0.561 0.91 0.879 0.91 0.822
P. vittata 0.91 0.64 0.855 0.694 0.91 0.478 0.717 0.608 0.939 0.91 0.939 0.855
L. japonicum 0.879 0.669 0.822 0.725 0.879 0.402 0.659 0.561 0.91 0.879 0.91 0.822
A. compressus 0.492 0.208 0.345 0.239 0.426 0.141 0.46 -0.184 0.542 0.426 0.486 0.431
C. aciculatus 0.822 0.508 0.759 0.573 0.822 0.473 0.529 0.469 0.855 0.822 0.855 0.759
C. citratus 0.91 0.64 0.855 0.694 0.91 0.515 0.657 0.608 0.939 0.91 0.939 0.855
C. dactylon 0.768 0.426 0.701 0.498 0.768 0.223 0.52 0.659 0.804 0.824 0.804 0.701
E. crusgalli 1 0.595 0.879 0.768 0.879 0.534 0.613 0.561 0.91 0.879 0.91 0.822
E. minor 0.595 1 0.508 0.239 0.595 -0.237 0.408 0.466 0.64 0.595 0.64 0.508
E. pilosa 0.879 0.508 1 0.8 0.822 0.534 0.647 0.469 0.855 0.884 0.855 0.815
I. aristatum 0.768 0.239 0.8 1 0.653 0.519 0.43 0.183 0.694 0.702 0.694 0.63
L. gracile 0.879 0.595 0.822 0.653 1 0.413 0.613 0.623 0.91 0.879 0.971 0.822
P. repens 0.534 -0.237 0.534 0.519 0.413 1 0.173 -0.166 0.5 0.472 0.446 0.506
P. scrobiculatum 0.613 0.408 0.647 0.43 0.613 0.173 1 0.277 0.711 0.674 0.657 0.582
R. repens 0.561 0.466 0.469 0.183 0.623 -0.166 0.277 1 0.608 0.603 0.669 0.549
C. roseus 0.91 0.64 0.855 0.694 0.91 0.5 0.711 0.608 1 0.91 0.939 0.855
S. rhombifolia 0.879 0.595 0.884 0.702 0.879 0.472 0.674 0.603 0.91 1 0.91 0.822
U. sp. 0.91 0.64 0.855 0.694 0.971 0.446 0.657 0.669 0.939 0.91 1 0.855
U. lobata 0.822 0.508 0.815 0.63 0.822 0.506 0.582 0.549 0.855 0.822 0.855 1
A. conyzoides 0.736 0.17 0.755 0.579 0.684 0.345 0.449 0.283 0.724 0.746 0.657 0.578
A. subulatus 0.822 0.508 0.88 0.681 0.822 0.538 0.645 0.469 0.855 0.88 0.855 0.883
B. bipinnata 0.768 0.426 0.701 0.498 0.898 0.242 0.449 0.591 0.804 0.823 0.87 0.701
B. pilosa 0.794 0.519 0.729 0.535 0.794 0.454 0.489 0.573 0.829 0.794 0.829 0.852
E. sonchifolia 0.555 0.004 0.493 0.57 0.418 0.297 0.467 0 0.57 0.466 0.48 0.441
E. acer 0.743 0.387 0.673 0.462 0.743 0.529 0.525 0.394 0.78 0.743 0.78 0.732
E. chinense 0.477 0.266 0.429 0.258 0.477 0.152 0.545 0.319 0.532 0.477 0.532 0.512
M. micrantha 0.891 0.426 0.88 0.83 0.768 0.585 0.562 0.382 0.804 0.824 0.804 0.701
P. indica 0.91 0.64 0.918 0.745 0.91 0.505 0.72 0.608 0.939 0.969 0.939 0.855
V. cinerea 0.768 0.426 0.876 0.653 0.768 0.46 0.689 0.434 0.869 0.825 0.804 0.818
W. triloba 0.794 0.515 0.729 0.535 0.794 0.508 0.489 0.425 0.829 0.794 0.829 0.788
J. linifolia 0.879 0.595 0.822 0.653 0.879 0.515 0.613 0.638 0.91 0.879 0.91 0.946
P. triquetrum 0.91 0.687 0.855 0.694 0.91 0.446 0.724 0.608 0.939 0.91 0.939 0.855
C. asiatica 0.743 0.387 0.802 0.692 0.743 0.545 0.638 0.398 0.78 0.806 0.78 0.804
C. exaltatus 0.97 0.64 0.912 0.761 0.91 0.533 0.657 0.608 0.939 0.91 0.939 0.855
C. compressus 0.91 0.64 0.855 0.694 0.91 0.524 0.657 0.608 0.939 0.91 0.939 0.855
K. brevifolia 0.879 0.638 0.822 0.716 0.879 0.548 0.613 0.561 0.91 0.879 0.91 0.822
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L. cernuum 0.91 0.716 0.855 0.694 0.91 0.446 0.704 0.608 0.939 0.91 0.939 0.855
A. philoxeroides 0.91 0.64 0.855 0.759 0.91 0.526 0.657 0.608 0.939 0.91 0.939 0.855
A. sessilis 0.91 0.64 0.855 0.694 0.91 0.509 0.657 0.608 0.939 0.91 0.939 0.911
I. triloba 0.91 0.64 0.918 0.745 0.91 0.505 0.72 0.608 0.939 0.969 0.939 0.855
P. purpurea 0.743 0.429 0.792 0.568 0.743 0.383 0.525 0.491 0.78 0.804 0.78 0.799
Z. pendula 0.909 0.551 0.91 0.792 0.85 0.555 0.627 0.514 0.882 0.85 0.882 0.79
O. corniculata 0.822 0.508 0.878 0.747 0.822 0.512 0.645 0.469 0.855 0.822 0.855 0.819
n=50.

E. E. M.
A. conyzoides A. subulatus B. bipinnata B. pilosa E. acer P. indica V. cinerea W. triloba J. linifolia
sonchifolia chinense micrantha
E. hirta 0.613 0.822 0.768 0.856 0.418 0.809 0.532 0.768 0.91 0.768 0.794 0.942
S. sebiferum 0.657 0.855 0.804 0.891 0.509 0.842 0.532 0.804 0.939 0.869 0.891 0.91
A. scarabaeoides 0.413 0.483 0.677 0.572 -0.075 0.468 0.227 0.298 0.569 0.419 0.337 0.553
D. gangeticum 0.657 0.855 0.804 0.829 0.516 0.78 0.575 0.804 0.939 0.804 0.829 0.91
D. triflorum 0.449 0.701 0.638 0.737 0.354 0.676 0.311 0.638 0.804 0.709 0.737 0.768
S. cannabina 0.613 0.879 0.768 0.794 0.418 0.809 0.544 0.768 0.91 0.768 0.853 0.879
P. vittata 0.657 0.855 0.804 0.829 0.48 0.847 0.588 0.804 0.939 0.804 0.829 0.91
L. japonicum 0.613 0.822 0.768 0.794 0.501 0.743 0.568 0.768 0.91 0.768 0.794 0.879
A. compressus 0.125 0.308 0.237 0.492 0.21 0.495 0.201 0.406 0.486 0.407 0.562 0.475
C. aciculatus 0.529 0.759 0.701 0.729 0.411 0.723 0.422 0.701 0.855 0.701 0.838 0.822
C. citratus 0.657 0.855 0.804 0.829 0.48 0.78 0.532 0.804 0.939 0.804 0.884 0.91
C. dactylon 0.706 0.701 0.864 0.669 0.465 0.608 0.378 0.638 0.804 0.694 0.669 0.768
E. crusgalli 0.736 0.822 0.768 0.794 0.555 0.743 0.477 0.891 0.91 0.768 0.794 0.879
E. minor 0.17 0.508 0.426 0.519 0.004 0.387 0.266 0.426 0.64 0.426 0.515 0.595
E. pilosa 0.755 0.88 0.701 0.729 0.493 0.673 0.429 0.88 0.918 0.876 0.729 0.822
I. aristatum 0.579 0.681 0.498 0.535 0.57 0.462 0.258 0.83 0.745 0.653 0.535 0.653
L. gracile 0.684 0.822 0.898 0.794 0.418 0.743 0.477 0.768 0.91 0.768 0.794 0.879
P. repens 0.345 0.538 0.242 0.454 0.297 0.529 0.152 0.585 0.505 0.46 0.508 0.515
P. scrobiculatum 0.449 0.645 0.449 0.489 0.467 0.525 0.545 0.562 0.72 0.689 0.489 0.613
R. repens 0.283 0.469 0.591 0.573 0 0.394 0.319 0.382 0.608 0.434 0.425 0.638
C. roseus 0.724 0.855 0.804 0.829 0.57 0.78 0.532 0.804 0.939 0.869 0.829 0.91
S. rhombifolia 0.746 0.88 0.823 0.794 0.466 0.743 0.477 0.824 0.969 0.825 0.794 0.879
U. sp. 0.657 0.855 0.87 0.829 0.48 0.78 0.532 0.804 0.939 0.804 0.829 0.91
U. lobata 0.578 0.883 0.701 0.852 0.441 0.732 0.512 0.701 0.855 0.818 0.788 0.946
A. conyzoides 1 0.637 0.775 0.488 0.404 0.411 0.103 0.732 0.715 0.679 0.488 0.613
A. subulatus 0.637 1 0.701 0.729 0.417 0.734 0.578 0.757 0.915 0.816 0.729 0.884
B. bipinnata 0.775 0.701 1 0.669 0.303 0.608 0.27 0.638 0.804 0.638 0.669 0.768
B. pilosa 0.488 0.729 0.669 1 0.27 0.824 0.45 0.669 0.829 0.736 0.763 0.856
E. sonchifolia 0.404 0.417 0.303 0.27 1 0.207 0.26 0.552 0.48 0.516 0.27 0.485
E. acer 0.411 0.734 0.608 0.824 0.207 1 0.54 0.608 0.78 0.675 0.756 0.743
E. chinense 0.103 0.578 0.27 0.45 0.26 0.54 1 0.383 0.532 0.393 0.372 0.556
M. micrantha 0.732 0.757 0.638 0.669 0.552 0.608 0.383 1 0.86 0.753 0.669 0.768
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P. indica 0.715 0.915 0.804 0.829 0.48 0.78 0.532 0.86 1 0.861 0.829 0.91
V. cinerea 0.679 0.816 0.638 0.736 0.516 0.675 0.393 0.753 0.861 1 0.736 0.768
W. triloba 0.488 0.729 0.669 0.763 0.27 0.756 0.372 0.669 0.829 0.736 1 0.794
J. linifolia 0.613 0.884 0.768 0.856 0.485 0.743 0.556 0.768 0.91 0.768 0.794 1
P. triquetrum 0.657 0.855 0.804 0.829 0.549 0.78 0.58 0.804 0.939 0.804 0.829 0.91
C. asiatica 0.57 0.872 0.608 0.64 0.483 0.633 0.505 0.72 0.843 0.73 0.64 0.809
C. exaltatus 0.719 0.855 0.804 0.829 0.54 0.78 0.532 0.86 0.939 0.804 0.829 0.91
C. compressus 0.657 0.855 0.804 0.829 0.48 0.833 0.586 0.804 0.939 0.804 0.886 0.91
K. brevifolia 0.613 0.822 0.768 0.794 0.48 0.743 0.477 0.768 0.91 0.768 0.858 0.879
L. cernuum 0.657 0.855 0.804 0.829 0.516 0.78 0.575 0.804 0.939 0.804 0.829 0.91
A. philoxeroides 0.657 0.855 0.804 0.829 0.527 0.78 0.532 0.804 0.939 0.804 0.829 0.91
A. sessilis 0.657 0.855 0.804 0.891 0.509 0.842 0.532 0.804 0.939 0.869 0.891 0.91
I. triloba 0.715 0.915 0.804 0.829 0.48 0.78 0.532 0.86 1 0.861 0.829 0.91
P. purpurea 0.517 0.791 0.608 0.707 0.297 0.577 0.313 0.667 0.842 0.721 0.755 0.812
Z. pendula 0.736 0.79 0.734 0.761 0.585 0.708 0.474 0.911 0.882 0.794 0.761 0.85
O. corniculata 0.685 0.82 0.701 0.729 0.553 0.673 0.487 0.823 0.855 0.816 0.729 0.822
n=50.

P. C. C. A. Z. O.
C. exaltatus K. brevifolia L. cernuum A. sessilis I. triloba P. purpurea
triquetrum asiatica compressus philoxeroides pendula corniculata
E. hirta 0.91 0.743 0.91 0.91 0.879 0.91 0.91 0.91 0.91 0.812 0.85 0.822
S. sebiferum 0.939 0.78 0.939 0.939 0.91 0.939 0.939 1 0.939 0.78 0.882 0.855
A. scarabaeoides 0.521 0.409 0.521 0.521 0.465 0.521 0.574 0.521 0.569 0.501 0.411 0.444
D. gangeticum 0.939 0.78 0.939 0.939 0.91 1 0.939 0.939 0.939 0.78 0.882 0.855
D. triflorum 0.804 0.665 0.804 0.804 0.768 0.804 0.866 0.87 0.804 0.608 0.734 0.701
S. cannabina 0.91 0.743 0.91 0.91 0.879 0.91 0.91 0.91 0.91 0.795 0.85 0.822
P. vittata 0.939 0.78 0.939 0.939 0.91 0.939 0.939 0.939 0.939 0.78 0.882 0.855
L. japonicum 0.91 0.743 0.91 0.91 0.879 0.969 0.91 0.91 0.91 0.743 0.85 0.822
A. compressus 0.552 0.211 0.486 0.568 0.455 0.549 0.522 0.563 0.486 0.305 0.445 0.417
C. aciculatus 0.855 0.732 0.855 0.911 0.822 0.855 0.855 0.855 0.855 0.739 0.79 0.816
C. citratus 0.939 0.78 0.939 0.939 0.91 0.939 0.939 0.939 0.939 0.78 0.882 0.855
C. dactylon 0.804 0.608 0.804 0.804 0.768 0.804 0.804 0.804 0.804 0.674 0.734 0.701
E. crusgalli 0.91 0.743 0.97 0.91 0.879 0.91 0.91 0.91 0.91 0.743 0.909 0.822
E. minor 0.687 0.387 0.64 0.64 0.638 0.716 0.64 0.64 0.64 0.429 0.551 0.508
E. pilosa 0.855 0.802 0.912 0.855 0.822 0.855 0.855 0.855 0.918 0.792 0.91 0.878
I. aristatum 0.694 0.692 0.761 0.694 0.716 0.694 0.759 0.694 0.745 0.568 0.792 0.747
L. gracile 0.91 0.743 0.91 0.91 0.879 0.91 0.91 0.91 0.91 0.743 0.85 0.822
P. repens 0.446 0.545 0.533 0.524 0.548 0.446 0.526 0.509 0.505 0.383 0.555 0.512
P. scrobiculatum 0.724 0.638 0.657 0.657 0.613 0.704 0.657 0.657 0.72 0.525 0.627 0.645
R. repens 0.608 0.398 0.608 0.608 0.561 0.608 0.608 0.608 0.608 0.491 0.514 0.469
C. roseus 0.939 0.78 0.939 0.939 0.91 0.939 0.939 0.939 0.939 0.78 0.882 0.855
S. rhombifolia 0.91 0.806 0.91 0.91 0.879 0.91 0.91 0.91 0.969 0.804 0.85 0.822
U. sp. 0.939 0.78 0.939 0.939 0.91 0.939 0.939 0.939 0.939 0.78 0.882 0.855
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U. lobata 0.855 0.804 0.855 0.855 0.822 0.855 0.855 0.911 0.855 0.799 0.79 0.819
A. conyzoides 0.657 0.57 0.719 0.657 0.613 0.657 0.657 0.657 0.715 0.517 0.736 0.685
A. subulatus 0.855 0.872 0.855 0.855 0.822 0.855 0.855 0.855 0.915 0.791 0.79 0.82
B. bipinnata 0.804 0.608 0.804 0.804 0.768 0.804 0.804 0.804 0.804 0.608 0.734 0.701
B. pilosa 0.829 0.64 0.829 0.829 0.794 0.829 0.829 0.891 0.829 0.707 0.761 0.729
E. sonchifolia 0.549 0.483 0.54 0.48 0.48 0.516 0.527 0.509 0.48 0.297 0.585 0.553
E. acer 0.78 0.633 0.78 0.833 0.743 0.78 0.78 0.842 0.78 0.577 0.708 0.673
E. chinense 0.58 0.505 0.532 0.586 0.477 0.575 0.532 0.532 0.532 0.313 0.474 0.487
M. micrantha 0.804 0.72 0.86 0.804 0.768 0.804 0.804 0.804 0.86 0.667 0.911 0.823
P. indica 0.939 0.843 0.939 0.939 0.91 0.939 0.939 0.939 1 0.842 0.882 0.855
V. cinerea 0.804 0.73 0.804 0.804 0.768 0.804 0.804 0.869 0.861 0.721 0.794 0.816
W. triloba 0.829 0.64 0.829 0.886 0.858 0.829 0.829 0.891 0.829 0.755 0.761 0.729
J. linifolia 0.91 0.809 0.91 0.91 0.879 0.91 0.91 0.91 0.91 0.812 0.85 0.822
P. triquetrum 1 0.78 0.939 0.939 0.91 0.939 0.939 0.939 0.939 0.78 0.882 0.855
C. asiatica 0.78 1 0.78 0.78 0.743 0.78 0.835 0.78 0.843 0.701 0.762 0.797
C. exaltatus 0.939 0.78 1 0.939 0.91 0.939 0.939 0.939 0.939 0.78 0.941 0.855
C. compressus 0.939 0.78 0.939 1 0.91 0.939 0.939 0.939 0.939 0.78 0.882 0.855
K. brevifolia 0.91 0.743 0.91 0.91 1 0.91 0.91 0.91 0.91 0.8 0.85 0.822
L. cernuum 0.939 0.78 0.939 0.939 0.91 1 0.939 0.939 0.939 0.78 0.882 0.855
A. philoxeroides 0.939 0.835 0.939 0.939 0.91 0.939 1 0.939 0.939 0.78 0.882 0.855
A. sessilis 0.939 0.78 0.939 0.939 0.91 0.939 0.939 1 0.939 0.78 0.882 0.855
I. triloba 0.939 0.843 0.939 0.939 0.91 0.939 0.939 0.939 1 0.842 0.882 0.855
P. purpurea 0.78 0.701 0.78 0.78 0.8 0.78 0.78 0.78 0.842 1 0.708 0.79
Z. pendula 0.882 0.762 0.941 0.882 0.85 0.882 0.882 0.882 0.882 0.708 1 0.913
O. corniculata 0.855 0.797 0.855 0.855 0.822 0.855 0.855 0.855 0.855 0.79 0.913 1
n=50.
Using the ID numbers of species in Table 2, the significant species pairs are listed as bellow
(1,2) (1,3) (1,4) (1,5) (1,6) (1,7) (1,8) (1,9) (1,10) (1,11) (1,12) (1,13) (1,14) (1,15) (1,16) (1,17) (1,18) (1,19)
(1,20) (1,21) (1,22) (1,23) (1,24) (1,25) (1,26) (1,27) (1,28) (1,29) (1,30) (1,31) (1,32) (1,33) (1,34) (1,35)
(1,36) (1,37) (1,38) (1,39) (1,40) (1,41) (1,42) (1,43) (1,44) (1,45) (1,46) (1,47) (1,48)
(2,3) (2,4) (2,5) (2,6) (2,7) (2,8) (2,9) (2,10) (2,11) (2,12) (2,13) (2,14) (2,15) (2,16) (2,17) (2,18) (2,19) (2,20)
(2,21) (2,22) (2,23) (2,24) (2,25) (2,26) (2,27) (2,28) (2,29) (2,30) (2,31) (2,32) (2,33) (2,34) (2,35) (2,36)
(2,37) (2,38) (2,39) (2,40) (2,41) (2,42) (2,43) (2,45) (2,46) (2,47) (2,48)
(3,4) (3,6) (3,7) (3,8) (3,10) (3,11) (3,12) (3,13) (3,15) (3,17) (3,20) (3,21) (3,22) (3,23) (3,24) (3,25) (3,26)
(3,27) (3,28) (3,30) (3,33) (3,34) (3,36) (3,37) (3,38) (3,39) (3,40) (3,41) (3,42) (3,43) (3,44) (3,45) (3,46)
(3,47) (3,48)
(4,5) (4,6) (4,7) (4,8) (4,9) (4,10) (4,11) (4,12) (4,13) (4,14) (4,15) (4,16) (4,17) (4,18) (4,19) (4,20) (4,21)
(4,22) (4,23) (4,24) (4,25) (4,26) (4,27) (4,28) (4,29) (4,30) (4,31) (4,32) (4,33) (4,34) (4,35) (4,36) (4,37)
(4,38) (4,39) (4,40) (4,41) (4,43) (4,44) (4,45) (4,46) (4,47) (4,48)
(5,6) (5,7) (5,8) (5,9) (5,10) (5,11) (5,12) (5,13) (5,14) (5,15) (5,16) (5,17) (5,19) (5,20) (5,21) (5,22) (5,23)
(5,24) (5,25) (5,26) (5,27) (5,28) (5,30) (5,32) (5,33) (5,34) (5,35) (5,36) (5,37) (5,38) (5,39) (5,40) (5,41)
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(5,42) (5,43) (5,44) (5,45) (5,46) (5,47) (5,48)

(6,7) (6,8) (6,9) (6,10) (6,11) (6,12) (6,13) (6,14) (6,15) (6,16) (6,17) (6,18) (6,19) (6,20) (6,21) (6,22) (6,23)
(6,24) (6,25) (6,26) (6,27) (6,28) (6,29) (6,30) (6,31) (6,32) (6,33) (6,34) (6,35) (6,36) (6,37) (6,38) (6,39)
(6,40) (6,41) (6,42) (6,43) (6,44) (6,45) (6,46) (6,47) (6,48)
(7,8) (7,9) (7,10) (7,11) (7,12) (7,13) (7,14) (7,15) (7,16) (7,17) (7,18) (7,19) (7,20) (7,21) (7,22) (7,23) (7,24)
(7,25) (7,26) (7,27) (7,28) (7,29) (7,30) (7,31) (7,32) (7,33) (7,34) (7,35) (7,36) (7,37) (7,38) (7,39) (7,40)
(7,41) (7,42) (7,43) (7,44) (7,45) (7,46) (7,47) (7,48)
(8,9) (8,10) (8,11) (8,12) (8,13) (8,14) (8,15) (8,16) (8,17) (8,18) (8,19) (8,20) (8,21) (8,22) (8,23) (8,24) (8,25)
(8,26) (8,27) (8,28) (8,29) (8,30) (8,31) (8,32) (8,33) (8,34) (8,35) (8,36) (8,37) (8,38) (8,39) (8,40) (8,41)
(8,42) (8,43) (8,44) (8,45) (8,46) (8,47) (8,48)
(9,10) (9,11) (9,13) (9,17) (9,19) (9,21) (9,22) (9,23) (9,24) (9,28) (9,30) (9,32) (9,33) (9,34) (9,35) (9,36)
(9,37) (9,39) (9,40) (9,41) (9,42) (9,43) (9,44) (9,45) (9,47) (9,48)
(10,11) (10,12) (10,13) (10,14) (10,15) (10,16) (10,17) (10,18) (10,19) (10,20) (10,21) (10,22) (10,23) (10,24)
(10,25) (10,26) (10,27) (10,28) (10,29) (10,30) (10,31) (10,32) (10,33) (10,34) (10,35) (10,36) (10,37) (10,38)
(10,39) (10,40) (10,41) (10,42) (10,43) (10,44) (10,45) (10,46) (10,47) (10,48)
(11,12) (11,13) (11,14) (11,15) (11,16) (11,17) (11,18) (11,19) (11,20) (11,21) (11,22) (11,23) (11,24) (11,25)
(11,26) (11,27) (11,28) (11,29) (11,30) (11,31) (11,32) (11,33) (11,34) (11,35) (11,36) (11,37) (11,38) (11,39)
(11,40) (11,41) (11,42) (11,43) (11,44) (11,45) (11,46) (11,47) (11,48)
(12,13) (12,14) (12,15) (12,16) (12,17) (12,19) (12,20) (12,21) (12,22) (12,23) (12,24) (12,25) (12,26) (12,27)
(12,28) (12,29) (12,30) (12,31) (12,32) (12,33) (12,34) (12,35) (12,36) (12,37) (12,38) (12,39) (12,40) (12,41)
(12,42) (12,43) (12,44) (12,45) (12,46) (12,47) (12,48)
(13,14) (13,15) (13,16) (13,17) (13,18) (13,19) (13,20) (13,21) (13,22) (13,23) (13,24) (13,25) (13,26) (13,27)
(13,28) (13,29) (13,30) (13,31) (13,32) (13,33) (13,34) (13,35) (13,36) (13,37) (13,38) (13,39) (13,40) (13,41)
(13,42) (13,43) (13,44) (13,45) (13,46) (13,47) (13,48)
(14,15) (14,17) (14,19) (14,20) (14,21) (14,22) (14,23) (14,24) (14,26) (14,27) (14,28) (14,30) (14,32) (14,33)
(14,34) (14,35) (14,36) (14,37) (14,38) (14,39) (14,40) (14,41) (14,42) (14,43) (14,44) (14,45) (14,46) (14,47)
(14,48)
(15,16) (15,17) (15,18) (15,19) (15,20) (15,21) (15,22) (15,23) (15,24) (15,25) (15,26) (15,27) (15,28) (15,29)
(15,30) (15,31) (15,32) (15,33) (15,34) (15,35) (15,36) (15,37) (15,38) (15,39) (15,40) (15,41) (15,42) (15,43)
(15,44) (15,45) (15,46) (15,47) (15,48)
(16,17) (16,18) (16,19) (16,21) (16,22) (16,23) (16,24) (16,25) (16,26) (16,27) (16,28) (16,29) (16,30) (16,32)
(16,33) (16,34) (16,35) (16,36) (16,37) (16,38) (16,39) (16,40) (16,41) (16,42) (16,43) (16,44) (16,45) (16,46)
(16,47) (16,48)
(17,18) (17,19) (17,20) (17,21) (17,22) (17,23) (17,24) (17,25) (17,26) (17,27) (17,28) (17,29) (17,30) (17,31)
(17,32) (17,33) (17,34) (17,35) (17,36) (17,37) (17,38) (17,39) (17,40) (17,41) (17,42) (17,43) (17,44) (17,45)
(17,46) (17,47) (17,48)
(18,21) (18,22) (18,23) (18,24) (18,26) (18,28) (18,30) (18,32) (18,33) (18,34) (18,35) (18,36) (18,37) (18,38)
(18,39) (18,40) (18,41) (18,42) (18,43) (18,44) (18,45) (18,46) (18,47) (18,48)
(19,21) (19,22) (19,23) (19,24) (19,25) (19,26) (19,27) (19,28) (19,29) (19,30) (19,31) (19,32) (19,33) (19,34)
(19,35) (19,36) (19,37) (19,38) (19,39) (19,40) (19,41) (19,42) (19,43) (19,44) (19,45) (19,46) (19,47) (19,48)
(20,21) (20,22) (20,23) (20,24) (20,26) (20,27) (20,28) (20,30) (20,32) (20,33) (20,34) (20,35) (20,36) (20,37)
(20,38) (20,39) (20,40) (20,41) (20,42) (20,43) (20,44) (20,45) (20,46) (20,47) (20,48)
(21,22) (21,23) (21,24) (21,25) (21,26) (21,27) (21,28) (21,29) (21,30) (21,31) (21,32) (21,33) (21,34) (21,35)
(21,36) (21,37) (21,38) (21,39) (21,40) (21,41) (21,42) (21,43) (21,44) (21,45) (21,46) (21,47) (21,48)
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(22,23) (22,24) (22,25) (22,26) (22,27) (22,28) (22,29) (22,30) (22,31) (22,32) (22,33) (22,34) (22,35) (22,36)
(22,37) (22,38) (22,39) (22,40) (22,41) (22,42) (22,43) (22,44) (22,45) (22,46) (22,47) (22,48)
(23,24) (23,25) (23,26) (23,27) (23,28) (23,29) (23,30) (23,31) (23,32) (23,33) (23,34) (23,35) (23,36) (23,37)
(23,38) (23,39) (23,40) (23,41) (23,42) (23,43) (23,44) (23,45) (23,46) (23,47) (23,48)
(24,25) (24,26) (24,27) (24,28) (24,29) (24,30) (24,31) (24,32) (24,33) (24,34) (24,35) (24,36) (24,37) (24,38)
(24,39) (24,40) (24,41) (24,42) (24,43) (24,44) (24,45) (24,46) (24,47) (24,48)
(25,26) (25,27) (25,28) (25,29) (25,30) (25,32) (25,33) (25,34) (25,35) (25,36) (25,37) (25,38) (25,39) (25,40)
(25,41) (25,42) (25,43) (25,44) (25,45) (25,46) (25,47) (25,48)
(26,27) (26,28) (26,29) (26,30) (26,31) (26,32) (26,33) (26,34) (26,35) (26,36) (26,37) (26,38) (26,39) (26,40)
(26,41) (26,42) (26,43) (26,44) (26,45) (26,46) (26,47) (26,48)
(27,28) (27,30) (27,32) (27,33) (27,34) (27,35) (27,36) (27,37) (27,38) (27,39) (27,40) (27,41) (27,42) (27,43)
(27,44) (27,45) (27,46) (27,47) (27,48)
(28,30) (28,31) (28,32) (28,33) (28,34) (28,35) (28,36) (28,37) (28,38) (28,39) (28,40) (28,41) (28,42) (28,43)
(28,44) (28,45) (28,46) (28,47) (28,48)
(29,32) (29,33) (29,34) (29,36) (29,37) (29,38) (29,39) (29,40) (29,41) (29,42) (29,43) (29,44) (29,45) (29,47)
(29,48)
(30,31) (30,32) (30,33) (30,34) (30,35) (30,36) (30,37) (30,38) (30,39) (30,40) (30,41) (30,42) (30,43) (30,44)
(30,45) (30,46) (30,47) (30,48)
(31,32) (31,33) (31,34) (31,35) (31,36) (31,37) (31,38) (31,39) (31,40) (31,41) (31,42) (31,43) (31,44) (31,45)
(31,47) (31,48)
(32,33) (32,34) (32,35) (32,36) (32,37) (32,38) (32,39) (32,40) (32,41) (32,42) (32,43) (32,44) (32,45) (32,46)
(32,47) (32,48)
(33,34) (33,35) (33,36) (33,37) (33,38) (33,39) (33,40) (33,41) (33,42) (33,43) (33,44) (33,46) (33,47) (33,48)
(34,35) (34,36) (34,37) (34,38) (34,39) (34,40) (34,41) (34,42) (34,43) (34,44) (34,45) (34,46) (34,47) (34,48)
(35,36) (35,37) (35,38) (35,39) (35,40) (35,41) (35,42) (35,43) (35,44) (35,45) (35,46) (35,47) (35,48)
(36,37) (36,38) (36,39) (36,40) (36,41) (36,42) (36,43) (36,44) (36,45) (36,46) (36,47) (36,48)
(37,38) (37,39) (37,40) (37,41) (37,42) (37,43) (37,44) (37,45) (37,46) (37,47) (37,48)
(38,39) (38,40) (38,41) (38,42) (38,43) (38,44) (38,45) (38,46) (38,47) (38,48)
(39,40) (39,41) (39,42) (39,43) (39,44) (39,45) (39,46) (39,47) (39,48)
(40,41) (40,42) (40,43) (40,44) (40,45) (40,46) (40,47) (40,48)
(41,42) (41,43) (41,44) (41,45) (41,46) (41,47) (41,48)
(42,43) (42,44) (42,45) (42,46) (42,47) (42,48)
(43,44) (43,45) (43,46) (43,47) (43,48)
(44,45) (44,46) (44,47) (44,48)
(45,46) (45,47) (45,48)
(46,47) (46,48)
(47,48)
3.4 Associations measured by point correlation coefficient

Inter-family point correlation coefficients are shown in Table 11.
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Table 11 Inter-family point correlation coefficients.

Euphorbiaceae 1 0.214 0.565 -0.051 0 -0.036 0.383 0.074 0.378
Leguminosae 0.214 1 0.121 -0.033 0 -0.167 0.226 0.197 -0.033
Pteridaceae 0.565 0.121 1 -0.029 0 -0.02 -0.057 0.042 -0.029
Lygodiaceae -0.051 -0.033 -0.029 1 0 -0.029 -0.082 0.06 -0.041
Gramineae 0 0 0 0 1 0 0 0 0
Apocynaceae -0.036 -0.167 -0.02 -0.029 0 1 -0.057 0.042 -0.029
Malvaceae 0.383 0.226 -0.057 -0.082 0 -0.057 1 0.118 0.505
Asteraceae 0.074 0.197 0.042 0.06 0 0.042 0.118 1 0.06
Onagraceae 0.378 -0.033 -0.029 -0.041 0 -0.029 0.505 0.06 1
Connaraceae -0.036 -0.167 -0.02 -0.029 0 -0.02 -0.057 0.042 -0.029
Umbelliferae -0.101 -0.007 -0.057 -0.082 0 -0.057 0.335 0.118 0.211
Cyperaceae -0.074 -0.346 -0.042 -0.06 0 -0.042 -0.118 0.086 -0.06
Lycopodiaceae -0.036 0.121 -0.02 0.699 0 -0.02 -0.057 0.042 -0.029
Amaranthaceae 0.378 0.173 -0.029 -0.041 0 -0.029 0.211 0.06 -0.041
Convolvulaceae 0.14 0.226 -0.057 -0.082 0 -0.057 0.335 0.118 0.211
Commelinaceae -0.063 -0.296 -0.036 -0.051 0 -0.036 -0.101 0.074 -0.051
Oxalidaceae -0.074 -0.047 -0.042 -0.06 0 -0.042 0.093 0.086 -0.06
n=50.
Table 11 Inter-family point correlation coefficients (Continue).

Euphorbiaceae -0.036 -0.101 -0.074 -0.036 0.378 0.14 -0.063 -0.074
Leguminosae -0.167 -0.007 -0.346 0.121 0.173 0.226 -0.296 -0.047
Pteridaceae -0.02 -0.057 -0.042 -0.02 -0.029 -0.057 -0.036 -0.042
Lygodiaceae -0.029 -0.082 -0.06 0.699 -0.041 -0.082 -0.051 -0.06
Gramineae 0 0 0 0 0 0 0 0
Apocynaceae -0.02 -0.057 -0.042 -0.02 -0.029 -0.057 -0.036 -0.042
Malvaceae -0.057 0.335 -0.118 -0.057 0.211 0.335 -0.101 0.093
Asteraceae 0.042 0.118 0.086 0.042 0.06 0.118 0.074 0.086
Onagraceae -0.029 0.211 -0.06 -0.029 -0.041 0.211 -0.051 -0.06
Connaraceae 1 -0.057 -0.042 -0.02 -0.029 -0.057 -0.036 -0.042
Umbelliferae -0.057 1 -0.118 -0.057 0.211 0.169 0.14 0.305
Cyperaceae -0.042 -0.118 1 -0.042 -0.06 0.093 0.235 -0.086
Lycopodiaceae -0.02 -0.057 -0.042 1 -0.029 -0.057 -0.036 -0.042
Amaranthaceae -0.029 0.211 -0.06 -0.029 1 -0.082 -0.051 -0.06
Convolvulaceae -0.057 0.169 0.093 -0.057 -0.082 1 -0.101 0.305
Commelinaceae -0.036 0.14 0.235 -0.036 -0.051 -0.101 1 0.546
Oxalidaceae -0.042 0.305 -0.086 -0.042 -0.06 0.305 0.546 1
n=50.
Table 11 indicates that at the 99 % confidence level, the associations of Gramineae to other families is not
significant and all remaining family pairs are significantly associated and about half of them are positive
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associations.
Interspecific point correlation coefficients are shown in Table 12.
Table 12 Interspecfic point correlation coefficients.

S. A. D. S. L. C. C.
E. hirta D. triflorum P. vittata A. compressus C. dactylon
sebiferum scarabaeoides gangeticum cannabina japonicum aciculatus citratus
E. hirta 1 -0.029 0.221 -0.029 -0.075 -0.041 0.699 -0.041 0.188 -0.06 -0.029 -0.075
S. sebiferum -0.029 1 -0.131 -0.02 0.386 -0.029 -0.02 -0.029 0.131 -0.042 -0.02 -0.052
A. scarabaeoides 0.221 -0.131 1 -0.131 -0.093 0.016 0.154 -0.188 -0.194 0.023 0.154 0.4
D. gangeticum -0.029 -0.02 -0.131 1 -0.052 -0.029 -0.02 0.699 0.131 -0.042 -0.02 -0.052
D. triflorum -0.075 0.386 -0.093 -0.052 1 -0.075 -0.052 -0.075 0.093 -0.108 -0.052 -0.136
S. cannabina -0.041 -0.029 0.016 -0.029 -0.075 1 -0.029 -0.041 -0.016 -0.06 -0.029 -0.075
P. vittata 0.699 -0.02 0.154 -0.02 -0.052 -0.029 1 -0.029 0.131 -0.042 -0.02 -0.052
L. japonicum -0.041 -0.029 -0.188 0.699 -0.075 -0.041 -0.029 1 -0.016 -0.06 -0.029 -0.075
A. compressus 0.188 0.131 -0.194 0.131 0.093 -0.016 0.131 -0.016 1 0.124 0.131 -0.153
C. aciculatus -0.06 -0.042 0.023 -0.042 -0.108 -0.06 -0.042 -0.06 0.124 1 0.484 -0.108
C. citratus -0.029 -0.02 0.154 -0.02 -0.052 -0.029 -0.02 -0.029 0.131 0.484 1 -0.052
C. dactylon -0.075 -0.052 0.4 -0.052 -0.136 -0.075 -0.052 -0.075 -0.153 -0.108 -0.052 1
E. crusgalli -0.041 -0.029 -0.188 -0.029 -0.075 -0.041 -0.029 -0.041 -0.016 -0.06 -0.029 -0.075
E. minor -0.127 -0.089 -0.396 0.229 0.318 -0.127 -0.089 0.1 -0.139 -0.183 -0.089 -0.23
E. pilosa -0.06 -0.042 0.023 -0.042 -0.108 -0.06 -0.042 -0.06 -0.171 -0.086 -0.042 -0.108
I. aristatum -0.108 -0.075 -0.199 -0.075 -0.047 -0.108 -0.075 0.137 0.102 -0.156 -0.075 -0.196
L. gracile -0.041 -0.029 0.221 -0.029 -0.075 -0.041 -0.029 -0.041 -0.221 -0.06 -0.029 -0.075
P. repens 0.14 0.097 0.116 -0.208 -0.142 0.14 0.097 -0.078 0.141 0.202 0.097 -0.142
P. scrobiculatum 0.111 -0.084 0.001 0.241 0.061 -0.12 0.241 0.111 0.272 -0.174 -0.084 -0.078
R. repens 0.078 -0.097 0.399 -0.097 0.01 -0.14 -0.097 -0.14 -0.485 -0.202 -0.097 0.406
C. roseus -0.029 -0.02 -0.131 -0.02 -0.052 -0.029 -0.02 -0.029 0.131 -0.042 -0.02 -0.052
S. rhombifolia -0.041 -0.029 0.221 -0.029 -0.075 -0.041 -0.029 -0.041 -0.221 -0.06 -0.029 0.238
U. sp. -0.029 -0.02 0.154 -0.02 -0.052 -0.029 -0.02 -0.029 -0.154 -0.042 -0.02 -0.052
U. lobata 0.316 0.484 0.023 -0.042 0.117 -0.06 -0.042 -0.06 -0.023 -0.086 -0.042 -0.108
A. conyzoides -0.12 -0.084 0.093 -0.084 -0.218 -0.12 -0.084 -0.12 -0.093 -0.174 -0.084 0.342
A. subulatus -0.06 -0.042 0.171 -0.042 -0.108 0.316 -0.042 -0.06 -0.319 -0.086 -0.042 -0.108
B. bipinnata -0.075 -0.052 0.4 -0.052 -0.136 -0.075 -0.052 -0.075 -0.276 -0.108 -0.052 0.621
B. pilosa 0.272 0.428 0.227 -0.047 0.082 -0.068 -0.047 -0.068 0.173 -0.098 -0.047 -0.123
E. sonchifolia -0.23 0.126 -0.313 0.126 0.203 -0.23 -0.161 0.18 0.313 -0.035 -0.161 0.079
E. acer 0.211 0.354 0.205 -0.057 0.028 0.211 0.354 -0.082 0.141 0.093 -0.057 -0.148
E. chinense 0.024 -0.126 -0.009 0.161 -0.203 0.024 0.161 0.23 0.009 -0.112 -0.126 -0.079
M. micrantha -0.075 -0.052 -0.217 -0.052 -0.136 -0.075 -0.052 -0.075 0.093 -0.108 -0.052 -0.136
P. indica -0.029 -0.02 0.154 -0.02 -0.052 -0.029 -0.02 -0.029 -0.154 -0.042 -0.02 -0.052
V. cinerea -0.075 0.386 0.029 -0.052 0.053 -0.075 -0.052 -0.075 0.093 -0.108 -0.052 0.053
W. triloba -0.068 0.428 -0.173 -0.047 0.082 0.272 -0.047 -0.068 0.173 0.393 0.428 -0.123
J. linifolia 0.479 -0.029 0.016 -0.029 -0.075 -0.041 -0.029 -0.041 -0.016 -0.06 -0.029 -0.075
P. triquetrum -0.029 -0.02 -0.131 -0.02 -0.052 -0.029 -0.02 -0.029 0.131 -0.042 -0.02 -0.052
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C. asiatica -0.082 -0.057 0.09 -0.057 0.028 -0.082 -0.057 -0.082 -0.205 0.093 -0.057 -0.148
C. exaltatus -0.029 -0.02 -0.131 -0.02 -0.052 -0.029 -0.02 -0.029 -0.154 -0.042 -0.02 -0.052
C. compressus -0.029 -0.02 -0.131 -0.02 -0.052 -0.029 -0.02 -0.029 0.131 0.484 -0.02 -0.052
K. brevifolia -0.041 -0.029 -0.188 -0.029 -0.075 -0.041 -0.029 -0.041 -0.016 -0.06 -0.029 -0.075
L. cernuum -0.029 -0.02 -0.131 1 -0.052 -0.029 -0.02 0.699 0.131 -0.042 -0.02 -0.052
A. philoxeroides -0.029 -0.02 0.154 -0.02 0.386 -0.029 -0.02 -0.029 0.131 -0.042 -0.02 -0.052
A. sessilis -0.029 1 -0.131 -0.02 0.386 -0.029 -0.02 -0.029 0.131 -0.042 -0.02 -0.052
I. triloba -0.029 -0.02 0.154 -0.02 -0.052 -0.029 -0.02 -0.029 -0.154 -0.042 -0.02 -0.052
P. purpurea 0.211 -0.057 0.205 -0.057 -0.148 0.211 -0.057 -0.082 -0.09 0.093 -0.057 0.028
Z. pendula -0.051 -0.036 -0.233 -0.036 -0.093 -0.051 -0.036 -0.051 0.064 -0.074 -0.036 -0.093
O. corniculata -0.06 -0.042 0.023 -0.042 -0.108 -0.06 -0.042 -0.06 0.124 0.184 -0.042 -0.108
n=50.
Table 12 Interspecfic point correlation coefficients (Continue).

I. P.
E. crusgalli E. minor E. pilosa L. gracile P. repens R. repens C. roseus S. rhombifolia U. sp. U. lobata
aristatum scrobiculatum
E. hirta -0.041 -0.127 -0.06 -0.108 -0.041 0.14 0.111 0.078 -0.029 -0.041 -0.029 0.316
S. sebiferum -0.029 -0.089 -0.042 -0.075 -0.029 0.097 -0.084 -0.097 -0.02 -0.029 -0.02 0.484
A. scarabaeoides -0.188 -0.396 0.023 -0.199 0.221 0.116 0.001 0.399 -0.131 0.221 0.154 0.023
D. gangeticum -0.029 0.229 -0.042 -0.075 -0.029 -0.208 0.241 -0.097 -0.02 -0.029 -0.02 -0.042
D. triflorum -0.075 0.318 -0.108 -0.047 -0.075 -0.142 0.061 0.01 -0.052 -0.075 -0.052 0.117
S. cannabina -0.041 -0.127 -0.06 -0.108 -0.041 0.14 -0.12 -0.14 -0.029 -0.041 -0.029 -0.06
P. vittata -0.029 -0.089 -0.042 -0.075 -0.029 0.097 0.241 -0.097 -0.02 -0.029 -0.02 -0.042
L. japonicum -0.041 0.1 -0.06 0.137 -0.041 -0.078 0.111 -0.14 -0.029 -0.041 -0.029 -0.06
A. compressus -0.016 -0.139 -0.171 0.102 -0.221 0.141 0.272 -0.485 0.131 -0.221 -0.154 -0.023
C. aciculatus -0.06 -0.183 -0.086 -0.156 -0.06 0.202 -0.174 -0.202 -0.042 -0.06 -0.042 -0.086
C. citratus -0.029 -0.089 -0.042 -0.075 -0.029 0.097 -0.084 -0.097 -0.02 -0.029 -0.02 -0.042
C. dactylon -0.075 -0.23 -0.108 -0.196 -0.075 -0.142 -0.078 0.406 -0.052 0.238 -0.052 -0.108
E. crusgalli 1 -0.127 0.316 0.384 -0.041 0.14 -0.12 -0.14 -0.029 -0.041 -0.029 -0.06
E. minor -0.127 1 -0.183 -0.331 -0.127 -0.527 0.036 0.145 -0.089 -0.127 -0.089 -0.183
E. pilosa 0.316 -0.183 1 0.555 -0.06 0.202 0.161 -0.202 -0.042 0.316 -0.042 0.184
I. aristatum 0.384 -0.331 0.555 1 -0.108 0.26 0.015 -0.364 -0.075 0.137 -0.075 0.021
L. gracile -0.041 -0.127 -0.06 -0.108 1 -0.078 -0.12 0.078 -0.029 -0.041 0.699 -0.06
P. repens 0.14 -0.527 0.202 0.26 -0.078 1 0.015 -0.264 0.097 0.14 -0.208 0.202
P. scrobiculatum -0.12 0.036 0.161 0.015 -0.12 0.015 1 -0.113 0.241 0.111 -0.084 -0.006
R. repens -0.14 0.145 -0.202 -0.364 0.078 -0.264 -0.113 1 -0.097 0.078 0.208 -0.044
C. roseus -0.029 -0.089 -0.042 -0.075 -0.029 0.097 0.241 -0.097 1 -0.029 -0.02 -0.042
S. rhombifolia -0.041 -0.127 0.316 0.137 -0.041 0.14 0.111 0.078 -0.029 1 -0.029 -0.06
U. sp. -0.029 -0.089 -0.042 -0.075 0.699 -0.208 -0.084 0.208 -0.02 -0.029 1 -0.042
U. lobata -0.06 -0.183 0.184 0.021 -0.06 0.202 -0.006 -0.044 -0.042 -0.06 -0.042 1
A. conyzoides 0.344 -0.369 0.497 0.345 0.111 0.211 0.168 -0.113 0.241 0.344 -0.084 -0.006
A. subulatus -0.06 -0.183 0.456 0.199 -0.06 0.202 0.161 -0.202 -0.042 0.316 -0.042 0.456
B. bipinnata -0.075 -0.23 -0.108 -0.196 0.552 -0.142 -0.218 0.274 -0.052 0.238 0.386 -0.108
B. pilosa -0.068 -0.059 -0.098 -0.177 -0.068 0.228 -0.197 0.057 -0.047 -0.068 -0.047 0.393
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E. sonchifolia 0.18 -0.165 0.112 0.373 -0.23 0.169 0.341 -0.255 0.126 -0.024 -0.161 0.112
E. acer -0.082 -0.251 -0.118 -0.214 -0.082 0.276 0.023 -0.153 -0.057 -0.082 -0.057 0.093
E. chinense -0.18 0.075 -0.112 -0.081 -0.18 -0.082 0.393 0.082 -0.126 -0.18 -0.126 0.035
M. micrantha 0.552 -0.23 0.571 0.695 -0.075 0.253 0.061 -0.253 -0.052 0.238 -0.052 -0.108
P. indica -0.029 -0.089 0.484 0.268 -0.029 0.097 0.241 -0.097 -0.02 0.699 -0.02 -0.042
V. cinerea -0.075 -0.23 0.571 0.249 -0.075 0.253 0.342 -0.121 0.386 0.238 -0.052 0.344
W. triloba -0.068 -0.059 -0.098 -0.177 -0.068 0.228 -0.197 -0.228 -0.047 -0.068 -0.047 0.147
J. linifolia -0.041 -0.127 -0.06 -0.108 -0.041 0.14 -0.12 0.078 -0.029 -0.041 -0.029 0.692
P. triquetrum -0.029 0.229 -0.042 -0.075 -0.029 -0.208 0.241 -0.097 -0.02 -0.029 -0.02 -0.042
C. asiatica -0.082 -0.251 0.305 0.342 -0.082 0.276 0.286 -0.153 -0.057 0.211 -0.057 0.305
C. exaltatus 0.699 -0.089 0.484 0.268 -0.029 0.097 -0.084 -0.097 -0.02 -0.029 -0.02 -0.042
C. compressus -0.029 -0.089 -0.042 -0.075 -0.029 0.097 -0.084 -0.097 -0.02 -0.029 -0.02 -0.042
K. brevifolia -0.041 0.1 -0.06 0.137 -0.041 0.14 -0.12 -0.14 -0.029 -0.041 -0.029 -0.06
L. cernuum -0.029 0.229 -0.042 -0.075 -0.029 -0.208 0.241 -0.097 -0.02 -0.029 -0.02 -0.042
A. philoxeroides -0.029 -0.089 -0.042 0.268 -0.029 0.097 -0.084 -0.097 -0.02 -0.029 -0.02 -0.042
A. sessilis -0.029 -0.089 -0.042 -0.075 -0.029 0.097 -0.084 -0.097 -0.02 -0.029 -0.02 0.484
I. triloba -0.029 -0.089 0.484 0.268 -0.029 0.097 0.241 -0.097 -0.02 0.699 -0.02 -0.042
P. purpurea -0.082 -0.123 0.305 0.064 -0.082 0.153 0.023 -0.029 -0.057 0.211 -0.057 0.305
Z. pendula 0.378 -0.157 0.546 0.475 -0.051 0.173 0.042 -0.173 -0.036 -0.051 -0.036 -0.074
O. corniculata -0.06 -0.183 0.456 0.377 -0.06 0.202 0.161 -0.202 -0.042 -0.06 -0.042 0.184
n=50.

E. E. M. V.
A. conyzoides A. subulatus B. bipinnata B. pilosa E. acer P. indica W. triloba J. linifolia
sonchifolia chinense micrantha cinerea
E. hirta -0.12 -0.06 -0.075 0.272 -0.23 0.211 0.024 -0.075 -0.029 -0.075 -0.068 0.479
S. sebiferum -0.084 -0.042 -0.052 0.428 0.126 0.354 -0.126 -0.052 -0.02 0.386 0.428 -0.029
A. scarabaeoides 0.093 0.171 0.4 0.227 -0.313 0.205 -0.009 -0.217 0.154 0.029 -0.173 0.016
D. gangeticum -0.084 -0.042 -0.052 -0.047 0.126 -0.057 0.161 -0.052 -0.02 -0.052 -0.047 -0.029
D. triflorum -0.218 -0.108 -0.136 0.082 0.203 0.028 -0.203 -0.136 -0.052 0.053 0.082 -0.075
S. cannabina -0.12 0.316 -0.075 -0.068 -0.23 0.211 0.024 -0.075 -0.029 -0.075 0.272 -0.041
P. vittata -0.084 -0.042 -0.052 -0.047 -0.161 0.354 0.161 -0.052 -0.02 -0.052 -0.047 -0.029
L. japonicum -0.12 -0.06 -0.075 -0.068 0.18 -0.082 0.23 -0.075 -0.029 -0.075 -0.068 -0.041
A. compressus -0.093 -0.319 -0.276 0.173 0.313 0.141 0.009 0.093 -0.154 0.093 0.173 -0.016
C. aciculatus -0.174 -0.086 -0.108 -0.098 -0.035 0.093 -0.112 -0.108 -0.042 -0.108 0.393 -0.06
C. citratus -0.084 -0.042 -0.052 -0.047 -0.161 -0.057 -0.126 -0.052 -0.02 -0.052 0.428 -0.029
C. dactylon 0.342 -0.108 0.621 -0.123 0.079 -0.148 -0.079 -0.136 -0.052 0.053 -0.123 -0.075
E. crusgalli 0.344 -0.06 -0.075 -0.068 0.18 -0.082 -0.18 0.552 -0.029 -0.075 -0.068 -0.041
E. minor -0.369 -0.183 -0.23 -0.059 -0.165 -0.251 0.075 -0.23 -0.089 -0.23 -0.059 -0.127
E. pilosa 0.497 0.456 -0.108 -0.098 0.112 -0.118 -0.112 0.571 0.484 0.571 -0.098 -0.06
I. aristatum 0.345 0.199 -0.196 -0.177 0.373 -0.214 -0.081 0.695 0.268 0.249 -0.177 -0.108
L. gracile 0.111 -0.06 0.552 -0.068 -0.23 -0.082 -0.18 -0.075 -0.029 -0.075 -0.068 -0.041
P. repens 0.211 0.202 -0.142 0.228 0.169 0.276 -0.082 0.253 0.097 0.253 0.228 0.14
P. scrobiculatum 0.168 0.161 -0.218 -0.197 0.341 0.023 0.393 0.061 0.241 0.342 -0.197 -0.12
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R. repens -0.113 -0.202 0.274 0.057 -0.255 -0.153 0.082 -0.253 -0.097 -0.121 -0.228 0.078
C. roseus 0.241 -0.042 -0.052 -0.047 0.126 -0.057 -0.126 -0.052 -0.02 0.386 -0.047 -0.029
S. rhombifolia 0.344 0.316 0.238 -0.068 -0.024 -0.082 -0.18 0.238 0.699 0.238 -0.068 -0.041
U. sp. -0.084 -0.042 0.386 -0.047 -0.161 -0.057 -0.126 -0.052 -0.02 -0.052 -0.047 -0.029
U. lobata -0.006 0.456 -0.108 0.393 0.112 0.093 0.035 -0.108 -0.042 0.344 0.147 0.692
A. conyzoides 1 0.161 0.482 -0.197 0.157 -0.239 -0.249 0.482 0.241 0.342 -0.197 -0.12
A. subulatus 0.161 1 -0.108 -0.098 -0.035 0.093 0.184 0.117 0.484 0.344 -0.098 0.316
B. bipinnata 0.482 -0.108 1 -0.123 -0.168 -0.148 -0.327 -0.136 -0.052 -0.136 -0.123 -0.075
B. pilosa -0.197 -0.098 -0.123 1 -0.241 0.441 -0.026 -0.123 -0.047 0.082 0.111 0.272
E. sonchifolia 0.157 -0.035 -0.168 -0.241 1 -0.222 0.136 0.203 -0.161 0.203 -0.241 -0.024
E. acer -0.239 0.093 -0.148 0.441 -0.222 1 0.222 -0.148 -0.057 0.028 0.249 -0.082
E. chinense -0.249 0.184 -0.327 -0.026 0.136 0.222 1 -0.079 -0.126 -0.079 -0.161 0.024
M. micrantha 0.482 0.117 -0.136 -0.123 0.203 -0.148 -0.079 1 0.386 0.242 -0.123 -0.075
P. indica 0.241 0.484 -0.052 -0.047 -0.161 -0.057 -0.126 0.386 1 0.386 -0.047 -0.029
V. cinerea 0.342 0.344 -0.136 0.082 0.203 0.028 -0.079 0.242 0.386 1 0.082 -0.075
W. triloba -0.197 -0.098 -0.123 0.111 -0.241 0.249 -0.161 -0.123 -0.047 0.082 1 -0.068
J. linifolia -0.12 0.316 -0.075 0.272 -0.024 -0.082 0.024 -0.075 -0.029 -0.075 -0.068 1
P. triquetrum -0.084 -0.042 -0.052 -0.047 0.126 -0.057 0.161 -0.052 -0.02 -0.052 -0.047 -0.029
C. asiatica 0.155 0.518 -0.148 -0.134 0.241 0.003 0.106 0.205 0.354 0.205 -0.134 0.211
C. exaltatus 0.241 -0.042 -0.052 -0.047 0.126 -0.057 -0.126 0.386 -0.02 -0.052 -0.047 -0.029
C. compressus -0.084 -0.042 -0.052 -0.047 -0.161 0.354 0.161 -0.052 -0.02 -0.052 0.428 -0.029
K. brevifolia -0.12 -0.06 -0.075 -0.068 -0.024 -0.082 -0.18 -0.075 -0.029 -0.075 0.272 -0.041
L. cernuum -0.084 -0.042 -0.052 -0.047 0.126 -0.057 0.161 -0.052 -0.02 -0.052 -0.047 -0.029
A. philoxeroides -0.084 -0.042 -0.052 -0.047 0.126 -0.057 -0.126 -0.052 -0.02 -0.052 -0.047 -0.029
A. sessilis -0.084 -0.042 -0.052 0.428 0.126 0.354 -0.126 -0.052 -0.02 0.386 0.428 -0.029
I. triloba 0.241 0.484 -0.052 -0.047 -0.161 -0.057 -0.126 0.386 1 0.386 -0.047 -0.029
P. purpurea 0.023 0.305 -0.148 0.057 -0.106 -0.162 -0.125 0.028 0.354 0.205 0.249 0.211
Z. pendula 0.426 -0.074 -0.093 -0.084 0.223 -0.101 -0.054 0.684 -0.036 0.165 -0.084 -0.051
O. corniculata 0.329 0.184 -0.108 -0.098 0.261 -0.118 0.035 0.344 -0.042 0.344 -0.098 -0.06
n=50.

P. C. C. K. A. P.
C. exaltatus L. cernuum A. sessilis I. triloba Z. pendula O. corniculata
triquetrum asiatica compressus brevifolia philoxeroides purpurea
E. hirta -0.029 -0.082 -0.029 -0.029 -0.041 -0.029 -0.029 -0.029 -0.029 0.211 -0.051 -0.06
S. sebiferum -0.02 -0.057 -0.02 -0.02 -0.029 -0.02 -0.02 1 -0.02 -0.057 -0.036 -0.042
A. scarabaeoides -0.131 0.09 -0.131 -0.131 -0.188 -0.131 0.154 -0.131 0.154 0.205 -0.233 0.023
D. gangeticum -0.02 -0.057 -0.02 -0.02 -0.029 1 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
D. triflorum -0.052 0.028 -0.052 -0.052 -0.075 -0.052 0.386 0.386 -0.052 -0.148 -0.093 -0.108
S. cannabina -0.029 -0.082 -0.029 -0.029 -0.041 -0.029 -0.029 -0.029 -0.029 0.211 -0.051 -0.06
P. vittata -0.02 -0.057 -0.02 -0.02 -0.029 -0.02 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
L. japonicum -0.029 -0.082 -0.029 -0.029 -0.041 0.699 -0.029 -0.029 -0.029 -0.082 -0.051 -0.06
A. compressus 0.131 -0.205 -0.154 0.131 -0.016 0.131 0.131 0.131 -0.154 -0.09 0.064 0.124
C. aciculatus -0.042 0.093 -0.042 0.484 -0.06 -0.042 -0.042 -0.042 -0.042 0.093 -0.074 0.184
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C. citratus -0.02 -0.057 -0.02 -0.02 -0.029 -0.02 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
C. dactylon -0.052 -0.148 -0.052 -0.052 -0.075 -0.052 -0.052 -0.052 -0.052 0.028 -0.093 -0.108
E. crusgalli -0.029 -0.082 0.699 -0.029 -0.041 -0.029 -0.029 -0.029 -0.029 -0.082 0.378 -0.06
E. minor 0.229 -0.251 -0.089 -0.089 0.1 0.229 -0.089 -0.089 -0.089 -0.123 -0.157 -0.183
E. pilosa -0.042 0.305 0.484 -0.042 -0.06 -0.042 -0.042 -0.042 0.484 0.305 0.546 0.456
I. aristatum -0.075 0.342 0.268 -0.075 0.137 -0.075 0.268 -0.075 0.268 0.064 0.475 0.377
L. gracile -0.029 -0.082 -0.029 -0.029 -0.041 -0.029 -0.029 -0.029 -0.029 -0.082 -0.051 -0.06
P. repens -0.208 0.276 0.097 0.097 0.14 -0.208 0.097 0.097 0.097 0.153 0.173 0.202
P. scrobiculatum 0.241 0.286 -0.084 -0.084 -0.12 0.241 -0.084 -0.084 0.241 0.023 0.042 0.161
R. repens -0.097 -0.153 -0.097 -0.097 -0.14 -0.097 -0.097 -0.097 -0.097 -0.029 -0.173 -0.202
C. roseus -0.02 -0.057 -0.02 -0.02 -0.029 -0.02 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
S. rhombifolia -0.029 0.211 -0.029 -0.029 -0.041 -0.029 -0.029 -0.029 0.699 0.211 -0.051 -0.06
U. sp. -0.02 -0.057 -0.02 -0.02 -0.029 -0.02 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
U. lobata -0.042 0.305 -0.042 -0.042 -0.06 -0.042 -0.042 0.484 -0.042 0.305 -0.074 0.184
A. conyzoides -0.084 0.155 0.241 -0.084 -0.12 -0.084 -0.084 -0.084 0.241 0.023 0.426 0.329
A. subulatus -0.042 0.518 -0.042 -0.042 -0.06 -0.042 -0.042 -0.042 0.484 0.305 -0.074 0.184
B. bipinnata -0.052 -0.148 -0.052 -0.052 -0.075 -0.052 -0.052 -0.052 -0.052 -0.148 -0.093 -0.108
B. pilosa -0.047 -0.134 -0.047 -0.047 -0.068 -0.047 -0.047 0.428 -0.047 0.057 -0.084 -0.098
E. sonchifolia 0.126 0.241 0.126 -0.161 -0.024 0.126 0.126 0.126 -0.161 -0.106 0.223 0.261
E. acer -0.057 0.003 -0.057 0.354 -0.082 -0.057 -0.057 0.354 -0.057 -0.162 -0.101 -0.118
E. chinense 0.161 0.106 -0.126 0.161 -0.18 0.161 -0.126 -0.126 -0.126 -0.125 -0.054 0.035
M. micrantha -0.052 0.205 0.386 -0.052 -0.075 -0.052 -0.052 -0.052 0.386 0.028 0.684 0.344
P. indica -0.02 0.354 -0.02 -0.02 -0.029 -0.02 -0.02 -0.02 1 0.354 -0.036 -0.042
V. cinerea -0.052 0.205 -0.052 -0.052 -0.075 -0.052 -0.052 0.386 0.386 0.205 0.165 0.344
W. triloba -0.047 -0.134 -0.047 0.428 0.272 -0.047 -0.047 0.428 -0.047 0.249 -0.084 -0.098
J. linifolia -0.029 0.211 -0.029 -0.029 -0.041 -0.029 -0.029 -0.029 -0.029 0.211 -0.051 -0.06
P. triquetrum 1 -0.057 -0.02 -0.02 -0.029 -0.02 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
C. asiatica -0.057 1 -0.057 -0.057 -0.082 -0.057 0.354 -0.057 0.354 0.169 0.14 0.305
C. exaltatus -0.02 -0.057 1 -0.02 -0.029 -0.02 -0.02 -0.02 -0.02 -0.057 0.565 -0.042
C. compressus -0.02 -0.057 -0.02 1 -0.029 -0.02 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
K. brevifolia -0.029 -0.082 -0.029 -0.029 1 -0.029 -0.029 -0.029 -0.029 0.211 -0.051 -0.06
L. cernuum -0.02 -0.057 -0.02 -0.02 -0.029 1 -0.02 -0.02 -0.02 -0.057 -0.036 -0.042
A. philoxeroides -0.02 0.354 -0.02 -0.02 -0.029 -0.02 1 -0.02 -0.02 -0.057 -0.036 -0.042
A. sessilis -0.02 -0.057 -0.02 -0.02 -0.029 -0.02 -0.02 1 -0.02 -0.057 -0.036 -0.042
I. triloba -0.02 0.354 -0.02 -0.02 -0.029 -0.02 -0.02 -0.02 1 0.354 -0.036 -0.042
P. purpurea -0.057 0.169 -0.057 -0.057 0.211 -0.057 -0.057 -0.057 0.354 1 -0.101 0.305
Z. pendula -0.036 0.14 0.565 -0.036 -0.051 -0.036 -0.036 -0.036 -0.036 -0.101 1 0.546
O. corniculata -0.042 0.305 -0.042 -0.042 -0.06 -0.042 -0.042 -0.042 -0.042 0.305 0.546 1
n=50.
Using the ID numbers of species in Table 2, the significant species pairs are listed as bellow
(1,2) (1,3) (1,4) (1,5) (1,6) (1,7) (1,8) (1,9) (1,10) (1,11) (1,12) (1,13) (1,14) (1,15) (1,16) (1,17) (1,18) (1,19)
(1,20) (1,21) (1,22) (1,23) (1,24) (1,25) (1,26) (1,27) (1,28) (1,29) (1,30) (1,31) (1,32) (1,33) (1,34) (1,35)
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(1,36) (1,37) (1,38) (1,39) (1,40) (1,41) (1,42) (1,43) (1,44) (1,45) (1,46) (1,47) (1,48)
(2,3) (2,4) (2,5) (2,6) (2,7) (2,8) (2,9) (2,10) (2,11) (2,12) (2,13) (2,14) (2,15) (2,16) (2,17) (2,18) (2,19) (2,20)
(2,21) (2,22) (2,23) (2,24) (2,25) (2,26) (2,27) (2,28) (2,29) (2,30) (2,31) (2,32) (2,33) (2,34) (2,35) (2,36)
(2,37) (2,38) (2,39) (2,40) (2,41) (2,42) (2,43) (2,44) (2,45) (2,46) (2,47) (2,48)
(3,4) (3,5) (3,6) (3,7) (3,8) (3,9) (3,10) (3,11) (3,12) (3,13) (3,14) (3,15) (3,16) (3,17) (3,18) (3,19) (3,20) (3,21)
(3,22) (3,23) (3,24) (3,25) (3,26) (3,27) (3,28) (3,29) (3,30) (3,31) (3,32) (3,33) (3,34) (3,35) (3,36) (3,37)
(3,38) (3,39) (3,40) (3,41) (3,42) (3,43) (3,44) (3,45) (3,46) (3,47) (3,48)
(4,5) (4,6) (4,7) (4,8) (4,9) (4,10) (4,11) (4,12) (4,13) (4,14) (4,15) (4,16) (4,17) (4,18) (4,19) (4,20) (4,21)
(4,22) (4,23) (4,24) (4,25) (4,26) (4,27) (4,28) (4,29) (4,30) (4,31) (4,32) (4,33) (4,34) (4,35) (4,36) (4,37)
(4,38) (4,39) (4,40) (4,41) (4,42) (4,43) (4,44) (4,45) (4,46) (4,47) (4,48)
(5,6) (5,7) (5,8) (5,9) (5,10) (5,11) (5,12) (5,13) (5,14) (5,15) (5,16) (5,17) (5,18) (5,19) (5,20) (5,21) (5,22)
(5,23) (5,24) (5,25) (5,26) (5,27) (5,28) (5,29) (5,30) (5,31) (5,32) (5,33) (5,34) (5,35) (5,36) (5,37) (5,38)
(5,39) (5,40) (5,41) (5,42) (5,43) (5,44) (5,45) (5,46) (5,47) (5,48)
(6,7) (6,8) (6,9) (6,10) (6,11) (6,12) (6,13) (6,14) (6,15) (6,16) (6,17) (6,18) (6,19) (6,20) (6,21) (6,22) (6,23)
(6,24) (6,25) (6,26) (6,27) (6,28) (6,29) (6,30) (6,31) (6,32) (6,33) (6,34) (6,35) (6,36) (6,37) (6,38) (6,39)
(6,40) (6,41) (6,42) (6,43) (6,44) (6,45) (6,46) (6,47) (6,48)
(7,8) (7,9) (7,10) (7,11) (7,12) (7,13) (7,14) (7,15) (7,16) (7,17) (7,18) (7,19) (7,20) (7,21) (7,22) (7,23) (7,24)
(7,25) (7,26) (7,27) (7,28) (7,29) (7,30) (7,31) (7,32) (7,33) (7,34) (7,35) (7,36) (7,37) (7,38) (7,39) (7,40)
(7,41) (7,42) (7,43) (7,44) (7,45) (7,46) (7,47) (7,48)
(8,9) (8,10) (8,11) (8,12) (8,13) (8,14) (8,15) (8,16) (8,17) (8,18) (8,19) (8,20) (8,21) (8,22) (8,23) (8,24) (8,25)
(8,26) (8,27) (8,28) (8,29) (8,30) (8,31) (8,32) (8,33) (8,34) (8,35) (8,36) (8,37) (8,38) (8,39) (8,40) (8,41)
(8,42) (8,43) (8,44) (8,45) (8,46) (8,47) (8,48)
(9,10) (9,11) (9,12) (9,13) (9,14) (9,15) (9,16) (9,17) (9,18) (9,19) (9,20) (9,21) (9,22) (9,23) (9,24) (9,25)
(9,26) (9,27) (9,28) (9,29) (9,30) (9,31) (9,32) (9,33) (9,34) (9,35) (9,36) (9,37) (9,38) (9,39) (9,40) (9,41)
(9,42) (9,43) (9,44) (9,45) (9,46) (9,47) (9,48)
(10,11) (10,12) (10,13) (10,14) (10,15) (10,16) (10,17) (10,18) (10,19) (10,20) (10,21) (10,22) (10,23) (10,24)
(10,25) (10,26) (10,27) (10,28) (10,29) (10,30) (10,31) (10,32) (10,33) (10,34) (10,35) (10,36) (10,37) (10,38)
(10,39) (10,40) (10,41) (10,42) (10,43) (10,44) (10,45) (10,46) (10,47) (10,48)
(11,12) (11,13) (11,14) (11,15) (11,16) (11,17) (11,18) (11,19) (11,20) (11,21) (11,22) (11,23) (11,24) (11,25)
(11,26) (11,27) (11,28) (11,29) (11,30) (11,31) (11,32) (11,33) (11,34) (11,35) (11,36) (11,37) (11,38) (11,39)
(11,40) (11,41) (11,42) (11,43) (11,44) (11,45) (11,46) (11,47) (11,48)
(12,13) (12,14) (12,15) (12,16) (12,17) (12,18) (12,19) (12,20) (12,21) (12,22) (12,23) (12,24) (12,25) (12,26)
(12,27) (12,28) (12,29) (12,30) (12,31) (12,32) (12,33) (12,34) (12,35) (12,36) (12,37) (12,38) (12,39) (12,40)
(12,41) (12,42) (12,43) (12,44) (12,45) (12,46) (12,47) (12,48)
(13,14) (13,15) (13,16) (13,17) (13,18) (13,19) (13,20) (13,21) (13,22) (13,23) (13,24) (13,25) (13,26) (13,27)
(13,28) (13,29) (13,30) (13,31) (13,32) (13,33) (13,34) (13,35) (13,36) (13,37) (13,38) (13,39) (13,40) (13,41)
(13,42) (13,43) (13,44) (13,45) (13,46) (13,47) (13,48)
(14,15) (14,16) (14,17) (14,18) (14,19) (14,20) (14,21) (14,22) (14,23) (14,24) (14,25) (14,26) (14,27) (14,28)
(14,29) (14,30) (14,31) (14,32) (14,33) (14,34) (14,35) (14,36) (14,37) (14,38) (14,39) (14,40) (14,41) (14,42)
(14,43) (14,44) (14,45) (14,46) (14,47) (14,48)
(15,16) (15,17) (15,18) (15,19) (15,20) (15,21) (15,22) (15,23) (15,24) (15,25) (15,26) (15,27) (15,28) (15,29)
(15,30) (15,31) (15,32) (15,33) (15,34) (15,35) (15,36) (15,37) (15,38) (15,39) (15,40) (15,41) (15,42) (15,43)
(15,44) (15,45) (15,46) (15,47) (15,48)
(16,17) (16,18) (16,19) (16,20) (16,21) (16,22) (16,23) (16,24) (16,25) (16,26) (16,27) (16,28) (16,29) (16,30)
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(16,31) (16,32) (16,33) (16,34) (16,35) (16,36) (16,37) (16,38) (16,39) (16,40) (16,41) (16,42) (16,43) (16,44)
(16,45) (16,46) (16,47) (16,48)
(17,18) (17,19) (17,20) (17,21) (17,22) (17,23) (17,24) (17,25) (17,26) (17,27) (17,28) (17,29) (17,30) (17,31)
(17,32) (17,33) (17,34) (17,35) (17,36) (17,37) (17,38) (17,39) (17,40) (17,41) (17,42) (17,43) (17,44) (17,45)
(17,46) (17,47) (17,48)
(18,19) (18,20) (18,21) (18,22) (18,23) (18,24) (18,25) (18,26) (18,27) (18,28) (18,29) (18,30) (18,31) (18,32)
(18,33) (18,34) (18,35) (18,36) (18,37) (18,38) (18,39) (18,40) (18,41) (18,42) (18,43) (18,44) (18,45) (18,46)
(18,47) (18,48)
(19,20) (19,21) (19,22) (19,23) (19,24) (19,25) (19,26) (19,27) (19,28) (19,29) (19,30) (19,31) (19,32) (19,33)
(19,34) (19,35) (19,36) (19,37) (19,38) (19,39) (19,40) (19,41) (19,42) (19,43) (19,44) (19,45) (19,46) (19,47)
(19,48)
(20,21) (20,22) (20,23) (20,24) (20,25) (20,26) (20,27) (20,28) (20,29) (20,30) (20,31) (20,32) (20,33) (20,34)
(20,35) (20,36) (20,37) (20,38) (20,39) (20,40) (20,41) (20,42) (20,43) (20,44) (20,45) (20,46) (20,47) (20,48)
(21,22) (21,23) (21,24) (21,25) (21,26) (21,27) (21,28) (21,29) (21,30) (21,31) (21,32) (21,33) (21,34) (21,35)
(21,36) (21,37) (21,38) (21,39) (21,40) (21,41) (21,42) (21,43) (21,44) (21,45) (21,46) (21,47) (21,48)
(22,23) (22,24) (22,25) (22,26) (22,27) (22,28) (22,29) (22,30) (22,31) (22,32) (22,33) (22,34) (22,35) (22,36)
(22,37) (22,38) (22,39) (22,40) (22,41) (22,42) (22,43) (22,44) (22,45) (22,46) (22,47) (22,48)
(23,24) (23,25) (23,26) (23,27) (23,28) (23,29) (23,30) (23,31) (23,32) (23,33) (23,34) (23,35) (23,36) (23,37)
(23,38) (23,39) (23,40) (23,41) (23,42) (23,43) (23,44) (23,45) (23,46) (23,47) (23,48)
(24,25) (24,26) (24,27) (24,28) (24,29) (24,30) (24,31) (24,32) (24,33) (24,34) (24,35) (24,36) (24,37) (24,38)
(24,39) (24,40) (24,41) (24,42) (24,43) (24,44) (24,45) (24,46) (24,47) (24,48)
(25,26) (25,27) (25,28) (25,29) (25,30) (25,31) (25,32) (25,33) (25,34) (25,35) (25,36) (25,37) (25,38) (25,39)
(25,40) (25,41) (25,42) (25,43) (25,44) (25,45) (25,46) (25,47) (25,48)
(26,27) (26,28) (26,29) (26,30) (26,31) (26,32) (26,33) (26,34) (26,35) (26,36) (26,37) (26,38) (26,39) (26,40)
(26,41) (26,42) (26,43) (26,44) (26,45) (26,46) (26,47) (26,48)
(27,28) (27,29) (27,30) (27,31) (27,32) (27,33) (27,34) (27,35) (27,36) (27,37) (27,38) (27,39) (27,40) (27,41)
(27,42) (27,43) (27,44) (27,45) (27,46) (27,47) (27,48)
(28,29) (28,30) (28,31) (28,32) (28,33) (28,34) (28,35) (28,36) (28,37) (28,38) (28,39) (28,40) (28,41) (28,42)
(28,43) (28,44) (28,45) (28,46) (28,47) (28,48)
(29,30) (29,31) (29,32) (29,33) (29,34) (29,35) (29,36) (29,37) (29,38) (29,39) (29,40) (29,41) (29,42) (29,43)
(29,44) (29,45) (29,46) (29,47) (29,48)
(30,31) (30,32) (30,33) (30,34) (30,35) (30,36) (30,37) (30,38) (30,39) (30,40) (30,41) (30,42) (30,43) (30,44)
(30,45) (30,46) (30,47) (30,48)
(31,32) (31,33) (31,34) (31,35) (31,36) (31,37) (31,38) (31,39) (31,40) (31,41) (31,42) (31,43) (31,44) (31,45)
(31,46) (31,47) (31,48)
(32,33) (32,34) (32,35) (32,36) (32,37) (32,38) (32,39) (32,40) (32,41) (32,42) (32,43) (32,44) (32,45) (32,46)
(32,47) (32,48)
(33,34) (33,35) (33,36) (33,37) (33,38) (33,39) (33,40) (33,41) (33,42) (33,43) (33,44) (33,45) (33,46) (33,47)
(33,48)
(34,35) (34,36) (34,37) (34,38) (34,39) (34,40) (34,41) (34,42) (34,43) (34,44) (34,45) (34,46) (34,47) (34,48)
(35,36) (35,37) (35,38) (35,39) (35,40) (35,41) (35,42) (35,43) (35,44) (35,45) (35,46) (35,47) (35,48)
(36,37) (36,38) (36,39) (36,40) (36,41) (36,42) (36,43) (36,44) (36,45) (36,46) (36,47) (36,48)
(37,38) (37,39) (37,40) (37,41) (37,42) (37,43) (37,44) (37,45) (37,46) (37,47) (37,48)
(38,39) (38,40) (38,41) (38,42) (38,43) (38,44) (38,45) (38,46) (38,47) (38,48)
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(39,40) (39,41) (39,42) (39,43) (39,44) (39,45) (39,46) (39,47) (39,48)

(40,41) (40,42) (40,43) (40,44) (40,45) (40,46) (40,47) (40,48)
(41,42) (41,43) (41,44) (41,45) (41,46) (41,47) (41,48)
(42,43) (42,44) (42,45) (42,46) (42,47) (42,48)
(43,44) (43,45) (43,46) (43,47) (43,48)
(44,45) (44,46) (44,47) (44,48)
(45,46) (45,47) (45,48)
(46,47) (46,48)
(47,48)
3.5 Cluster analysis

The fuzzy cluster trees for family classification and species classification are indicated in Fig. 4 and 5
respectively. ID numbers for families and species can be found in Table 1 and Table 2.
It can be found from Table 1 and 2 that in the family classification, the family pair,
Euphorbiaceae-Amaranthaceae is the most associated, seconded by the family pair,
Commelinaceae-Oxalidaceae. In the species classification, the species pairs, S. sebiferum-A. sessilis, D.
gangeticum-S. rhombifolia, and P. indica-I. triloba, are the most associated.
Fig. 4 Fuzzy cluster tree for family classification.
4 Discussion
In summary, most of the significant associations between grass species/families are positive. Similar to the
results of Zhang (2011), the present study further proves that most associations are weak; positive associations
accounted for the most of the significant associations. It means that the competition/interference between grass
species/families is not significant.
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Fig. 5 Fuzzy cluster tree for species classification.
Spearman correlation (association) is a quasi-linear association (Zhang, 2011, 2012a, 2012b). A

significant Pearson correlation (association) usually means a strong Spearman rank correlation (association)
but a weak Pearson correlation (association) does not mean that Spearman rank correlation (association) is
certainly weak. A significant Spearman rank correlation (association) may likely implicit a stronger linear
correlation (association). A weak Spearman rank correlation (association) will certainly mean a weak linear
correlation (association).
We always want to know direct interspecific associations (Schoener, 1993). A direct interspecific
association (actually the interspecific interaction in this case) can be obtained by using net correlation measure.
The other measures, Pearson correlation, Spearman rank correlation and point correlation may yield both
indirect and direct associations. So net correlation is particullarly suggested for using in such studies. To
achieve better results, in particular for that from net correlation analysis, more samples are suggested to be
collected in the sampling survey.
For plant communities, a direct and positive association (direct interaction) means the facilitation (Bruno
et al., 2003; Ollerton et al., 2003), mutualism, commensalism, proto-cooperation, complementary
resource-partitioning, or parasitism (it rarely occurs in the nature) between two plant species (Callaway, 1995;
Palmer et al., 2003; Hegland et al., 2009), if the possibility for sharing the similar requirements of
environmental conditions has been excluded. However, the negative association in the same case means the
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interspecific competition or inerspecfic interference (excretion of poisonous chemicals, etc.), or two species
have different resource-utilizing patterns: They require different habitats and their ecological niches are
separated, and co-existence of the two species in the same community is nearly impossible.
Community succession is the process of change in the species structure of an ecological community over
time. It is a self-organizing process by which an ecological community undergoes orderly and predictable
changes following initial colonization of new habitat or a disturbance (Wikipedia:
http://en.wikipedia.org/wiki/Ecological_succession). I hold that species with significant positive or negative
associations are generally keystone species in the community. About why the significant positive associations
(interactions), with respect to significant negative associations, always dominate the community, a reasonable
explanation is that although both negative and positive associations (interactions) occur in the process of
assembly and successsion of community, the adaptation and selection will finally result in the successful
coexistence of the species with significant positive associations in the climax community (Fig. 6). Dominance
of significant positive associations (interactions) means the relative stablility and equilibrium of the
community. According to Clements (1916), community succession involves several phases as follows, and
here I use “-“ for significant negative association, “+” for significant positive association and “*” for no
significant association in order to show the change of negative vs. positive associations
Fig. 6 Illustration of change of significant positive/negative interspecific associations. No significant associations account for the
majority of possible interspecific associations.
(1) Nudation (******): Succession begins with the development of a bare site, i.e., disturbance.
(2) Migration (**********-+): It refers to arrival of propagules.
(3) Ecesis (********---+): It involves establishment and initial growth of vegetation.
(4) Competition (******-----+): As vegetation became well established, grew, and spread, various
species began to compete for space, light and nutrients.
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(5) Reaction (Go to step (1) for re-starting; and near the end stage: ******---+++): During this phase
autogenic changes affect the habitat resulting in replacement of one plant community by another.
(6) Stabilization (******-+++++): Reaction phase leads to development of a climax community.
It should be noted that no significant associations usually account for the majority of possible
interspecific associations at each phase of community succession. They guarantee the robustness of community.
They are candidates of keystone species, i.e., lose of some existing keystone species might be filled with some
species previously with no significant associations.
Acknowledgment
I would like to thank all students of Department of Ecology, Sun Yat-sen University, for their field sampling in
Zhuai.
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Book Review
A review on the book, Ecological Modeling
GuangHua Liu
Guangdong AIB Polytech College, Guangzhou 510507, China
E-mail: ghliu@gdaib.edu.cn
Received 15 April 2014; Accepted 30 May 2014; Published online 1 September 2014
Abstract
The book, Ecological Modeling, edited by WenJun Zhang and published by Nova Science Publishers, USA,
was briefly reviewed in present report.
Keywords network biology; book; review.
Selforganizology
1 Introduction
Ecological modelling is a fast growing science. It is a powerful tool to handle complex ecological problems.
More and more innovative methodologies and theories on ecological modelling are emerging around the world.
Most of them have been published on interantional journals. Due to less page limitations, however, some
quality papers may also be published in a careful planned book. The book, Ecological Modelling, is published
to reflect recent achievements of scientists in ecological modelling. This book includes a lot of models,
methods and theories on ecological modelling, for example, artificial neural networks, individual-based
modelling, ecological nich models, landscape and GIS modelling, population dynamics, nutritional ecology,
remote sensing, decision support systems, etc.
Articles in this book are contributed by nearly 40 ecologists, geo-scientists, physicist and mathematicians
from the United States, Canada, China, Australia, Austria, Brazil, Russia, Greece, Italy, Spain, Portugal and
Tunis, etc. It will provide researchers with diverse aspects of the latest advances in ecological modelling. It is a
valuable reference for the scientists, university teachers and graduate students in ecology, environmental
sciences, geography and computational science.
2 Contents
The book covered such contens as follows
Chapter 1 (Zhang and Liu, 2012): Probability distribution functions have been widely used to model the
spatial distribution of arthropods. Aggregation types (i.e., randomly distributed, uniformly distributed,
aggregately distributed, etc.) of arthropods can be detected based on probability distribution functions, but the
abundance at given location is not able to be predicted by them. This study aimed to present an artificial neural
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network to simulate spatial distribution of arthropods. Response surface model and spline function were
compared and evaluated against the neural network model for their simulation performance. The results
showed that the artificial neural network exhibited good simulation performance. Simulated spatial distribution
was highly in accordant with the observed one. Overall the neural network performed better in the case of
lower total abundance of arthropods. Response surface model could fit the spatial distribution of arthropods
but the simulation performance was worse than neural network. Cross validation revealed that neural network
performed better than response surface model and spline function in predicting spatial distribution of
arthropods. Confidence interval of predicted abundance could be obtained using randomized submission of
quadrate sequences in the neural network simulation. It is concluded that artificial neural network is a valuable
model to simulate the spatial distribution of arthropods.
Chapter 2 (Petropoulos et al., 2012): Remote sensing has generally demonstrated a great potential in
mapping spatial patterns of vegetation. By employing the amount of reflected radiation at particular regions of
the electromagnetic spectrum, it is possible to make estimates on certain characteristic of vegetation. The use
of radiometric vegetation indices is a fast and efficient method for vegetation monitoring, explotiting
information acquired from remote sensing data. These indices are dimensionless, radiometric measures that
function as indicators of relative abundance and activity of green vegetation. Throughout the years, a large
number of multispectral vegetation indices have been formulated. Each has variable degree of efficiency in
estimating one or more vegetation parameters such as, health status, nutrient or water deficiency, crop yield,
vegetation cover fraction, leaf area index, absorbed photosynthetically active radiation, net primary production
and above-ground biomass. Additionally some of them also consider atmospheric effects and/ or the soil
background for an enhanced retrieval. The present chapter aims in providing an overview on the use of
radiometric vegetation indices developed over the last few decades utilizing spectral information acquired
from multispectral remote sensing sensors. Before that, an introduction to some important principles of remote
sensing relevant to the vegetation spectral response is made available, as this was considered necessary to
better understand the context of the present overview.
Chapter 3 (Altartouri et al., 2012): According to the Water Framework Directive (WFD, 2000/60/EC),
Integrated River Basin Management Plans (RBMP) are required at different scales, in order to prevent amongst
other things, water resource deterioration and ensure water pollution reduction. An integrated river basin
management approach underpins a risk-based land management framework for all activities within a spatial
land-use planning framework. To this end, a risk assessment methodology is required to identify water
pollution hazards in order to set appropriate environmental objectives and in turn design suitable mitigation
measures. Surface water pollution as a result of Olive Mill Waste (OMW) discharge is a serious hazard in the
olive oil producing regions of the Mediterranean. However, there is no standardised method to assess the risk
of water pollution from olive mill waste for any given river basin. The present chapter shows the results from a
study conducted addressing the above issue by designing a detailed risk assessment methodology, which
utilises GIS modelling to classify within a watershed individual sub-catchment risk of water pollution
occurring from olive mill waste discharges. The chapter presents the proposed criteria and calculations
required to estimate sub-catchment risk significance and comments on the methods potential for wider
application. It combines elements from risk assessment frameworks, Multi Criteria Analysis (MCA), and
Geographic Information Systems (GIS). MCA is used to aggregate different aspects and elements associated
with this environmental problem, while GIS modeling tools helped in obtaining many criterion values and
providing insight into how different objects interact in nature and how these interactions influence risk at the
watershed level. The proposed method was trialled in the Keritis watershed in Crete, Greece and the results
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indicated that this method has the potential to be a useful guide to prioritise risk management actions and
mitigation measures which can subsequently be incorporated in river basin management plans.
Chapter 4 (Nedorezov, 2012): Publication is devoted to the problem of population time series analysis
with various discrete time models of population dynamics. Applications of various statistical criterions, which
i
are normally used for determination of mathematical model parameters, are u nder the discussion. With a
particular example on green oak leaf roller (Tortrix viridana L.) population fluctuations, which had been
presented in publications by Rubtsov (1992), and Korzukhin and Semevskiy (1992) for three different
locations in Europe, the possibilities of considering approach to the analysis of population dynamics are
demonstrated. For approximations of empirical datasets the well-known models of population dynamics with a
discrete time (Kostitzin model, Skellam model, Moran – Ricker model, Morris – Varley – Gradwell model,
and discrete logistic model) were applied. For every model the final decision about the possibility to use the
concrete model for approximation of datasets are based on analyses of deviations between theoretical (model)
and empirical trajectories: the correspondence of distribution of deviations to Normal distribution with zero
average was checked with Kolmogorov – Smirnov and Shapiro – Wilk tests, and existence/absence of serial
correlation was determined with Durbin – Watson criteria. It was shown that for two experimental trajectories
Kostitzin model and discrete logistic model give good approximations; it means that population dynamics can
be explained as a result of influence of intra-population self-regulative mechanisms only. The third considering
empirical trajectory needs in use more complicated mathematical models for fitting.
Chapter 5 (Cianelli et al., 2012): In the last decades, numerical modelling has gained increasing
consensus in the scientific world, and particularly in the framework of behavioural and population ecology.
Through numerical models it is possible to reconstruct what is observed in the environment or in the laboratory
and to get a more in-depth comprehension of the factors regulating the phenomena under examination.
Numerous approaches have been developed in this framework, but probably one of the most promising is the
individual-based modelling. With this type of approach it is relatively straightforward to investigate aspects
related to the ecology of a population starting from the characterisation of processes taking place at the scale of
the individual organism. This contribution is intended to provide a general view of the main features of the
individual-based models and of their peculiarities in comparison to other modelling strategies. Special
emphasis will be given to applications in the field of phyto- and zooplankton ecology and behaviour, and
results from the available literature on this topic will be used as examples.
Chapter 6 (Watts et al., 2012): The larval phase of most blowfly species is considered a critical
developmental period in which intense limitation of feeding resources frequently occurs. Furthermore, such a
period is characterised by complex ecological processes occurring at both individual and population levels.
These processes have been analysed by means of traditional statistical techniques such as simple and multiple
linear regression models. Nonetheless, it has been suggested that some important explanatory variables could
well introduce non-linearity into the modelling of the nutritional ecology of blowflies. In this context, dynamic
aspects of the life history of blowflies could be clarified and detailed by the deployment of machine learning
approaches such as artificial neural networks (ANNs), which are mathematical tools widely applied to the
resolution of complex problems. A distinguishing feature of neural network models is that their effective
implementation is not precluded by the theoretical distribution of the data used. Therefore, the principal aim of
this investigation was to use neural network models (namely multi-layer perceptrons and fuzzy neural
networks) in order to ascertain whether these tools would be able to outperform a general quadratic model (that
is, a second-order regression model with three predictor variables) in predicting pupal weight values (output)
of experimental populations of Chrysomya megacephala (F.) (Diptera: Calliphoridae), using initial larval
density (number of larvae), amount of available food, and pupal size as input variables. These input variables
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may have generated non-linear variation in the output values, and fuzzy neural networks provided more
accurate outcomes than the general quadratic model (i.e. the statistical model). The superiority of fuzzy neural
networks over a regression-based statistical method does represent an important fact, because more accurate
models may well clarify several intricate aspects regarding the nutritional ecology of blowflies. Additionally,
the extraction of fuzzy rules from the fuzzy neural networks provided an easily comprehensible way of
describing what the networks had learned.
Chapter 7 (Newton, 2012): An ecological-based decision-support system and corresponding algorithmic
analogue for managing natural black spruce (Picea mariana (Mill) BSP.) and jack pine (Pinus banksiana
Lamb.) mixed stands was developed. The integrated hierarchical system consisted of six sequentially-linked
estimation modules. The first module consisted of a key set of empirical yield-density relationships and
theoretically-based functions derived from allometry and self-thinning theory that was used to describe overall
stand dynamics including temporal size-density interrelationships and expected stand development trajectories.
The second module was comprised of a Weibull-based parameter prediction equation system and an
accompanying composite height-diameter function which was used to recover diameter and height
distributions. The third module included a set of species-specific composite taper equations which was used to
derive log product distributions and volumetric yields. The fourth module was composed of a set of
species-specific allometric-based composite biomass equations which was used to estimate mass distributions
and associated carbon-based equivalents for each above-ground component (bark, stem, branch and foliage).
The fifth module incorporated a set of species-specific end-product and value equations which was used to
predict chip and lumber volumes and associated monetary equivalents by sawmill type (stud and randomized
length mill configurations). The sixth module encompassed a set of species-specific composite equations that
was used to derive wood and log quality metrics (specific gravity and mean maximum branch diameter,
respectively). The stand dynamic and structural recovery modules were developed employing 382 stand-level
measurements derived from 155 permanent and temporary sample plots situated throughout the central portion
of the Canadian Boreal Forest Region, the taper and end-product modules were developed employing
published results from taper and sawmill simulation studies, and the biomass and fibre attribute modules were
developed using data from density control experiments. The potential of the system in facilitating the
transformative change towards the production of higher value end-products and a broader array of ecosystem
services was exemplified by simultaneously contrasting the consequences of density management regimes
involving commercial thinning treatments, in terms of overall productivity, end-product yields, economic
efficiency, and ecological impact. This integration of quantitative relationships derived from applied ecology,
plant population biology and forest science into a common analytical platform, illustrates the synergy that can
be realized through a multi-disciplinary approach to forest modeling.
Chapter 8 (Barbosa et al., 2012): The authors present a review of the concepts and methods associated to
ecological niche modeling illustrated with the published works on amphibians and reptiles of the
Mediterranean Basin, one of the world's biodiversity hotspots for conservation priorities. The authors start by
introducing ecological niche models, analyzing the various concepts of niche and the modeling methods
associated to each of them. The authors list some conceptual and practical steps that should be followed when
modeling, and highlight the pitfalls that should be avoided. The authors then outline the history of ecological
modeling of Mediterranean amphibians and reptiles, including a variety of aspects: identification of the
ecological niche; detection of common distribution areas (chorotypes) and other biogeographical patterns;
analysis and prediction of species richness patterns; analysis of the expansion of native and invasive species;
integration of molecular data with spatial modeling; identification of contact zones between related taxa;
assessment of species' conservation status; and prediction of future conservation problems, including the
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effects of global change. The authors conclude this review with a discussion of the research that still needs to
be developed in this area.
Chapter 9 (Krivtsov and Jago, 2012): Numerical techniques (e.g. correlation, multiple regression and
factor analysis, path analysis, methods of network analysis, and, in particular, simulation modelling) may be
very helpful in investigations of indirect relationships in aquatic ecosystems. Here the authors give a brief
overview of some examples of the relevant studies, and focus on 1) a case study of a freshwater eutrophic lake,
where statistical analysis of the datasets obtained within a comprehensive monitoring programme, and
sensitivity analysis by a mathematical model ‘Rostherne’, helped to reveal the previously overlooked
relationships between Si and P biogeochemical cycles coupled through the dynamics of primary producers,
and 2) give an overview of how the coupling of physical, chemical, and biological processes in the marine
ecosystem models offers a basis for investigations of indirect interactions in continental shelf seas. Complex
aquatic ecosystem models provide a numerical simulation of biogeochemical fluxes underpinned by coupling
physical forcing functions with definitions simulating biological and chemical processes, and offer a potential
for quantitative interpretation of sediment proxies in the stratigraphic record. Combination of models and
sediment proxies, calibrated by training sets, can provide information on water column structure, surface
heating, mixing, and water depth, thus providing a basis for reconstruction of the past, and predicting the
future environmental dynamics.
Chapter 10 (Schmickl et al., 2012): In the evolution of social insects, the colony and not the (often sterile)
individual worker should be considered the major unit of selection. Thus, social insect colonies are considered
to be 'super-organisms', which have – like all other organisms – to perform behaviors which affect their outside
environment and which alter their own future internal status. The way these behaviors are coordinated is by
means of communication, which is either direct or indirect and which involves information exchange either by
transmitting signals or by exploiting cues. Therefore, social insect colonies perform information processing in
a rather similar way as multicellular organisms do, where behaviors result from the exchange of information
among their sub-modules (cells). In many cases, self-organization allows a colony to evaluate massive
amounts of information in parallel and to decide about the colony's future behavioral responses. Many
feedback systems that govern self-organization of workers have been investigated empirically and theoretically.
Here, the authors discuss models which have been proposed to explain division of labor and task selection in
social insects. The authors demonstrate how the collective regulation of labor in eusocial insect colonies is
studied by means of top-down modeling and by bottom-up models, often analyzed with multi-agent computer
simulations.
Chapter 11 (Gamez, 2012): The paper is a review of a research line initiated two decades ago. At the
beginning the research was concentrated on basic qualitative properties of ecological and population-genetic
models, such as observability and controllability. For population system, observability means that, e.g. from
partial observation of the system (observing only certain indicator species), in principle the whole state process
can be recovered. Recently, for different ecosystems, the so-called observer system (or state estimators) have
been constructed that enables us to effectively estimate the whole state process from the observation. The
methodology of observer design can be also applied to estimate unknown changes in ecological parameters of
the system. Clearly, both observation (i.e. monitoring) and control are important issues in conservation
ecology. For an ecological system, in an appropriate setting, controllability implies that a disturbed ecosystem
can be steered beck to an equilibrium state by an abiotic human intervention. Recent research concern the
effective calculation of such control functions. While the considered ecological models are density-dependent,
observability and controllability problems also naturally arise in frequency-dependent models of population
genetics. As for the frequency-dependent case, observation systems typically occur in case of phenotypic
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observation of genetic processes; control systems can be used to model e.g. artificial selection. In this survey,
in addition to the basic methodology and its applications, the recent developments of the field are also
reported.
Chapter 12 (Spagnolo et al., 2012): The authors analyse the effects of environmental noise in three
different biological systems: (i) mating behaviour of individuals of Nezara viridula; (ii) polymer translocation
in crowded solution; (iii) an ecosystem described by a Verhulst model with a multiplicative Lévy noise.
Specifically, the authors report on experiments on the behavioural response of N. viridula individuals to
sub-threshold deterministic signals in the presence of noise. The authors analyse the insect response by
directionality tests performed on a group of male individuals at different noise intensities. The percentage of
insects which react to the sub-threshold signal shows a non-monotonic behavior, characterized by the presence
of a maximum, for increasing values of the noise intensity. This is the signature of the non-dynamical
stochastic resonance phenomenon. By using a ”hard” threshold model the authors find that the maximum of
the output cross correlation occurs in the same range of noise intensity values for which the behavioral
activation shows a maximum. In the second system, the noise driven translocation of short polymers in
crowded solutions is analyzed. An improved version of the Rouse model for a flexible polymer has been
adopted to mimic the molecular dynamics, by taking into account both the interactions between adjacent
monomers and introducing a Lennard-Jones potential between non-adjacent beads. A bending recoil torque has
also been included in our model. The polymer dynamics is simulated in a two-dimensional domain by
numerically solving the Langevin equations of motion. Thermal fluctuations are taken into account by
introducing a Gaussian uncorrelated noise. The mean first translocation time of the polymer centre of inertia
shows a minimum as a function of the frequency of the oscillating forcing field. In the third ecosystem, the
transient dynamics of the Verhulst model perturbed by arbitrary non-Gaussian white noise is investigated.
Based on the infinitely divisible distribution of the Lévy process the authors study the nonlinear relaxation of
the population density for three cases of white non-Gaussian noise: (i) shot noise, (ii) noise with a probability
density of increments expressed in terms of Gamma function, and (iii) Cauchy stable noise. The authors obtain
exact results for the probability distribution of the population density in all cases, and for Cauchy stable noise
the exact expression of the nonlinear relaxation time is derived. Moreover starting from an initial delta
function distribution, the authors find a transition induced by the multiplicative Lévy noise, from a trimodal
probability distribution to a bimodal probability distribution in asymptotics. Finally the authors find a
nonmonotonic behavior of the nonlinear relaxation time as a function of the Cauchy stable noise intensity.
Chapter 13 (Ferrarini, 2012): Landscape modelling is founded on the idea that the patterning of landscape
elements strongly influences ecological characteristics, thus the ability to quantify landscape structure is a
prerequisite to the study of landscape function and change over time as well. For this reason, much emphasis
has been placed until now on developing methods to quantify landscape structure. Unfortunately, on one side
landscape (i.e., landcover or landuse) and vegetation maps are very complex mosaics of thousands of patches,
and this makes the interpretation of their structure very challenging. On the other side, methods developed so
far to quantify landscape structure just return numerical results, that are not linked to cartographic outputs. Last,
landscape pattern indices are numerous, and the need for a synthetic representation is more and more impelling.
The author provide here the description and application of a novel approach to landscape structural modelling
based on the combined use of GIS (Geographical Information Systems) and multivariate statistics. First,
landscape structure of the study area (Ceno valley, Italy) is analyzed through 5 patch-based, non-redundant
indicators (area, isolation, compactness, shape complexity, interspersion) with indirect link to functional
aspects. Second, PCA (principal component analysis) is used in order to synthesize structural indicators, and
cartographic output is given. Third, KCA (k-means cluster analysis) is applied in order to group landscape
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patches into homogeneous clusters, and again GIS output is supplied. Last, LDA (linear discriminant analysis)
is employed to provide evidence for the differences among clusters. This modelling approach provides the
chance for a deep and cost-effective exegesis of landscape structure, with promising consequences on
conjecture formulation about functional aspects as well.
Chapter 14 (Sellami, 2012): Our days, the climatic change, manifested by strong and brutal precipitation,
violent wind and long drought, has as direct consequence to damage the plant canopies (forests, sylviculture,
oasis, pastoral lands and agricultural fields) so menacing the human feeding either from plants or animals
(caprine, ovine, bovine, cameline..), exhausting the water resources, increasing the need for energy in
buildings used for all activities (industrial, agricultural and services). Which solution the ecological modelling
is capable to participate with, at short and long dated, in order to buffer the climatic change effect and to
assume the need of food and clean energy for human? In this chapter the authors present the basic concepts to
model the plant architecture (species, densities, positions and orientation) the most adaptable to the sudden
calamity, the energy use efficiency in building (material of construction, isolation system, organisation of
accessories and apparatus), and the produce of clean energy from the wind velocity (founding wind sources
and evaluating regional wind potential offshore and inshore, conceptualising wind turbine and testing their
efficiencies
References
Altartouri A, Pediaditi K, Petropoulos GP, et al. 2012. Development of a decision support system for the
estimation of surface water pollution risk from olive mill waste discharges. In: Ecological Modeling
(WenJun Zhang, ed). 41-64, Nova Science Publishers, New York, USA
Barbosa AM, Sillero N, Martínez-Freiría F, et al. 2012. Ecological niche models in mediterranean herpetology:
past, present, and future. In: Ecological Modeling (WenJun Zhang, ed). 173-204, Nova Science Publishers,
New York, USA
Cianelli D, Uttieri M, Zambianchi E. 2012. Individual based modelling of planktonic organisms. In: Ecological
Modeling (WenJun Zhang, ed). 83-96, Nova Science Publishers, New York, USA
Ferrarini A. 2012. Landscape structural modeling. a multivariate cartographic exegesis. In: Ecological
Gámez M. 2012. Observation and control in density- and frequency-dependent population models. In:
Ecological Modeling (WenJun Zhang, ed). 267-288, Nova Science Publishers, New York, USA
Krivtsov V, Jago CF. 2012. Some aspects of phytoplankton and ecosystem modelling in freshwater and marine
environments: consideration of indirect interactions, and the implications for interpreting past and future
overall ecosystem functioning. In: Ecological Modeling (WenJun Zhang, ed). 205-222, Nova Science
Publishers, New York, USA
Nedorezov LV. 2012. Analysis of green oak leaf roller population dynamics in various locations. In:
Newton PF. Development and utility of an ecological-based decision-support system for managing mixed
coniferous forest stands for multiple objectives. In: Ecological Modeling (WenJun Zhang, ed). 115-172,
Nova Science Publishers, New York, USA
Petropoulos GP, Kalaitzidis C. 2012. Multispectral vegetation indices in remote sensing: An overview. In:
Schmickl T, Crailsheim K. 2012. Modellig population dynamics, division of labour and nutrient economics of
social insect colonies. In: Ecological Modeling (WenJun Zhang, ed). 223-265, Nova Science Publishers,
New York, USA
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Sellami MH. 2012. Basic concepts for modelling in different and complementary ecological fields: plants
canopies conservation, thermal efficiency in buildings and wind energy producing. In: Ecological
Spagnolo B, Valenti D, Spezia S, et al. 2012. Environmental noise and nonlinear relaxation in biological
systems. In: Ecological Modeling (WenJun Zhang, ed). 289-323, Nova Science Publishers, New York,
USA
Watts MJ, Bianconi A, Serapiao ABS, et al. 2012. The effectiveness of artificial neural networks in modelling
the nutritional ecology of a blowfly species. In: Ecological Modeling (WenJun Zhang, ed). 97-114, Nova
Science Publishers, New York, USA
Zhang WJ, Liu GH. 2012. Artificial Neural Network Simulation of Spatial Distribution of Arthropods: A
Multi-model Comparison. In: Ecological Modeling (WenJun Zhang, ed). 1-14, Nova Science Publishers,
New York, USA
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Selforganizology
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Selforganizology
Volume 1, Number 2, 1 September 2014
Articles
Structure and dynamics of Lithocolletis ringoniella-Parasitoids food web in

apple orchards of Shaanxi, China
Xin Li, YuYu Liu 51-61
Two-dimensional ordered cluster analysis of component groups in

self-organization
WenJun Zhang, YanHong Qi, ZhiGuo Zhang 62-77
Diversity and aggregation patterns of plant species in a grass community

Ran Li, DanTing Chen, GongWei Liang, et al. 78-88
Interspecific associations and community structure: A local survey and

analysis in a grass community
WenJun Zhang 89-129
A review on the book, Ecological Modeling

GuangHua Liu 130-137
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Selforganizology

Vol. 1, No. 2, 1 September 2014

International Academy of Ecology and Environmental Sciences

Editorial Office: selforganizology@iaees.org

Publisher: International Academy of Ecology and Environmental Sciences

Structure and dynamics of Lithocolletis ringoniella-Parasitoids food

Xin Li, Yuyu Liu

2 Materials and Methods

3 Results and Analysis

sometimes reaches 41%-98%.

Table 2 Emergence rate of A. theivorae in different days (Yangling, Shaanxi, China,

(2) S. sericeicornis and S. Foerst

Table 5 Parasitic mechanism of four dominant parasitoids of L. ringoniella.

3.2 Dynamics of food web

Fig. 3 Dynamics of L. ringoniella (Yangling, Shaanxi, China, 2008).

Fig. 5 Dynamics of parasitization effect to L. ringoniella (Yangling, Shaanxi, 2008).

discontinuously grows from late May to December.

Date No. A. theivorae S. sericeicornis No. Emergence Cocoon/cystics

4/18 0 0 0 0 0 0 0.00 0.00

3.2.3 Dynamics of parasitization effect

Two-dimensional ordered cluster analysis of component groups in

WenJun Zhang1,2, YanHong Qi3, ZhiGuo Zhang4

Keywords two-dimensional cluster analysis; components; groups.

2.1 Choose distance measures

x2= n..(∑pi=1∑qj=1nij2/(ni. n.j)-1)

yij= yij max j dij i=1,2,…,n

g1(i,j)={k, i | yk12>( yj12- yi12)( yk11- yi11)/( yj11- yi11)+ yi12}

min pbar=(∑i2-1i=i1∑i3-1j=i2 pij)/(( i2- i1)( i3- i2))

Fig. 1 Applet window of the algorithm-Input.

Fig. 2 Applet window of the algorithm-Output.

Fig. 3 Geographical distribution of sampling sites in a meadow of Zhuhai.

P_test value=0.0017<=0.05, statistically significant

P_test value=0.2045>0.05, not statistically significant

P_test value=0.3349>0.05, not statistically significant

(74 ) (59 69 70 78 79 80 88 90 ) (81 82 83 84 85 86 91 92 93 100 ) (89 ) (99 ) (98 ) (97 ) (96 )

Fig. 4 indicates 2-dimensional classification for p=0.0482 (<=0.05, statistically significant).

Fig. 4 Classification for p=0.0482 (<=0.05, statistically significant).

3.2 Two-dimensional vegetation distribution in Ngari

Fig. 5 Geographical distribution of sampling sites in Ngari.

P_test value=0.0028<=0.05, statistically significant

Between-sampling site p values matrix is listed as the following:

Diversity and aggregation patterns of plant species in a grass

2 Materials and Methods

Fig. 1 The grassland for sampling survey.

Fig. 2 Sampling design.

pr= (k+r-1)!pr / (r!(k-1)!Qk+r) r>0

IV = (relative cover + relative frequency + relative density) / 3

The summed dominance ratio for two factors (SDR2) is expressed as

SDR2= (density ratio + relative frequency) / 2 × 100%

2.2.3 Diversity of community

Simpson index: Sp=1 - Σxj(xj-1) / (N(N-1))

where S: total number of plant species, Pj= xj / N, N: total number of individuals.

Table 1 Aggregation patterns of 47 species.

3.2 Importance values of plant species

Table 2 Importance values of 47 species.

3.3 Diversity of plant species

Plant species SDR2(%) Plant species SDR2(%)

4 Discussion and Main Conclusion

Interspecific associations and community structure: A local survey

2 Materials and Methods

Fig. 1 A profile of the grass community.

If χ2>χ2α(1) (χ20.01(1)=6.64, χ20.05(1)=3.84), then interspecific (inter-family) association is statistically