Changes of carbon stocks in bamboo stands in China during 100 years

Forest Ecology and Management 258 (2009) 1489–1496

Changes of carbon stocks in bamboo stands in China during 100 years

Xiangang Chen a,*, Xiaoquan Zhang b, Yiping Zhang c, Trevor Booth d, Xinhua He e

a Department of the Environmental Science & Engineering, SWFC, Kunming 650224, PR Chinab Institute of Forest Ecology, Environment & Protection, CAF, Beijing 100091, PR Chinac Xishuangbanna Tropical Botanical Garden, CAS, Kunming 650223, PR Chinad CSIRO Forest Biosciences, PO Box E4008, Kingston, Canberra, ACT 2604, Australiae Department of the Foreign Languages, SWFC, Kunming 650224,PR China

A R T I C L E I N F O

Article history:

Received 14 February 2009

Received in revised form 27 June 2009

Accepted 30 June 2009

Keywords:

Bamboo stand

Biomass

Soil organic carbon

Carbon stock

A B S T R A C T

Bamboo stands are one of the most important forest types in China, covering an area of about 4.99

million hectares, and estimation of their carbon stocks is vital for China’s national carbon accounting.

Bamboo biomass and carbon fraction, as well as soil bulk density and soil organic matter content, data

were collated from 40 publications describing conditions at 35 sites in 10 Chinese provinces where most

bamboo stands are distributed. Carbon stocks and its changes in the living biomass and soil organic

matter in bamboo stands in China in the past five decades were estimated based on these collated data

together with the area of bamboo stands and number of bamboo culms derived from the National

Forestry Inventory (NFI). Our estimates indicate that the carbon stocks in bamboo stands in China have

been increasing since the 1950s with estimated values of 318.55 Tg C (1950–1962), 427.37 Tg C (1977–

1981), 463.80 Tg C (1984–1988), 493.00 Tg C (1989–1993), 548.79 Tg C (1994–1998) and 631.58 Tg C

(1999–2003) accompanying the increase of bamboo stand area. Based on correlation between forest area

and bamboo area, as well as the trends of forest area predicted in government strategy documents for

forest development over the next five decades, the carbon stocks in bamboo stands for 2010, 2020, 2030,

2040 and 2050 are estimated to be 727.08 Tg C, 839.16 Tg C, 914.43 Tg C, 966.803 Tg C and 1017.64 Tg C,

respectively.

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1. Introduction

Forests play a major role in global terrestrial carbon cycling. Onthe one hand, forests, through photosynthesis, sequester carbondioxide from the atmosphere and accumulate biomass (includingtrunks, branches, leaves and roots) as well as contributing to organiccarbon in soils. Though forests account for only 38% of terrestriallands on the Earth, they store 893 Pg C or 45.70% of the terrestrialcarbon stocks (Yoda, 1993). On the other hand, deforestation is animportant source of carbon dioxide emissions and second only to thefossil fuel combustion. The net emission of carbon dioxide due todeforestation from 1860 to 1980 has been estimated to be as high as135–220 Pg C (Houghton et al., 1983). Bamboo stands are one ofimportant forest types in the world and account for about 3% of thetotal forest area in China (SFAPRC, 2005). How the carbon stocks ofthese bamboo stands change should be a worthy issue for estimatingthe carbon stocks of China’s forests.

Bamboo includes about 1500 species under 87 genera in thesubfamily of Bambusiodeae worldwide (Ohrnberger, 1999). China,

* Corresponding author.

E-mail address: [email protected] (X. Chen).

0378-1127/$ – see front matter � 2009 Elsevier B.V. All rights reserved.

doi:10.1016/j.foreco.2009.06.051

with about 500 bamboo species belonging to 48 genera in thesubfamily of Bambusiodeae, is the largest producer of bamboo in theworld (Zhou, 1998). According to the statistic data obtained from the6th National Forestry Inventory (NFI) (1999–2003) (SFAPRC, 2005),there are 4.99 million hectares (Mha) of bamboo stands in China,among which 3.37 Mha is Phyllostachys pubescens accounting forabout 70% of the total. In the past three decades, the area of bamboostands in the mainland of China has increased by 51.40%, from3.20 Mha in the late 1970s to almost 5.00 Mha early this century(Fig. 1) due to afforestation on wasteland. Bamboo stand has becomean important forest type in China. Judging by the statistics of 6th NFIdata, about 98% of bamboo stands are distributed in southern China,including such 12 provinces as Fujian, Jiangxi, Zhejiang, Hunan,Guangdong, Sichuan, Guangxi, Anhui, Hubei, Chongqing, Guizhouand Yunnan. Bamboo stands in Fujian, Jiangxi and Zhejiangprovinces account for about half the total area (Fig. 2).

Living biomass and soil organic matter content in bamboo standshave been sampled and measured at stand level by many researchteams (Appendices A and B). There are also a few reports on thecarbon fraction or carbon stock of bamboo stands in some areas(Isagi et al., 1997; Lin et al., 1998b; Zhou and Jiang, 2004; Li et al.,2006). However, carbon stocks and its changes in bamboo stands atthe regional scale have not been individually assessed in previous

Fig. 1. Changes of bamboo stand area in China since 1950 (data are from the database of National Forest Inventory).

Fig. 2. Distribution of bamboo stands in China.

X. Chen et al. / Forest Ecology and Management 258 (2009) 1489–14961490

studies. So much bamboo stands have been excluded to someone’sstudies reporting on total carbon stocks in China’s forests (e.g., Fangand Chen, 2001). This paper estimates past carbon stock changes inbamboo stands in China as well as projecting likely future changes,thus filling a gap in carbon accounting in the forest sector.

2. Data and methods

The carbon stock in bamboo stand contains biomass carbon andsoil organic carbon (SOC). The biomass of bamboo is consisted ofliving biomass (stem, root, branch and leave), litter and deadbamboo. However when estimating the biomass of bamboo standmost researchers did not take account of litter and dead bamboo asthe two parts are generally very small proportion against livingbiomass (Appendix A). In this paper the carbon stock in bamboostand is expressed as the carbon stock in living biomass and soilorganic matter.

2.1. Carbon stocks in living biomass of bamboo stands

Bamboo, with height growth usually finished within 2–4months, is regarded as one of the fastest growing plants and hasquick renewing capacity (Zhou et al., 2005). When bamboo stand

become ripe its living biomass tends to a maximum and preservesa dynamic balance (Shang, 2002). A biomass loss due to harvestingcan be rapidly compensated for by subsequent regrowth ofbamboo. Therefore, the biomass of ripe bamboo stands can beassumed be steady-state.

Carbon stock in living biomass of bamboo stands was estimatedfor two bamboo categories, i.e. P. pubescens and other bamboospecies:

CB ¼ 0:5B1 þ 0:45B2 (1)

where CB is the total carbon stock (Tg C) in the living biomass ofbamboo stands; B1 and B2 represent the biomass of P. pubescens

and other bamboo species (Tg d.m.), respectively; 0.5 and 0.45 arecarbon fractions of P. pubescens (Zhou, 2004) and other bamboospecies (Lin et al., 1998b; Li et al., 2006), respectively.

The national forestry inventory provides two sets of data, i.e.,the area of bamboo stands and the number of standing bambooculms. Two methods are applied to calculate carbon stocks in livingbiomasses of bamboo stands:

� area-based method:

Bi ¼ BaiAi � 10�6 (2)

Table 1Estimating mean values of the biomass and soil bulk density and organic matter content of bamboo stands.

Bamboo species Statistic parameter Biomass (Mg ha�1) Rod biomass (kg culm�1) Bulk density

(g cm�3)

Organic matter con-

tent (%)

0–20 20–40 0–20 20–40

Phyllostachys pubescens Average 159.86 63.46 1.02 1.07 3.33 1.82

Standard deviation 137.50 51.59 0.15 0.12 0.89 0.64

Number of samples 21 14 20 15 20 15

Other bamboo species Average 95.36 2.51

Standard deviation 82.21 1.50

Number of samples 36 16

X. Chen et al. / Forest Ecology and Management 258 (2009) 1489–1496 1491

where Bi is the total biomass (Tg d.m.) of bamboo category i; Bai

represents the average biomass per hectare of bamboo category i

(Mg d.m. ha�1); Ai represents the area of bamboo category i (ha);

i = 1 (means P. pubescens), 2 (means other bamboo species).� culm-based method:

Bi ¼ BciNi � 10�9 (3)

where Bci represents the mean biomass of individual bamboo

category i (kg d.m. culm�1), Ni is the culm number of standing

bamboo category i (culm); i is same as in Eq. (2).

The biomass of bamboo stand per hectare varies with bamboospecies, geographical location, standing density, site conditionsand management practices. P. pubescens and Dendrocalamopsis

oldhami usually have relatively high biomass stock. Studies onbamboo biomass have been reviewed and the related biomassdata have been collated (Appendix A). Using these data meanbiomass stock per hectare or per bamboo culm was assessed forthe two bamboo groups, i.e., P. pubescens and other bamboospecies (Table 1). The values of Ai and Ni are derived from thedatabase of NFI.

Since 1950 the increases of bamboo stand area have beenclosely related to the increases of forest area across the wholecountry (Fig. 3). Two scenarios for the future changes of bamboostand area were considered:� Scenario A: The future area of bamboo stands was estimated by

extrapolating the trend of area changes of bamboo stands in thepast five decades (i.e. 1.79% annual increment) (Fig. 4):� Scenario B: The future area of bamboo stands was calculated

using relationship in Fig. 3 according to the projected forest areain the national strategy for forestry development (RGCFSDS,2003) (Fig. 4).

Fig. 3. Correlation of area between bamboo stands and total forests in China since

1950 (data are from the National Forest Inventory).

2.2. Carbon stocks in soil organic matter of bamboo stands

The carbon in soil organic matter means soil organic carbon. TheSOC stocks in bamboo stands are estimated based on the area ofbamboo stands and the SOC stocks per hectare:

SOCt ¼X

SOCiAi � 10�6 (4)

where SOCi represents the total SOC stocks in bamboo stands (Tg C),SOCi expresses the SOC stocks on unit area of bamboo category i

(Mg C ha�1), Ai presents the area of bamboo category i (ha).Bamboo is a shallow-rooted plant (EBFC, 1996) with its roots

generally concentrated in the upper 40 cm of soil. Most studiesreported soil organic matter content rather than SOC stock, so SOCstock was estimated via soil bulk density and organic mattercontent, using the following formula:

SOCi ¼ 0:58 �X

SDi jSOMi jDi j � 102 (5)

where 0.58 is Blemmelen ratio (adimensionless quantity relating soilorganic matter to organic carbon) (Jenny, 1988); SDij represents thesoil bulk density in layer j (g cm�3), SOMij is the soil organic mattercontent in soil layer j (%); Dij is the thickness of soil layer j (cm); j

represents surface soil layer (0–20 cm) and lower soil layer (20–40 cm), respectively. In generally the right of Eq. (5) needs to bemultiplied by a factor 1� P (P represents the proportion of gravel insoil) because the soil bulk density is estimated only for the fine soil.But in the existing studies on bamboo stand soil in China the effect ofgravel content in soil was almost removed when surveying andcalculating the value of soil bulk density. So Eq. (5) is suitable to suchstudies in China.

The values of SDij and SOMij are derived from the literatures(Appendix B). For cases without soil bulk density, SDij was calculatedusing the following formula (Post et al., 1982; Post and Mann, 1990):

SDi j ¼ a j þ b jZ j þ c jlog SOMi j (6)

where Zj is the depth from soil surface to the center of the layer j

(cm); aj, bj and cj are regressive coefficients, which are fitted as

Fig. 4. Predicted area of bamboo stands in China in the next five decades.

X. Chen et al. / Forest Ecology and Management 258 (2009) 1489–14961492

0.456, 0.056 and 0.012 for surface layer (0–20 cm) of soil and 0.484,0.02 and�0.113 for lower layer (20–40 cm) of soil according to theknown values of soil bulk density in P. pubescens stands inAppendix B, respectively. These parameters are also applied to theother bamboo species which have no relevant parameters.

3. Results

3.1. Carbon stock changes of bamboo stands in the past five decades

The carbon stocks in bamboo stands in the last six NFI periodscalculated using the area-based method, are 318.55 Tg C (1950–1962), 427.37 Tg C (1977–1981), 463.80 Tg C (1984–1988),493.00 Tg C (1989–1993), 548.79 Tg C (1994–1998) and631.58 Tg C (1999–2003), respectively (Fig. 5). Among them thecarbon stocks in living biomass are 166.71 Tg C, 229.38 Tg C,244.36 Tg C, 258.543 Tg C, 288.23 Tg C and 331.93 Tg C, respectively,accounting for around 46–48% of the total carbon stock. SOC stocksaccounts for around 52–54% over this period from 1950 to 2003.

The carbon stocks in bamboo stands in China, calculated usingthe culm-based method, are 286.60 Tg C (1950–1962), 341.81 Tg C(1977–1981), 414.54 Tg C (1984–1988), 436.28 Tg C (1989–1993),504.82 Tg C (1994–1998), 605.51 Tg C (1999–2003), respectively(Fig. 5). Among them the living biomass carbon stocks are134.75 Tg C, 143.82 Tg C, 195.10 Tg C, 201.71 Tg C, 244.26 Tg Cand 305.86 Tg C, respectively.

In the past five decades, the carbon stocks in bamboo stands,estimated using both methods, have been increasing (Fig. 5). Theiraverage annual increment estimated using the area-based methodand culm-based method are correspondingly 4.73 Tg C year�1 and2.40 Tg C year�1 from the first to second NFI period,5.20 Tg C year�1 and 10.39 Tg C year�1 from 2nd to 3rd NFIperiods, 5.84 Tg C year�1 and 4.35 Tg C year�1 from 3rd to 4thNFI periods, 6.08 Tg C year�1 and 13.71 Tg C year�1 from 4th to 5thNFI periods, and 16.56 Tg C year�1 and 20.14 Tg C year�1 from 5thto 6th NFI periods.

3.2. Projection of the carbon stock in bamboo stands

The carbon stocks in bamboo stands in China are estimated at741.61 Tg C, 885.45 Tg C, 1 057.18 Tg C, 1 262.23 Tg C and 1

Fig. 5. Changes of carbon stocks in bambo

507.04 Tg C, respectively by the years 2010, 2020, 2030, 2040and 2050 under Scenario A, and727.08 Tg C, 839.16 Tg C,914.43 Tg C, 966.803 Tg C and 1017.64 Tg C, respectively by theyears 2010, 2020, 2030, 2040 and 2050 under Scenario B (Fig. 5),showing a apparent increase in potential carbon sequestrationover the next five decades.

4. Discussion

4.1. Biomass in bamboo stand

The mean biomass of P. pubescens stands across the standsreviewed in Table 1 is 159.86 Mg d.m. ha�1, which is smaller thannatural broadleaf evergreen forests (169.4 Mg d.m. ha�1), but largerthan some broadleaf evergreen plantation (103.74 Mg d.m. ha�1)and typical subtropical needle-leaf forests (56.46–152.36Mg d.m. ha�1 for Cunninghamia lanceolata and Pinus massoniana)in the region, and very close to subtropical deciduous mixedbroadleaf forests (170.99 Mg d.m. ha�1) (Table 2)(Feng et al., 1999).Chen et al. (2000) reported that the biomass of P. pubescens stands ishigher than that of 10-year-old pure Chinese fir stand with similarsite conditions (namely 63.25 Mg d.m. ha�1). The average biomassof the other species of bamboo stands is 95.36 Mg d.m. ha�1

(Table 1), and is close to mean biomass of subtropical broadleafevergreen plantations, i.e., Chinese fir forests and P. massoniana

forests (Table 2). As bamboo stand is known to reach maximumbiomass within very short periods it is inferred that bamboo standcompares very favorably with many different types of forests in itsability to quickly store larger amounts of carbon.

4.2. Differences caused by estimation methods

The carbon stocks in bamboo stands estimated using the area-based method are 26.07–85.56 Tg C, larger than that estimatedusing the culm-based method (Fig. 5). One probable reason whythe notable differences appear is that the mean biomass of bamboostands is overestimated as the majority of sample sites used forstudding biomass density locate in bamboo stands with higherculm density. The difference between estimated values by twomethods tends to reduce gradually from 4th to 6th NFI periods,which can be explained by the fast increase of culm numbers due

o stands in China from 1950 to 2050.

Table 2Comparison of the biomass of several subtropical forest types.

Forest types Standard age (year) Number of samples Biomass (Mg ha�1)

Range Average

Subtropical broadleaf evergreen forests 12–42 8 80.40–323.35 169.4

>100 5 334.22–491.17 388.69

Subtropical broadleaf evergreen plantations 5–30 17 13.40–255.40 103.74

Subtropical mixed deciduous broadleaf forests Middle aged 2 147.38–194.60 170.99

Chinese fir forests 3–10 17 16.60–99.69 56.46

10–20 34 59.3–131.68 105

20–30 17 82.55–334.25 152.36

Pinus massoniana forests 6–14 7 29.92–112.06 84.8

20–30 5 33.40–155.17 106

X. Chen et al. / Forest Ecology and Management 258 (2009) 1489–1496 1493

to the increasing culm density of bamboo stands. For example,judging by the statistics of NFI data the average culm density of P.

pubescens stands increases by 81 culm ha�1 from 4th to 5th NFIperiods and 211 culm ha�1 from 5th to 6th NFI periods. However itshould be noted that the biomass per culm probably decreaseswith the increase of culm density.

4.3. Evaluation of projection scenarios

Carbon stocks predicted under Scenario A are larger than thatunder Scenario B due to the larger projected area of bamboo stands.This projected area of bamboo stands under Scenario A isestimated by extrapolating the past increment trend of bamboostands. So the great increase of bamboo stand area in China in last30 years results in a larger area of bamboo stands being projectedunder Scenario A in future. But, the bamboo stand area projectedunder Scenario A has little of possibility as the non-vegetationareas can be used for afforestation is limited. The project area ofbamboo stands under Scenario B is got based on the nationalforestry development strategy and the fitted relationships withtotal forest area, which conforms more to reality with standarderror only 0.1 of the predicted value. The increment rate of forestarea tends to decrease due to the limitation of available lands forforestry development. As a result, the expansion rate of bamboostand tends to decrease.

4.4. Contribution of bamboo stands in carbon stock in forest of China

In the past five decades, the area of bamboo stands hasaccounted for 2.9% (1950–1962), 2.8% (1977–1981), 2.8% (1984–1988), 2.7% (1989–1993), 2.6% (1994–1998) and 3.0% (1999–2003)of all forest area in China according to the data from NFI (Fig. 3). Theproportion of bamboo stands area to all forest area decreasesslowly from 1950s to 1990s and increases rapidly after the end of1990s. Fang and Chen (2001) reported that the carbon stocks inliving biomass of China’s forests (excluding bamboo stands) are4380 Tg C (1977–1981), 4450 Tg C (1984–1988), 4630 Tg C (1989–1993) and 4750 Tg C (1994–1998). The carbon stocks in living

biomass of bamboo stands estimated above accounts for 7.80–9.76%, 9.32–10.42%, 9.42–10.65% and 10.63–11.55% of that ofChina’s forests, respectively for the 2nd, 3rd, 4th and 5th NFIperiods. The share of the carbon stocks of bamboo stands in totalcarbon stocks of forest rises continually and is remarkable in thepast. This indicates bamboo stands is one of the importantcomponents of carbon stock in China’s forests.

5. Conclusion

The statistics of NFI data show that the area of bamboo standsincreased along with forested lands and accounted for about 3% oftotal forest area in China in last five decades. P. pubescens is mainbamboo species and its area accounts for about 70% of that of allbamboo stands. Bamboo stands are an important forest type. Thestatistics of the sample data, collected from 21 sites for P.

pubescens and 36 sites for other species of bamboo, indicate thatthe average biomass of P. pubescens is 159.86 Mg d.m. ha�1 or63.46 kg d.m. culm�1 and that of other bamboo species is95.36 Mg d.m. ha�1 or 2.35 kg d.m. culm�1. Calculation showsthat the carbon stocks in bamboo stands increased obviously inlast five decades, and it contributes around 10% of carbon stocks inliving biomass of forests in China. It is expected that the area ofbamboo stands will increase significantly along with theincreasing forest area and the carbon stocks in bamboo standswill rise apparently in China in next five decades. The estimatedvalues of carbon stocks in bamboo stands under Scenario B arepreferable results because this scenario conforms more to reality.The bamboo stands in China will play an important role of carbonsinks in forest systems in coming five decades.

Acknowledgements

We would like to give our appreciation to the Natural ScienceFoundation of Yunnan Province for its financial support, and toprofessor Wuyuan Yun for his correcting the Latin of bamboospecies in this paper.

Appendix A. (*Mean value is used; **unpublished data)

Bamboo species Location Culm density

(culm ha�1)

Biomass

(Mg ha�1)

Culm biomass

(kg culm�1)

Reference

Phyllostachys pubescens Guizhou 1620 264.72 163.41 Wu (1983)

2055 260.39 126.71

2385 218.73 91.71

3825 271.42 70.96

3990 301.64 75.6

Qinshimen in Zhejiang 572.29 Wen (1990)

Anji in Zhejiang 162.91

Fuyang in Zhejiang 2700 120.12 44.49 Huang et al. (1993)

3750 182.38 48.63

Nanjing in Fujian 80.11 Li et al. (1993)

Dagangshan in Jiangxi 2788 60.98 21.87 Nie (1994)

3900 86.29 22.12

4545 99.54 21.9

Jian-ou in Fujian 81.74 Lan et al. (1999)

2280 341.08 149.6 Zheng et al. (1997)

Shaxian in Fujian 1350–1650* 25.29 16.86 **

2100–2400* 41.3 18.36

3900–3200* 57.64 16.24

Fujian 37.6 Zheng et al. (1998a)

Wuping in Fujian 67.18 Chen et al. (2000)

Jigong mountain in Henan 23.7 Yan et al. (2004)

Phyllostachys heteroclada Shucheng in Anhui 30000 93.67 3.12 Sun et al. (1986)

37500 156.26 4.17

45000 102.02 2.27

52500 121.08 2.31

Dendrocalamopsis oldhami Ruian in Zhejiang 148.19 Wen (1990)

Hua-an in Fujian 156.04 Lin et al. (1998a)

Dendrocalamopsis oldhami Wenzhou in Zhejiang 100.94 Wen (1990)

Dendrocalamus latiflorus Cangnan in Zhejiang 81.61

Phyllostachys nidularis Anji in Zhejiang 93.97

Phyllostachys viridis 114.87

Phyllosachys atro-aginata 296.58

Lophatherum gracile Xiaoshan in Zhejiang 202.44

Bambusa albo-lineata Shaoan in Fujian 41.12

98.11

Phyllostachys iridenscens Hengxian in Zhejiang 370.37

Pleioblastus amarus Yuhang in Zhejiang 280.58

102.82 Lin et al. (2004)

Western Sichuan 100.96 Li et al. (2006)

Neosinocalamus affinis Jinyunshan mountain in Chongqing 70180 156.41 2.23 Su and Zhong (1991)

Pleioblastus maculatus 24.06 Liu and Zhong (1996)

Oligostachyum oedogonatum Jian-ou in Fujian 15000 10.58 0.71 Zheng and Chen (1998)

30000 40.62 1.35

60000 70.72 1.18

90000 38.76 0.43

120000 30.06 0.25

Fujian 85004 34.63 0.41 Zheng et al. (1998b)

Qiongzhuea tumidinoda Daguan inYunnan 48.17 Dong et al. (2002)

Phyllostachys meyeri Songxi in Fujian 11584 52.73 4.55 Xu et al. (2004)

Gelidocalamus stellatus Jingang mountain in Jiangxi 16.03 Zhou and Jiang (2004)

Phyllostachys bambusoides Jigong mountain in Henan 87 Yan et al. (2004)

Arundianaria fargesii Zhenba in Shanxi 69.09 Wang et al. (2005)

Pseudosasa amabilis Mingqing in Fujian 6750 30.48 4.52 Lin (2005)

9000 32.51 3.61

11250 39.72 3.53

13500 40.63 3.01

Chimonobusa quadrangularis Nianping in Fujian 14.96 Tong (2007)

29.60

X. Chen et al. / Forest Ecology and Management 258 (2009) 1489–14961494

Appendix B

Some parameters relating to the soil attribute of Phyllostachys pubescens stand (*The italic figure means the calculation result using formula

(6); **the value in this column comes from 0.58�SD�SOM in formula (5); ***unpublished data).

Location Sampling site condition Bulk density

(g cmS3)

Organic matter

content (%)

SOC density

(10S2 g cmS3)

Reference

0–20 20–40 0–20 20–40 0–20** 20–40**

Anji in Zhejiang No understory 1.02* 3.89 2.31 Jiang and Hong (1987)

Herbal understory 1.02* 5.07 3.01

Mixed bamboo and arbor 1.02* 4.04 2.40

Fuyang in Zhejiang No understory 1.02* 2.52 1.49

Herbal understory 1.02* 2.99 1.77

Shaxian in Zhejiang No understory 1.02* 3.69 2.19

Herbal + bushes 1.02* 4.27 2.54

Fuyang in Zhejiang Bamboo fungus in stripes 0.94 3.77 2.06 Chen and Pei (1991)

Bamboo fungus in patches 0.70 3.86 1.57

Comparison sites 1.14 2.38 1.57

Fujian 0.90 1.02 2.25 1.67 1.17 0.99 Zheng et al. (1998a)

Fuzhou in Fujian Deep plough and fertilizing 1.18 1.21 4.28 2.50 2.93 1.75 Chen (1999)

Deep plough 0.98 1.00 3.30 1.17 1.87 0.68

Complete clearing of herbal 1.07 1.19 2.64 1.21 1.64 0.84

Grass removed 1.10 1.18 2.55 1.13 1.62 0.77

Comparison sites 1.01 1.02 1.42 0.97 0.83 0.57

Changting in Zhejiang 1.02* 1.07* 2.58 1.43 1.53 0.89 Zheng et al. (2000)

Anji in Zhejiang High production 1.02* 1.06* 3.48 1.80 2.06 1.10 Xu et al. (2000)

Low production 1.02* 1.06* 3.56 1.60 2.11 0.98

Zhuji Mts in Zhejiang 1.19 1.09 4.16 3.23 2.88 2.04 Huang (2001)

Zhe Jiang NE slope 0.94 2.20 1.20 ***

SW slope 1.14 2.00 1.32

Lower part of slope 0.81 2.50 1.17

Higher part of slope 1.14 2.00 1.32

X. Chen et al. / Forest Ecology and Management 258 (2009) 1489–1496 1495

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