Documente Academic
Documente Profesional
Documente Cultură
BIBIAN DIWAY
NURUL FARHANA ZAKARIA
SULIANA CHARLES
KEVIN NG KIT SIONG
LUCY CHONG
Copyright 2010
Sarawak Forest Department
Sarawak Forestry Corporation
+6(082) 319158
+6(082) 445639
TABLE OF CONTENTS
ii
LIST OF TABLES
iii
LIST OF FIGURES
iv
ACKNOWLEDGEMENT
EXECUTIVE SUMMARY
vi
1.0 INTRODUCTION
1.1.1
Protection of Ramin
1.1.2
2.0 OBJECTIVES
3.0 METHODOLOGY
3.1.1
3.1.2
DNA Extraction
3.1.3
3.1.4
Data analysis
4.1.1
Genetic Diversity
4.1.2
Populations differentiation
5.0 CONCLUSIONS
11
6.0 RECOMMENDATIONS
11
REFERENCES
12
PHOTOS
14
19
ii
LIST OF TABLES
Table 1: Export destinations for Ramin products (January to November 2009) for
Sarawak.
iii
LIST OF FIGURES
Figure 2: Example of some fragment data showing variable alleles size from three
individuals of Gonystylus bancanus.
10
Figure 4: Cluster of population based on Structure ver 2.3. Population divided into
two subregion: Red and Blue. Population 1: Sedilu;2: Lingga; 3: Serapau;
4: Loagan Bunut; 5: Kayangeran;6: Maludam; 7: Manggut; 8: Betong and
9: Naman.
11
iv
ACKNOWLEDGEMENT
The authors would like to acknowledge Datu Len Talif Salleh, the Director of Sarawak Forest
Department cum CEO of Sarawak Forestry Corporation, Mr. Ngui Siew Kong, the Asistant
Director of Sarawak Forest Department and Mr. Nigel Lim, the General Manager of Sarawak
Forestry Corporation for permitting us to conduct this research. We would like to thank Dr. Lee
Soon Leong from FRIM, managers and colleagues from Applied Forest Science and Industrial
Development of Sarawak Forestry Corporation for their fruitful suggestions and comments. Not
forgetting all staff of Sarawak Forestry Corporation and Forest Research Institute Malaysia
involved in implementing the project. Special thanks also go to Mr Tsuyoshi Harata for assisting
in statistical analysis and Dr. Koichi Kamiya for editing the report, Miss Stephanie Ronny, Mr.
Mohamad Shahbudin Sabki and Mr.Azahari Omar for their assistant in this project, and Rantai
Jawa and Andy Samuel for their photos.
Last but not least we would like to thank ITTO-CITES for funding this project and NRE the
executing agencies.
EXECUTIVE SUMMARY
Gonystylus bancanus popularly known as ramin is an endemic species to Peat Swamp Forest.
High demand for the ramin products and depletion of habitat due to land conversion has put
ramin in the Appendix II CITES - listed species. Various protection measures has been taken by
Sarawak government including imposed on ban on export of ramin log and for export of ramin
products will need verification from Customs on the CITES Export Permit. However the control
measure by the CITES could not guarantee no illegal trading of ramin. Often the illegal trading
happened through false declaration on the species name and origin of the products. This project
is to develop the DNA database for Gonystylus bancanus in Sarawak to enhance the enhance
Sarawak and Malaysia government as well as CITES effort to combat illegal trading. It is an
early stage in the effort to use DNA for tracing and tracking of timber origin and verification of
species.
The plant materials consisted of leaves and barks samples were collected from nine ramin
populations throughout. For extraction of DNA several methods were used including plant
extraction kit however only cetyltrimethyl ammonium bromide (CTAB) method with
modification yielded a good quality of DNA. From a total of 25 microsatellite markers tested, 18
markers were selected to amplify the DNA and to generate alleles frequencies data for the
database. A total of sixty seven unique alleles were detected providing useful information for
tracing of timber origin. Genetic diversity measured was generally high for most populations
except for Kayangeran Forest Reserve and Loagan Bunut National Park. The expected
heterozygosity values ranged from 0.682 to 0.803 indicated that genetic diversity was generally
high. The level of genetic diversity of ramin in Sarawak populations was as high as level of
genetic diversity for other species of tropical forest.
The genetic identity based on Neis 1978 identity index, ranged from 0.62994 between
populations of Kayangeran and Serapau to 0.982971 between populations of Manggut and
Lingga. UPGMA cluster analysis using Neis 1978 genetic distance clustered the populations
into two clusters which positively related to geographical location. Sedilu, Lingga, Serapau,
Maludam, Manggut, Betong and Naman were clustered into one group and the isolated
populations Kayangeran and Loagan Bunut were clustered in another group. The closest genetic
distance was 0.022656 between populations of Manggut and Lingga whereas the furthest was
0.462131 between populations of Kayangeran and Serapau. The high value of Fst indicated the
ramin populations in Sarawak in former days were connected but now becoming fragmented.
vi
Unique alleles detected within populations and cluster analysis indicated the ability and
usefulness of microsatellite markers selected for tracking and tracing of ramin populations in
Sarawak. Thus they are effective for the development of DNA database. However for the high
accuracy and reliability of microsatellite markers for tracking and tracing timber origin and
species identification should base on many loci. In addition, similar project should be extended
to other ramin population in other states and neighboring countries in order to enhance
implementation of CITES legislation/regulations and the enforcement officials capacity to curb
illegal logging within this region.
vii
1.0 Introduction
Gonystylus bancanus locally know as ramin telur belongs to the family Thymelaeaceae is a
native species of fragile Peat Swamp Forest (PSF). The wood is creamy yellow in colour with
moderately fine and even texture, is easy to saw, bore and turn and has smooth planned surface.
It is one of the most valuable tropical hardwoods and highly sought after for decorative cabinet
timber, furniture, squash court flooring and interior decorative work (Lim et al., 1999).
The largest importer of ramin sawn timber and products are countries such as China including
Hong Kong, United States and Japan (Table 1). From January to November 2009 the production
and export of Ramin for Sarawak reported were 2,591 m3, a decline compared to production in
2008, 2007 and 2006 (Table 2). The decline in ramin production for 2009 as reported by
Malaysia during the Fifty-ninth CITES meeting at Doha (Qatar) on 12 March 2010 was the result
of global economic recession faced by the timber sector since late 2008.
Table 1 Export destinations for Ramin products (January to November 2009) for Sarawak
Export destination
Market
China (including Hong Kong)
USA
Japan
Others
Total
Volume (m3)
623
454
448
1,066*
2,591
Ramin products
Products
Sawn Timber
Volume (m3)
1,153
Dowels/Mouldings
1,438
Total
2,591
2006
22,000
12,161
2007
12,875
10,434.31
81.04
2008
3,178
3,063.09 (5,112.1214m3)**
96.38
2009
3,178
2,591
81.53
The reduction in ramin production over the years was also the result of decline in areas covered
by peat swamp forest. Based on the 1995/1996 satellite images, 9.2 % or 1.13 million ha of
Sarawaks land area was covered by Peat Swamp Forest. However, in 2002, the Landsat and
SPOT data showed that 98.5 % of the peat swamp forest has been opened up for various
purposes leaving only 18,920.45 ha relatively untouched PSF (Wong 2002). From the total area
1
opened up, 78,260.96 ha has been converted into plantation area. The total area converted to
plantation was further increased to 389,483 ha in 2004 (Figure provided by Sarawak Forestry
Department GIS unit and reported by Lee 2005).
Due to the reduction in PSF within PFE and the decline in ramin production, Malaysia
particularly Sarawak has been reported by Environmental Investigation Agency (EIA) and
Telapak of being involved in smuggling and laundering of illegal Indonesian ramin into
number of importing countries, including US and UK (Traffic 2004, EIA/Telapak 2004). Worse
still, Malaysia has also been accused of failing to carry out a proper non-detriment finding for
ramin export and has no effective control in place on ramin harvesting (EIA/Telapak 2006). The
Malaysian government through Malaysia Timber Council (MTC, 2004) responded that the
country does not condone any illegal trade in timber and has implemented various measures to
combat the entry of illegal logs.
database developed for N. heimii can serve for the random match probability estimation of an
illegal logging with its original stumps (Tnah et al., 2010).
2.0 Objective
The aims of the study are; (i) to develop a DNA database as initial stage to enhance efforts of
tracing and tracking of Gonystylus bancanus timber, (ii) to estimate the genetic diversity of
ramin populations in Sarawak, and (iii) to determine extents of genetic differentiation among
populations that contribute to baseline information for the tracing and tracking system using
DNA markers.
3.0 Methodology
3.1 Study sites and samples collection
Samples were collected from known geographical range of peat swamp forest in Sarawak (Fig.
1). Peat swamp areas in the central part of Sarawak were not surveyed for this project because
the areas are earmarked for further development. A total of nine populations were sampled
(Table 3). At least 30 individuals were collected from each population, though few areas
involved intensive search due to limited stand density remaining.
Kayangeran
Loagan
Bunut
Sedilu
Lingga
Serapau
Manggut
Maludam
Betong
Naman
Locality
Nearest
town
Latitude
Longitude
1
2
3
4
Sedilu
Lingga
Serapau
Loagan Bunut
Simunjan
Sri Aman
Sri Aman
Miri
N01022'
N01021'
N01019'
N03047'
E110049'
E111010'
E111012'
E114014'
No
of
individuals
sampled
31
30
30
101
5
6
7
8
Kayangeran
Maludam
Manggut
Betong
Lawas
Sebuyau
Betong
Betong
N4o53
N01o38
N01o29
N01o 25
E115o25
E111o 03
E111o 16
E111o 23
29
36
53
40
Naman
Sibu
N02o11
E111o 50E
55
FSTAT, ver. 2.9.3.2 (Goudet,2002) was used to calculate the allelic richness, Ar (Petit et al.,
1998), Wrights (1951) F statistics (Fst, Fis and Fit) based on Weir and Cockerhams (1984) to
determine the distribution of genetic variation within and among populations and R-statistics,
Rst, (Slatkin,1995; Goodman,1997) calculation, to estimate a population differentiation. Fit and
Fis measure heterozygosity of individual relative to the total population and its subpopulations,
respectively; Fst is a measure of genetic differentiation among populations.
Cluster analysis based on genetic distances estimated from Neis (1978) index of similarity was
performed using Genetic Data Analysis (GDA) program (Lewis and Zaykin, 2001). The
population dendrograms were performed with the tree drawing software TreeView (Page, 2001).
Sedilu
Lingga
Serapau
Gba082
Gba092
Gba100
Gba108
Gba129
Gba147
Gba175
Gba348
Gba430
WGb17
WGb22
WGb24
WGb29
WGb32
WGb37
WGb38
WGB39
Total
19
9
11
21
14
16
2
7
13
13
12
9
12
19
6
7
8
207
20
11
6
22
15
17
2
7
13
14
11
10
15
17
6
6
8
206
21
9
9
19
14
16
3
11
11
14
11
7
14
15
6
7
6
200
Loagan
Bunut
16
7
5
14
11
16
2
10
11
16
6
10
9
16
5
5
4
168
Kayangeran
Maludam
Manggut
Betong
Naman
Total
8
4
4
8
10
14
2
6
7
7
2
7
4
10
6
3
4
109
20
9
10
19
12
20
2
9
11
15
10
10
14
16
6
7
6
202
22
11
10
21
15
24
2
11
11
17
14
11
15
20
7
6
6
227
23
7
10
24
16
20
2
11
12
15
11
10
19
17
7
6
7
222
24
9
12
21
17
21
3
9
18
17
11
9
17
20
7
9
8
239
32
15
18
35
25
37
3
17
20
24
15
17
27
28
9
11
11
356
A total of 356 alleles were detected from 18 polymorphic loci (Table 4). The average number of
alleles and allelic richness was relatively high for most populations except for Kayangeran (6.06
and 5.95, respectively) and Loagan Bunut (9.33 and 7.38, respectively) (Table 5). The average
number of alleles over all populations was 10.99. Figure 2 shows examples of some observed
alleles derived using GeneMapper ver 3.7 software. Polymorphic information content (PIC) was
the lowest at Kayangeran (0.621) and the highest at Sedilu (0.76) (Table 5).
6
Sixty seven of the 356 alleles detected occurred only in one of each of the population. Sedilu had
13 unique alleles that were not detected in other populations, Naman and Manggut had 9 unique
alleles, while Lingga and Betong each had 8 unique alleles. Other populations had less than 7
unique alleles. These unique alleles are very informative used for population identification.
Figure 2 Example of some fragment data showing variable alleles size from three
individuals of Gonystylus bancanus.
Observed and expected heterozygosities, Ho and He were generally high for all populations with
no value less than 0.6. When all populations were compared together, Kayangeran and Loagan
Bunut had the lowest value observed. On average, the expected heterozygosity was 0.766, which
was within the range observed for other tropical species (Table 6). However, the values of Fis
were significantly different from zero for three populations, an indication of inbreeding within
population (Table 5).
The value of measured genetic diversity at Kayangeran and Loagan Bunut are low, but this may
be a function of sampling error. Because the trees at both sites grow in a clump, most of the
samples were collected from trees less than 10 meters apart from one to another. Sedilu,
Manggut and Naman had high value of genetic diversity but these populations had significant
departure from Hardy Weinberg Expectation, with a deficiency of heterozygotes.
PIC
MNA
Ar
Ho(SD)
He(SD)
Fis
Sedilu
Lingga
Serapau
Loagan Bunut
Kayangeran
Maludam
Manggut
Betong
Naman
Mean
31
30
30
101
29
36
53
40
55
0.762
0.732
0.738
0.677
0.621
0746
0.746
0.748
0.761
0.725
11.50
11.44
11.11
9.33
6.06
11.22
12.61
12.33
13.28
10.99
10.96
10.8
10.66
7.38
5.95
10.34
10.33
10.67
10.96
9.78
0.688(0.019)
0.738(0.019)
0.747(0.019)
0.706(0.011)
0.655(0.021)
0.748(0.017)
0.731(0.014)
0.745(0.016)
0.752(0.014)
0.723(0.016)
0.803(0.036)
0.770(0.039)
0.775(0.036)
0.719(0.032)
0.682(0.036)
0.785(0.030)
0.782(0.034)
0.787(0.030)
0.792(0.030)
0.766(0.033)
0.145*
0.042
0.037
0.018
0.039
0.048
0.065*
0.053
0.052*
0.056
No of
samples
No of
Loci
40
173
156
30
90
32
8
7
7
4
7
2
No of
populations
1
1
1
1
5
1
He
0.64
0.70
0.64
0.78
0.71
0.75
Reference
Ujino et al., 1998
Ng et al., 2004.
Ng et al., 2004
Konuma et al., 2000
Lim et al., 2002
Konishi et al., 2004
Lingga
Serapau
Kayangeran
Maludam
Manggut
Betong
Naman
0.912005
Loagan
Bunut
0.761285
Sedilu
0.921376
0.716141
0.926301
0.903312
0.889029
0.861203
Lingga
0.081887
0.980047
0.766813
0.660457
0.980337
0.982971
0.956784
0.884990
Serapau
Loagan
Bunut
Kayangeran
0.092110
0.020154
0.750547
0.629940
0.968779
0.977598
0.951722
0.896850
0.272747
0.265513
0.286952
0.843882
0.774306
0.757500
0.801043
0.848819
Maludam
0.333878
0.414823
0.462131
0.169743
0.669202
0.659585
0.717615
0.726191
0.076556
0.019859
0.031719
0.255788
0.401670
0.976887
0.970750
0.899795
Manggut
0.101687
0.017175
0.022656
0.277732
0.416145
0.023384
0.968556
0.883434
Betong
0.117625
0.044177
0.049482
0.221840
0.331823
0.029686
0.031949
0.908063
Naman
0.149425
0.122179
0.108867
0.163909
0.319942
0.105589
0.123939
0.096441
Genetic identity and distances differed among populations (Table 7). The genetic identity (Nei
1978) ranged from 0.629940 between Kayangeran and Serapau to 0.982971 between Manggut
and Lingga. The results show that the populations are divided into at least two geographical
regions. Populations from the same geographical region such as Serapau, Maludam, Lingga,
Betong, Sedilu and Manggut have high values of more than 0.9. Similarly, when UPGMA
cluster analysis derived based on Neis gentic distance the populations were clustered into two,
namely the west cluster consisting of Naman, Sedilu, Betong, Maludam, Serapau, Lingga and
Manggut, and the east cluster consisting of Loagan Bunut and Kayangeran (Fig. 3). However it
seems that the west cluster was further separated and positively related to geographical location
when Lingga, Serapau, Manggut, Maludam and Betong populations showed short genetic
distance. The shortest genetic distance was Manggut and Lingga. The five populations were
connected through Lupar Rivers. Similar results were obtained when cluster analysis was
conducted using structure software ver. 2.3 computer program (Fig. 4).
To determine if populations were significantly distinguishable from each other, the pairwise
comparisons of populations was performed. Based on the Fst values, the populations were
significantly differentiated (P< 0.05) from each other except for Manggut, Serapau, Lingga and
Maludam. The Fst values ranged from 0.0042 between Lingga and Manggut to 0.0939 between
Sedilu and Kayangeran (Table 8). The low value of Fst observed between Lingga and Serapau,
Lingga and Maludam, and Lingga and Manggut indicated that they are similar to each other and
is more likely that gene flow would occur along the watercourse.
S e d ilu
15
14
B e to n g
13
M a lu d a m
12
S e ra p a u
17
11
L in g g a
10
M a n g g u t
L o a g a n B u n u t
16
K a y a n g e ra n
___________________________________________________
0.16
0.12
0.08
0.04
0.00
0 .1
Sedilu
Lingga
Serapau
Loagan
Bunut
Kayangeran
Maludam
Manggut
Betong
Sedilu
0.0000
Lingga
0.0200
0.0000
Serapau
Loagan
Bunut
0.0220
0.0052
0.0000
0.0753
0.0768
0.0811
0.0000
Kayangeran
0.0939
0.1155
0.1234
0.0619
0.0000
Maludam
0.0176
0.0050
0.0081
0.0729
0.1096
0.0000
Manggut
0.0240
0.0042
0.0056
0.0773
0.1113
0.0057
0.0000
Betong
0.0268
0.0116
0.0127
0.0649
0.0951
0.0072
0.0080
0.0000
Naman
0.0332
0.0307
0.0271
0.0502
0.0908
0.0256
0.0301
0.0234
10
Naman
0.0000
Figure 4 Cluster of population based on Structure ver. 2.3. Population divided into two
clusters: Red and Blue. Population 1: Sedilu; 2: Lingga; 3: Serapau; 4: Loagan
Bunut; 5: Kayangeran; 6: Maludam; 7: Manggut; 8: Betong and 9: Naman.
5.0 Conclusions
The effectiveness and efficiency of microsatellite markers to evaluate genetic diversity and
population differentiation demonstrated that microsatellite markers are useful for the
development of DNA database for ramin. The unique alleles detected in specific populations and
signification populations differentiation revealed by cluster analysis and Fst value indicated that
microsatellite markers could be used for population identification. Thus, microsatellite markers
are suitable to generate DNA database for individual timber identification and useful for tracking
of wood in production chain (Tnah et al., 2010; Koopman and Diemont 2004; Smulder et al.,
2008). However, the use of microsatellite markers for identification should not be based on a
limited number of markers but on many loci (Butler 2005). Furthermore, before a marker system
can be introduced into forensic casework, a population database must be established for
statistical evaluation of identity match proof (Tnah, 2010).
6.0 Recommendations
The current ramin DNA database was developed using leaves and bark samples. For actual
timber species identification and tracing and tracking system, samples from wood must be tested.
A new proposal entitled Use of DNA for identification of Gonystylus species and geographical
origin has been submitted to ITTO for funding. The unique alleles detected in specific
populations in Sarawak based on the 18 markers could be detected in other population outside
Sarawak. Thus, the use of more microsatellite markers and sampling across all ramin populations
to other states in Malaysia and neighboring countries is important until population specific
alleles detected. To implement this, support and commitment from respective countries, local
government, policy makers and various NGOs are needed. Once the database is completed, only
then it would be reliable in court proceeding to curb illegal logging enhance the implementation
of CITES legislation/regulations in this region.
11
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14
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149
163
149
149
153
153
149
157
149
155
149
155
149
163
149
149
149
149
155
163
149
28
153
173
151
149
163
170
163
155
155
149
163
161
171
155
170
155
171
149
155
149
161
149
169
170
155
155
149
169
175
155
142
131
146
148
146
118
142
138
146
136
138
138
134
0
126
118
142
140
126
146
134
142
130
130
118
146
146
138
142
154
142
142
156
157
155
146
173
138
146
140
140
150
146
0
142
138
146
150
140
165
142
156
146
146
173
173
146
163
150
154
186
189
189
186
186
186
189
189
189
186
186
189
189
186
186
189
189
189
186
189
189
186
186
189
189
186
186
186
186
189
186
189
189
186
189
186
189
189
189
186
186
189
189
186
186
189
189
189
189
189
189
186
189
189
189
189
186
186
189
189
116
124
116
114
122
116
116
114
116
114
116
126
116
116
116
116
130
114
116
116
116
130
114
122
116
108
126
108
122
114
130
128
124
122
126
130
122
130
130
130
128
126
130
122
122
126
130
122
116
122
122
130
122
122
130
122
137
116
130
137
120
112
112
106
112
116
106
118
124
116
116
112
118
120
116
116
118
108
108
112
106
112
118
120
120
106
120
116
118
116
120
122
120
112
124
120
120
120
134
120
122
130
124
134
124
134
134
120
114
116
116
118
120
130
120
116
124
118
118
120
MN044
MN045
MN046
MN047
MN048
MN049
MN050
MN051
MN052
MN053
MN054
MN055
BE001
BE002
BE003
BE004
BE005
BE006
BE007
BE008
BE009
BE010
BE011
BE012
BE013
BE014
BE015
BE016
BE017
BE018
236
250
248
258
230
262
250
256
250
248
246
262
250
248
248
228
248
248
248
248
244
256
250
248
250
248
222
254
226
248
248
268
258
280
268
264
250
264
272
248
266
264
264
278
264
248
248
250
250
264
246
258
290
290
250
270
266
276
246
258
88
88
96
90
90
90
90
88
98
88
90
90
88
90
90
90
90
90
88
90
90
98
88
88
90
90
96
88
90
90
90
96
98
96
90
90
90
96
102
108
90
106
98
104
90
90
96
104
90
104
98
104
98
90
98
90
98
90
96
90
145
145
145
125
125
145
145
145
145
145
145
145
145
145
145
125
125
153
145
155
143
145
145
125
125
145
145
125
125
125
153
145
145
145
145
145
145
167
145
145
145
147
145
155
155
153
125
153
145
155
155
163
155
145
145
145
145
125
167
125
174
174
182
174
204
178
182
182
174
166
186
186
184
186
186
190
194
159
159
186
186
184
182
188
181
170
181
188
174
174
194
182
201
201
204
186
182
194
182
201
195
186
186
190
197
196
196
186
194
188
206
186
190
190
184
184
188
204
188
180
149
149
155
151
155
149
149
149
155
153
153
149
155
153
149
155
153
149
149
149
163
149
149
153
153
149
149
149
149
149
29
155
163
162
163
163
149
149
153
171
153
167
155
167
153
170
171
163
155
155
169
163
163
165
169
172
155
163
171
167
167
142
120
136
136
146
142
142
142
118
144
138
118
130
138
150
134
146
138
148
138
138
136
126
126
138
142
150
126
136
146
146
146
146
150
150
146
142
146
146
173
140
138
138
140
150
168
150
146
157
144
146
144
150
126
150
146
165
138
142
146
189
189
186
186
186
189
189
189
189
186
186
186
186
189
186
186
189
189
189
186
186
186
189
189
186
186
189
189
186
186
189
189
189
186
186
189
189
189
189
189
186
189
189
189
189
189
189
189
189
186
186
186
189
189
189
186
189
189
189
186
114
114
122
124
114
122
126
122
114
128
124
116
126
122
116
122
126
114
116
116
116
112
126
116
132
116
114
126
116
116
114
130
122
130
122
122
130
130
126
128
126
126
130
126
130
130
130
126
116
126
126
130
139
139
132
126
132
134
130
126
114
116
112
106
106
116
118
118
106
118
120
106
116
114
114
106
106
106
116
112
120
116
106
106
116
106
118
112
124
124
120
124
122
120
122
116
120
120
116
120
122
134
124
116
120
116
116
118
116
116
122
126
116
116
116
126
124
124
124
124
BE019
BE020
BE021
BE022
BE023
BE024
BE025
BE026
BE027
BE028
BE029
BE030
BE031
BE032
BE033
BE034
BE035
BE036
BE037
BE038
BE039
BE040
NA001
NA002
NA003
NA004
NA005
NA006
NA007
NA008
248
226
250
250
248
262
248
248
248
248
222
248
248
244
252
248
250
248
226
238
238
248
262
270
246
250
248
248
246
248
272
274
252
258
286
268
252
264
264
250
268
252
258
252
276
276
268
254
250
286
238
250
268
274
262
260
250
256
246
256
90
90
88
90
90
90
88
90
90
90
96
98
90
96
90
90
88
90
90
90
90
90
90
90
90
88
90
90
90
104
114
90
90
98
90
90
88
98
90
98
98
98
98
98
90
100
90
98
98
98
98
96
90
104
104
90
90
104
90
104
125
145
163
145
125
145
145
145
143
145
145
145
145
125
145
125
145
145
145
125
125
125
149
125
145
145
125
145
125
145
145
145
163
155
145
145
151
161
145
155
145
145
155
145
145
145
145
145
147
145
145
155
149
145
145
145
157
157
145
155
186
186
186
181
170
180
174
174
189
188
174
174
174
180
197
188
166
170
174
170
174
193
190
190
180
194
190
188
180
159
194
192
200
191
186
201
181
180
196
192
186
182
174
186
201
192
182
186
194
174
187
198
192
204
188
194
194
190
186
197
149
159
149
163
153
155
153
145
149
153
155
149
158
155
149
155
155
149
155
149
153
155
149
153
149
149
149
149
149
155
30
149
163
169
163
175
155
155
155
167
157
155
162
175
163
149
167
163
155
163
153
170
155
170
171
157
149
155
169
163
175
116
136
140
142
136
116
142
140
138
146
150
146
140
118
150
146
142
146
142
146
138
124
0
142
122
142
140
142
142
138
138
146
142
146
146
161
150
144
150
150
161
156
144
140
156
150
146
160
146
146
140
126
0
160
146
160
156
156
142
150
189
186
186
186
186
189
189
189
186
189
186
186
189
186
186
189
189
186
189
186
186
186
186
186
186
186
186
186
186
189
189
189
186
186
189
189
189
189
189
189
186
189
189
189
189
189
189
186
189
186
186
189
189
186
190
189
186
186
186
189
122
122
114
116
114
116
116
114
116
116
116
114
114
122
122
130
116
114
114
114
130
122
126
114
126
126
130
122
126
106
130
122
114
130
124
124
118
124
122
122
122
122
134
126
130
130
126
122
130
130
130
139
130
116
126
130
130
130
128
116
116
106
116
108
118
116
106
116
106
118
114
114
114
116
106
116
112
106
104
116
118
112
106
106
116
106
122
122
106
116
116
112
124
112
118
122
112
118
112
118
116
116
114
118
116
140
118
118
112
116
118
124
130
106
120
122
122
134
116
124
NA009
NA010
NA011
NA012
NA013
NA014
NA015
NA016
NA017
NA018
NA019
NA020
NA021
NA022
NA023
NA024
NA025
NA026
NA027
NA028
NA029
NA030
NA031
NA032
NA033
NA034
NA035
NA036
NA037
NA038
266
216
236
236
276
248
252
262
268
256
252
246
246
258
262
250
224
250
236
228
246
250
248
248
248
236
246
254
268
250
272
216
254
236
276
254
270
262
270
272
272
250
272
274
276
274
256
264
264
228
248
250
276
248
256
252
248
272
272
256
90
88
90
90
90
90
90
90
90
90
88
88
98
98
90
90
90
90
90
88
88
90
90
88
90
90
90
104
90
90
90
90
104
90
100
90
90
106
98
96
90
104
98
104
104
90
90
104
90
88
92
90
90
104
90
90
90
104
102
102
125
125
125
145
125
125
145
145
147
145
143
145
0
145
125
145
145
145
125
125
125
125
125
125
145
145
145
145
145
125
155
125
145
145
155
125
161
161
161
147
143
159
0
155
125
147
151
155
125
145
145
149
145
145
153
149
153
145
155
145
192
192
192
190
195
194
184
180
187
178
186
182
188
178
187
180
180
186
186
184
178
190
190
180
180
188
187
182
184
188
206
200
200
204
201
204
195
182
195
186
194
190
188
204
194
187
190
188
194
184
188
194
192
184
190
188
190
188
196
188
153
160
160
155
149
149
155
149
149
153
149
149
157
149
170
149
155
149
157
149
157
155
149
149
155
163
163
155
155
149
31
155
164
164
157
157
157
157
157
170
170
175
149
157
149
175
158
158
157
175
155
165
175
157
167
167
169
163
163
169
175
136
126
138
146
122
142
142
142
142
144
138
142
142
142
118
144
142
134
160
136
142
144
142
140
140
122
138
142
140
144
142
130
150
150
142
150
154
144
154
146
146
144
142
150
144
146
144
142
163
160
155
156
144
142
142
134
148
144
156
154
189
189
189
189
189
189
189
189
189
186
186
186
0
186
189
189
186
186
186
186
189
186
189
186
186
189
186
186
186
186
189
189
189
189
189
189
189
189
189
189
189
189
0
186
189
189
186
189
189
189
189
186
189
189
189
189
189
189
189
189
116
0
0
126
116
126
126
114
126
126
114
124
128
106
126
126
126
114
122
114
114
126
114
114
126
122
126
126
106
116
130
0
0
130
130
130
126
126
130
126
126
126
128
116
126
128
126
122
122
126
122
126
116
122
130
130
126
130
126
126
116
125
114
122
116
120
114
114
114
122
124
114
0
118
118
118
114
124
116
114
106
112
118
116
116
116
114
122
126
108
122
125
140
140
140
140
116
114
124
134
140
122
0
122
118
134
128
142
120
116
106
120
120
134
118
120
116
126
142
116
NA039
NA040
NA041
NA042
NA043
NA044
NA045
NA046
NA047
NA048
NA049
NA050
NA051
NA052
NA053
NA054
NA055
246
262
248
272
226
246
256
246
246
246
238
246
238
248
228
256
230
274
268
248
276
274
262
264
276
266
250
266
260
248
276
274
272
280
92
90
90
90
90
90
90
88
90
90
88
88
90
90
90
88
88
92
104
90
90
106
90
102
90
104
104
90
98
90
90
104
88
90
125
125
145
125
145
153
145
125
145
149
125
125
125
147
125
145
125
155
145
145
145
153
164
145
145
145
159
145
143
145
151
145
145
164
186
188
194
192
182
170
176
190
188
159
180
182
170
194
188
184
184
190
196
198
200
187
178
190
192
192
190
186
188
184
194
192
204
204
153
161
149
149
153
155
0
149
149
153
155
157
170
155
151
149
153
32
175
163
157
170
167
171
0
163
155
163
173
170
170
163
170
153
175
142
140
136
134
144
136
124
142
136
144
136
142
144
142
132
142
142
142
142
156
136
144
142
150
142
142
156
142
150
148
144
144
144
158
189
186
189
189
186
189
186
186
189
189
189
189
186
186
189
189
186
189
189
189
189
189
189
186
189
189
189
189
189
189
189
189
189
189
126
114
114
120
122
126
116
114
126
114
120
114
114
122
126
122
116
130
126
126
130
126
126
130
126
126
128
122
122
122
122
130
126
126
110
106
106
114
122
106
108
114
120
114
118
118
118
122
120
108
118
114
136
124
126
122
106
116
120
120
114
120
118
122
122
134
112
122
Ind.
SE001
SE002
SE003
SE004
SE005
SE006
SE007
SE008
SE009
SE010
SE011
SE012
SE013
SE014
SE016
SE017
SE018
SE019
SE020
SE021
SE022
SE023
SE024
SE025
SE026
SE027
SE028
SE029
SE030
WGb06
196 199
196 199
199 199
187 199
196 196
196 196
187 196
199 199
195 195
187 199
183 198
195 198
199 199
187 199
187 196
199 199
187 196
192 196
190 199
196 199
199 199
199 199
177 199
196 196
199 199
199 199
187 196
196 199
187 199
WGb17
321 333
321 342
318 352
339 342
324 330
324 324
324 358
330 336
321 321
318 339
321 321
321 321
321 339
318 321
318 321
324 339
318 349
318 336
349 370
318 342
318 321
324 330
339 349
324 324
321 352
349 352
330 336
327 330
318 327
WGb22
282 284
276 288
272 284
270 284
272 272
0
0
242 284
270 294
272 272
284 296
0
0
0
0
284 284
284 284
270 284
272 286
242 286
284 284
272 284
282 282
270 282
270 296
270 270
272 272
270 284
272 286
284 292
272 280
270 286
WGb24
332 345
345 349
332 349
345 347
332 332
332 332
332 345
332 332
332 336
332 347
332 332
332 332
323 332
345 351
345 351
332 332
332 349
332 349
332 332
349 349
323 351
332 334
328 345
332 332
332 332
323 345
332 347
332 332
332 351
WGb29
135 146
135 135
118 122
111 137
102 102
104 104
135 135
135 141
102 102
122 135
102 104
104 104
122 122
128 143
135 135
135 141
104 135
122 135
135 135
104 135
143 143
122 135
126 135
104 104
111 135
111 135
135 135
128 135
111 128
33
WGb32
295 317
303 307
305 315
309 331
293 303
285 293
303 309
309 311
293 293
293 297
293 303
285 293
297 309
305 325
309 333
278 323
297 309
309 319
299 305
297 323
299 305
303 305
278 315
293 303
291 291
291 303
303 307
303 323
305 309
WGb37
169 173
169 171
162 169
171 171
169 175
169 169
173 173
171 173
169 169
162 171
169 169
169 169
169 171
171 173
173 173
162 169
162 173
173 173
169 173
162 169
162 171
169 173
162 169
169 169
162 162
162 173
173 173
169 173
169 173
WGb38
275 275
275 275
281 283
281 283
269 269
269 269
281 283
275 283
269 269
279 283
259 269
269 269
279 281
281 281
281 283
281 283
281 283
275 283
279 281
281 283
283 283
281 283
281 287
269 269
283 283
275 283
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164 164
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BE020
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BE024
BE025
BE026
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BE030
BE031
BE032
BE033
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BE035
BE036
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BE039
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NA010
NA011
NA012
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NA014
NA015
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NA020
NA021
NA022
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NA041
NA042
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NA044
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NA048
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NA050
NA051
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196
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160
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148
160
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160
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162
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47