**Use of the Pilodyn for Assessing Wood Properties in Standing Trees of** *Eucalyptus* **Clones<sup>1</sup>**

Wu Shijun1, Xu Jianmin1\*, Li Guangyou1, Risto Vuokko2, Lu Zhaohua1, Li Baoqi1 and Wang Wei1 *1Research Institute of Tropical Forestry, Chinese Academy of Forestry, Guangdong Guangzhou 2Guangxi StoraEnso Forestry Corporation Ltd., Nanning, Guangxi China* 

#### **1. Introduction**

196 New Advances and Contributions to Forestry Research

Torelli, N. (1994). Relationship between Tree Growth Characteristics, Wood Structure and

Jahrgang, Heft 6, 112–116.

Utilization of Beech (*Fagus sylvatica* L.). *Holzforschung und Holzverwertung*, 45.

As one of the major timer production species in the world, eucalyptus is characterized as fast-growing, high-yielding, and well adapted to different flat and mountainous environments with extreme low temperature of -5℃ ((Qi 2007)). Most of eucalypts species are naturally distributed in the continental Australia of Oceania, and a few native to the Timor Island of Indonesia and Papua New Guinea (Qi 2007).Identification and selection of superior trees in forest management and breeding programs provide a means to improve the properties and value of future wood products (Knowles et al. 2004). In recent years, breeding objectives in tree improvement have moved from volume per hectare alone, to include also wood properties and their impact on industrial end products (Wei and Borralho 1997). Wood basic density is considered as one of the most important wood properties which has a major impact on the freight costs, chipping properties, pulp yield per unit mass of wood and paper quality (Pliura et al. 2007; Laurence et al 1999). Wood basic density generally shows a high heritability and responds well to genetic improvement. But the genetics of wood density has not been studied in great detail (Macdonald et al. 1997). Currently, published information on genetic variation in wood basic density in eucalyptus is limited with a few studies conducted in China (Kien et al. 2008; Lu 2000; Luo 2003).

Measurement of wood density is expensive and time consuming and also create varying degrees of damage to experimental materials, and that has restricted the number and accuracy of the studies published (Hansen 2000; Wei et al. 1997). However, pilodyn sampling is faster, cheaper, and not destructive, thus resulting in overall higher expected gains for selection of

<sup>1</sup> This study belongs to the project of National Eleventh Five-Year Science and Technology" Breeding of High yield and High Quality Fast-growing Wood Species of Eucalyptus" (2006BAD01A15-4)

Author: Wu Shijun (1984-- ) male, Shandong Weifang, Under Post-graduate Student, wushijun0128@163.com

<sup>\*</sup>Corresponding Author: Xu Jianmin Professor

Use of the Pilodyn for Assessing Wood Properties in Standing Trees of *Eucalyptus* Clones 199

The pilodyn wood tester is an instrument originally developed in Switzerland for determining the degree of soft rot in wooded telephone poles (Raymond et al. 1998; Hansen 2000). Pilodyn penetration (PP), an indirect method for determining wood basic density, has been effective in assessing large number of trees in eucalyptus (Wei et al. 1997; Kien et al. 2008; Macdonald et al. 1997; Raymond et al. 1998) and other species (Ishiguri et al. 2008; Pliura et al. 2006). PP was measured using a 6-J Forest pilodyn with 2.5mm steel needle, by over the bark and removing a small section of bark (approximately 40mm × 20mm) at 1.3m respectively and taking two pilodyn shots on each of four aspects (north, south west and east) from an average tree per plot. The pilodyn is attractive in that it is rapid, does not require the use of an increment borer (destructive sampling), and is, in principle, free of operator bias (Cown 1978; Hansen 2000). To avoid introducing additional sources of error, all clones were sampled by the same team of people, minimizing the potential for operator

FAKOPP microsecond timer is able to measure acoustic velocity in standing trees, by timing the acoustic wave as it travels along the stem between points a known distance apart (Knowles et al. 2004; Chauhan et al. 2006). The results signals were engendered by start and stop transducers and recorded on an oscilloscope. Stress wave velocity (SWV) was then calculated by dividing the test span by the measurement stress wave transmission time

Where L=1500 mm is the distance between two probes, t is the transmission time in

The SWV is combined with density measurements to give an estimated of dynamic MoE

Where MoE is the dynamic modulus of elasticity, ρ is the average green density of the stem,

Wood basic density was defined as oven-dry wood mass per unit volume of green wood, and was measured using the water displacement method (Kube and Raymond 2002; Tappi 1989). Five mm increment cores from pith to bark were extracted at a height of 1.3 m in the south-north orientation from an average tree per plot, immediately stored in plastic tubes with both ends sealed (Kien et al. 2008).Wood basic density was determined using the water displacement method, with two weights for every sample: weight of water displaced by immersion of wedge (w1) and oven dry weight (w2) (Kien et al. 2008). Basic DEN was then

SWV = L / t (1)

MoE = ρω2 (2)

**2.2 Assessments of wood properties** 

**2.2.1 Pilodyn penetration** 

error (Raymond et al. 1998).

**2.2.2 Modulus of elasticity** 

(Wang et. al. 2000).

microseconds (μs).

ω is the SWV.

calculated as:

(Knowles et. al., 2004).

**2.2.3 Wood basic density** 

trees or culling of seedling seed orchards in comparison with the more destructive direct assessment of density (Greaves et al. 1996). Kube and Raymond (2002) reported that core sampling for basic density is assumed to cost \$10.5 per tree, which includes field collection and laboratory processing, whereas the cost of pilodyn measurements is assumed to be \$1.5 per tree.

The primary objective of this study is to test the effectiveness of pilodyn for evaluating wood basic density, modulus of elasticity (MoE) and other traits of eucalyptus clones in standing trees. This information will be used to develop appropriate selection strategies for eucalyptus breeding programs in southern China.

#### **2. Materials and methods**

#### **2.1 Trial description**

The trial was established at Shankou town in Guangxi (21°34' N, 112°42 E, 29m asl.), and is affected by the north tropical monsoon with annual mean temperature of 23℃ and annual mean rainfall of 1589mm. The lateritic red earth was derived from sandstone and contains 0.15% of organic matter (0-20cm). Previous vegetation was a plantation of Eucalyptus. Indigenous vegetation was found on site. 22 eucalyptus clones (table 1) were planted in April 2004. Field design was randomized complete blocks with 7 replications and 5-tree plot in a spacing of 4m × 2m. Measurements and increment cores were collected in December 2008, at which time the trial was aged 56 months.


Note: Male parents of U6, W5 and SH1 were not clear.

Table 1. Details of clones in the analysis

#### **2.2 Assessments of wood properties**

#### **2.2.1 Pilodyn penetration**

198 New Advances and Contributions to Forestry Research

trees or culling of seedling seed orchards in comparison with the more destructive direct assessment of density (Greaves et al. 1996). Kube and Raymond (2002) reported that core sampling for basic density is assumed to cost \$10.5 per tree, which includes field collection and laboratory processing, whereas the cost of pilodyn measurements is assumed to be \$1.5 per tree. The primary objective of this study is to test the effectiveness of pilodyn for evaluating wood basic density, modulus of elasticity (MoE) and other traits of eucalyptus clones in standing trees. This information will be used to develop appropriate selection strategies for

The trial was established at Shankou town in Guangxi (21°34' N, 112°42 E, 29m asl.), and is affected by the north tropical monsoon with annual mean temperature of 23℃ and annual mean rainfall of 1589mm. The lateritic red earth was derived from sandstone and contains 0.15% of organic matter (0-20cm). Previous vegetation was a plantation of Eucalyptus. Indigenous vegetation was found on site. 22 eucalyptus clones (table 1) were planted in April 2004. Field design was randomized complete blocks with 7 replications and 5-tree plot in a spacing of 4m × 2m. Measurements and increment cores were collected in December

Clone number Clone Identity. Parental Combination Style of Seedling 1 GRDH32-26 *E.urophylla* ×*E.grandis* Cuttings 2 W5 ABL 12×Unknown Tissue culture 3 GRDH32-29 *E.urophylla* ×*E.grandis* Cuttings 4 M1 *E.urophylla* ×*E.grandis* Tissue culture 5 GRDH32-28 *E.urophylla*×*E.grandis* Cuttings 6 SH1 Leizhou NO.1×Unknown Tissue culture 7 GRDH33-9 *E.urophylla* ×*E.grandis* Cuttings 8 U6 *E.urophylla* x *E.tereticornis* Tissue culture 9 GRDH32-25 *E.urophylla*×*E.grandis* Cuttings 10 DH32-29 *E.urophylla*×*E.grandis* Tissue culture 11 GRDH42-6 *E.grandis* ×*E.urophylla* Cuttings 12 RGD3 *E.urophylla*×*E.camaldulensis* Tissue culture 13 DH196 *E.urophylla*×*E.grandis* Cuttings 14 DH32-28 *E.urophylla*×*E.grandis* Tissue culture 15 GRDH30-10 *E.urophylla*×*E.grandis* Cuttings 16 TH9224 *E.urophylla*×*E.camaldulensis* Tissue culture 17 GRDH33-27 *E.urophylla*×*E.grandis* Cuttings 18 LH1 *E.urophylla*×*E.tereticornis* Tissue culture 19 TH9224 *E.grandis* x *E.camaldulensis* Cuttings 20 DH32-22 *E.urophylla*×*E.grandis* Tissue culture 21 DH32-13 *E.urophylla*×*E.grandis* Tissue culture 22 DH32-25 *E.urophylla*×*E.grandis* Tissue culture

eucalyptus breeding programs in southern China.

2008, at which time the trial was aged 56 months.

Note: Male parents of U6, W5 and SH1 were not clear.

Table 1. Details of clones in the analysis

**2. Materials and methods** 

**2.1 Trial description** 

The pilodyn wood tester is an instrument originally developed in Switzerland for determining the degree of soft rot in wooded telephone poles (Raymond et al. 1998; Hansen 2000). Pilodyn penetration (PP), an indirect method for determining wood basic density, has been effective in assessing large number of trees in eucalyptus (Wei et al. 1997; Kien et al. 2008; Macdonald et al. 1997; Raymond et al. 1998) and other species (Ishiguri et al. 2008; Pliura et al. 2006). PP was measured using a 6-J Forest pilodyn with 2.5mm steel needle, by over the bark and removing a small section of bark (approximately 40mm × 20mm) at 1.3m respectively and taking two pilodyn shots on each of four aspects (north, south west and east) from an average tree per plot. The pilodyn is attractive in that it is rapid, does not require the use of an increment borer (destructive sampling), and is, in principle, free of operator bias (Cown 1978; Hansen 2000). To avoid introducing additional sources of error, all clones were sampled by the same team of people, minimizing the potential for operator error (Raymond et al. 1998).

#### **2.2.2 Modulus of elasticity**

FAKOPP microsecond timer is able to measure acoustic velocity in standing trees, by timing the acoustic wave as it travels along the stem between points a known distance apart (Knowles et al. 2004; Chauhan et al. 2006). The results signals were engendered by start and stop transducers and recorded on an oscilloscope. Stress wave velocity (SWV) was then calculated by dividing the test span by the measurement stress wave transmission time (Wang et. al. 2000).

$$\mathbf{SWV} = \mathbf{L} \;/\; \mathbf{t} \tag{1}$$

Where L=1500 mm is the distance between two probes, t is the transmission time in microseconds (μs).

The SWV is combined with density measurements to give an estimated of dynamic MoE (Knowles et. al., 2004).

$$\text{MoE} \equiv \text{p} \text{o}^2 \tag{2}$$

Where MoE is the dynamic modulus of elasticity, ρ is the average green density of the stem, ω is the SWV.

#### **2.2.3 Wood basic density**

Wood basic density was defined as oven-dry wood mass per unit volume of green wood, and was measured using the water displacement method (Kube and Raymond 2002; Tappi 1989). Five mm increment cores from pith to bark were extracted at a height of 1.3 m in the south-north orientation from an average tree per plot, immediately stored in plastic tubes with both ends sealed (Kien et al. 2008).Wood basic density was determined using the water displacement method, with two weights for every sample: weight of water displaced by immersion of wedge (w1) and oven dry weight (w2) (Kien et al. 2008). Basic DEN was then calculated as:

Use of the Pilodyn for Assessing Wood Properties in Standing Trees of *Eucalyptus* Clones 201

pilody penetration and pith taken from clones. The results indicated that PP was generally reliable as an indirect measure of wood basic density. The correlations between pilodyn and MoE were significantly and negative. However, the relationship between pilodyn and MoE does not seem to be documented. And further research is needed to clarify in further. The correlations between pilodyn and heartwood density were slightly positive to strongly positively, lower than the correlations between pilodyn and other wood properties because

Clone number Mean value of PP (mm) wood basic density (g.cm-3) MoE (GPa) 16 9.44 0.4395 6.48 4 10.03 0.4913 7.42 12 10.28 0.4638 7.53 2 10.66 0.4145 4.93 15 10.81 0.4384 6.14 19 11.03 0.4302 6.31 8 11.15 0.362 3.94 20 11.41 0.4236 5.76 6 11.44 0.4295 5.25 18 11.47 0.4371 5.85 21 11.47 0.4262 5.84 10 11.63 0.4627 6.33 1 12 0.4614 5.88 3 12.22 0.4237 5.52 14 12.69 0.3938 5.48 9 12.97 0.4106 5.32 22 13.09 0.4172 5.65 17 13.5 0.4266 5.92 7 14.03 0.4164 5.47 11 14.94 0.3924 4.45 13 15.34 0.3899 4.35 5 15.41 0.3514 4.29

of the short length of steel needle.

Variance analysis of pilodyn

Table 2. The mean value of Pilodyn penetration and wood properties

The correlations between pilodyn and wood properties

Table 4. Variance analysis of pilodyn

Treatment PP over the bark PP with bark removal

Table 3. The mean value and variation coefficient of pilodyn penetration on four directions

Source DF F Value Pr≥F Treatment 1 16.47 < 0.0001 Directions 6 21.13 < 0.0001 Clones 21 8.10 < 0.0001

Index East West South North Mean East Wes South North Mean Mean PP (mm) 14.50 14.99 14.59 14.62 14.67 12.04 12.33 12.15 12.02 12.14 C V (%) 10.42 11.83 9.15 10.94 10.41 14.40 14.45 13.40 14.06 13.57

$$\text{BasicDEN} \left(\text{g} \right. \left. \text{cm} \text{-} \text{3} \right) = \text{w}\_2 \text{ / } \text{w}\_1 \tag{3}$$

Wood basic density, outer wood basic density and heartwood basic density were tested respectively.

#### **2.3 Statistic analysis**

The SAS software package was used to analyze the variance of different Pilodyn penetration and the relationship between the Pilodyn penetration and wood density or MOE, respectively.

The mean by ramet at each clone of sampling was submitted to a variance and a covariance analysis according to the following linear model (Hansen et. al. 1997):

$$\mathbf{y}\_{\overline{i}\overline{\eta}} = \mathbf{\mu} + \mathbf{c}\mathbf{\overline{i}} + \|\beta\_{\overline{i}} + \mathbf{c}\_{\overline{\eta}}\tag{4}$$

where yij is the performance of ramet of ith clone within jth block, μ is the general mean, αi is the random effect of the ith clone, βj is the random effect of the jth block, εij is the random error.

#### **3. Results and discussion**

#### **3.1 Comparison between Pilodyn penetration and wood properties**

The mean values of Pilodyn penetration and wood properties of 22 clones are listed in Table 2. The mean value ranged from 9.44 to 15.41 mm for Pilodyn penetration, 0.3514 to 0.4913 g.cm-3 for wood basic density and 3.94 to 7.53 GPa for MoE, which were smaller than previous studies on the same species (Knowles et al. 2004; Kien et al. 2008; Wei et al. 1997) as well as other species (Jacques 2004; Zhu et al. 2008; Zhu et al. 2009). The most suitable range of basic density for pulpwood in eucalyptus is 0.48 to 0.57 g • cm-3 and pulp yield decrease sharply when basic density falls below 0.4 exceeds 0.60 (Dean 1995; Ikemori et. al. 1986). There were considerably lower density values than those found in this study. Consequently, wood basic density should be improved substantially to about 0.55 g • cm-3, and this would benefit pulp production in southern China (Kien et al. 2008). Clones of M1, RGD3 and TH9224 had higher basic density and MoE, meanwhile, clones of DH32-28, GRDH42-6 and DH196 had lower basic density and MoE.

The variation coefficient of Pilodyn penetration over the bark was ranged from 9.15% to 11.83%, whereas those measured by removing the bark was ranged from 13.40% to 14.45% (Table 3). One possible explanation could be that bark thickness and branch cluster frequency could affect this value. This agreed well with previously published results by Wei (1997) and Yin (2008).

The analysis of variance of pilodyn is presented in Table 4. There were significant (1% level) differences between pilodyn penetration of different treatment, different directions and different clones, indicating that selection of clones for pilodyn would be effective.

The regression equations and phenotypic correlations between pilodyn penetration and wood properties are given in Table 5 and Table 6, respectively. Generally strongly negative correlations were found between pilodyn and wood properties, ranging from -0.433 to - 0.755, slightly lower than previously published study (Wei et al. 1997; Chapola 1994). The possible explanation could be at least in part to the relatively small age of materials or less pilody penetration and pith taken from clones. The results indicated that PP was generally reliable as an indirect measure of wood basic density. The correlations between pilodyn and MoE were significantly and negative. However, the relationship between pilodyn and MoE does not seem to be documented. And further research is needed to clarify in further. The correlations between pilodyn and heartwood density were slightly positive to strongly positively, lower than the correlations between pilodyn and other wood properties because of the short length of steel needle.


Variance analysis of pilodyn

200 New Advances and Contributions to Forestry Research

Wood basic density, outer wood basic density and heartwood basic density were tested

The SAS software package was used to analyze the variance of different Pilodyn penetration and the relationship between the Pilodyn penetration and wood density or MOE,

The mean by ramet at each clone of sampling was submitted to a variance and a covariance

 yij = μ +αi + βi + εij (4) where yij is the performance of ramet of ith clone within jth block, μ is the general mean, αi is the random effect of the ith clone, βj is the random effect of the jth block, εij is the random error.

The mean values of Pilodyn penetration and wood properties of 22 clones are listed in Table 2. The mean value ranged from 9.44 to 15.41 mm for Pilodyn penetration, 0.3514 to 0.4913 g.cm-3 for wood basic density and 3.94 to 7.53 GPa for MoE, which were smaller than previous studies on the same species (Knowles et al. 2004; Kien et al. 2008; Wei et al. 1997) as well as other species (Jacques 2004; Zhu et al. 2008; Zhu et al. 2009). The most suitable range of basic density for pulpwood in eucalyptus is 0.48 to 0.57 g • cm-3 and pulp yield decrease sharply when basic density falls below 0.4 exceeds 0.60 (Dean 1995; Ikemori et. al. 1986). There were considerably lower density values than those found in this study. Consequently, wood basic density should be improved substantially to about 0.55 g • cm-3, and this would benefit pulp production in southern China (Kien et al. 2008). Clones of M1, RGD3 and TH9224 had higher basic density and MoE, meanwhile, clones of DH32-28, GRDH42-6 and

The variation coefficient of Pilodyn penetration over the bark was ranged from 9.15% to 11.83%, whereas those measured by removing the bark was ranged from 13.40% to 14.45% (Table 3). One possible explanation could be that bark thickness and branch cluster frequency could affect this value. This agreed well with previously published results by Wei

The analysis of variance of pilodyn is presented in Table 4. There were significant (1% level) differences between pilodyn penetration of different treatment, different directions and

The regression equations and phenotypic correlations between pilodyn penetration and wood properties are given in Table 5 and Table 6, respectively. Generally strongly negative correlations were found between pilodyn and wood properties, ranging from -0.433 to - 0.755, slightly lower than previously published study (Wei et al. 1997; Chapola 1994). The possible explanation could be at least in part to the relatively small age of materials or less

different clones, indicating that selection of clones for pilodyn would be effective.

analysis according to the following linear model (Hansen et. al. 1997):

**3.1 Comparison between Pilodyn penetration and wood properties** 

respectively.

respectively.

**2.3 Statistic analysis** 

**3. Results and discussion** 

DH196 had lower basic density and MoE.

(1997) and Yin (2008).

Basic DEN (g • cm-3) = w2 / w1 (3)

Table 2. The mean value of Pilodyn penetration and wood properties


Table 3. The mean value and variation coefficient of pilodyn penetration on four directions


The correlations between pilodyn and wood properties

Table 4. Variance analysis of pilodyn

Use of the Pilodyn for Assessing Wood Properties in Standing Trees of *Eucalyptus* Clones 203

1. The mean value of Pilodyn penetration, wood basic density and MoE ranged from 9.44

2. There were significant differences between pilodyn penetration of different treatment, different directions and different clones. The coefficient of variation ranged from 9.15% to 11.83% for Pilodyn penetration over the bark and ranged from 13.40% to 14.45% for

3. The correlations between pilodyn and wood properties were generally strongly negative, and the coefficients ranged from -0.433 to -0.755. The results indicated that wood basic density and MoE can be predicted by using pilodyn. Results from this study also tend to confirm those of Cown (1981) who concluded that Pilodyn is not an accurate equipment for measurement, but it does provide an effective and efficient

We thank Huang Hongjian, Tan Peitao and Hu Yang from Xinhui Forest Bureau for their assistance. Comments from K. Harding, D. Pegg, Dr. Zeng Jie and an anonymous reviewer

Chapola Gbj. 1994. Assessment of some wood properties of eucalyptus species grown in Malawi using pilodyn method. Discovery and Innovation. 6(1):98-109 Chauhan S.S., Walker J.C.F. 2006. Variation in acoustic and density with age, and their interrelationships in radiation pine. Forest Ecology and Management. 229:388-394 Cown, D.J.. 1981. Use of the pilodyn wood tester for estimating wood density in standing

Cown D J . 1978. Comparison of the Pilodyn and torsiometer methods for the rapid assessment

Dean, G.H. 1995. Objectives for wood fibre quality and uniformity. In: pott, B.M., Borralho,

Ikemori Y. K., Martins F. C. G. and Zobel, B. J.. 1986 The impact of accelerated breeding on

Ishiguri, F., Matsui, R., Lizuka, K., Yokota, S. and Yoshizawa, N. 2008. Prediction of the

of 36-year-old Japanese larch trees. Oiginal Arbelten · Originals. 66: 275–280 Greaves, B.L., Borralho, N.M.G, Raymond, C.A. and Farrington, A. 1996. Use of a pilodyn

Hansen C. P. 2000. Application of the Pilodyn in forest tree improvement. DFSC Series of Technical Notes. TN55. Danida Forest Seed Centre, Humlebaek, Denmark. Hansen J.K., Roulund H. 1996. Genetic parameters for spiral grain, stem form, pilodyn and

N.M.G., Reid, J. B., Cromer, R. N., Tibbits, W.N. and Raymond, C.A. (eds). Eucalypts plantations: improving fibre yield and quality. CR-IUFRO Conf., Hobart, 19-24 Feb. 483

31 wood Properties. In proceedings of the 18th IUFRO World 32 Conference

mechanical properties of lumber by stress-wave velocity and Pilodyn penetration

for the indirect selection of basic density in *Eucalyptus nitens.*Canadian Journal of

growth in 13 years old clones of *Sitka Spruce* (Picea sitchensis (Bong.) Carr.). Silva

trees – in fluce of site and tree age. World Forestry Conference.

Division 5: Forest products. Ljubljana, Yugoslavia. p.358-368

Forest Research. 26(9):1643-1650

Genetica. 46 (2-3):107-113

of wood density in living trees. N ZJ For Sci ,:384 – 3911

to 15.41 mm, 0.3514 to 0.4913 g.cm-3 and 3.94 to 7.53 GPa, respectively.

Pilodyn penetration with bark removal.

means of estimating wood properties.

**5. Acknowledgements** 

are appreciated.

**6. References** 

pp.




Table 6. Regression analysis of wood properties (y) to pilodyn penetration(x) with bark removal on four directions

#### **4. Conclusion**

In the present study, the effectiveness of pilodyn for assessing wood properties of eucalyptus clones in standing trees was discussed. The results obtained are as follows:


#### **5. Acknowledgements**

We thank Huang Hongjian, Tan Peitao and Hu Yang from Xinhui Forest Bureau for their assistance. Comments from K. Harding, D. Pegg, Dr. Zeng Jie and an anonymous reviewer are appreciated.

#### **6. References**

202 New Advances and Contributions to Forestry Research

Directions Wood properties Regression equation R2 R

Table 5. Regression analysis of wood properties (y) to pilodyn penetration (x) over the bark

Directions Wood properties regression equation R2 R

Table 6. Regression analysis of wood properties (y) to pilodyn penetration(x) with bark

In the present study, the effectiveness of pilodyn for assessing wood properties of eucalyptus clones in standing trees was discussed. The results obtained are as follows:

MoE y = 0.0341x2 - 1.3743x + 18.331 0.365 -0.604\*\* Basic density y = 0.0003x2 - 0.0189x + 0.6422 0.281 -0.530\* Outer wood density y = 0.0007x2 - 0.0356x + 0.8051 0.417 -0.646\*\* Heartwood density y = -0.0003x2 + 0.0008x + 0.4794 0.188 -0.433\*

MoE y = -0.0037x2 - 0.1913x + 9.3565 0.374 -0.611\*\* Basic density y = -2E-05x2 - 0.0103x + 0.581 0.363 -0.603\*\* Outer wood density y = 0.0009x2 - 0.0409x + 0.8485 0.482 -0.695\*\* Heartwood density y = -0.0005x2 + 0.0076x + 0.4308 0.274 -0.523\*

MoE y = 0.05x2 - 1.9367x + 23.169 0.424 -0.651\*\* Basic density y = 0.0019x2 - 0.0735x + 1.0769 0.395 -0.629\*\* Outer wood density y = 0.0031x2 - 0.1139x + 1.4198 0.521 -0.722\*\* Heartwood density y = 0.0011x2 - 0.0459x + 0.8543 0.289 -0.538\*\*

MoE y = -0.0105x2 - 0.0191x + 8.1845 0.357 -0.597\*\* Basic density y = -0.0003x2 - 0.0042x + 0.5399 0.357 -0.598\*\* Outer wood density y = 0.0014x2 - 0.0579x + 0.9693 0.464 -0.681\*\* Heartwood density y = -0.0012x2 + 0.0255x + 0.3069 0.284 -0.533\*

MoE y = 0.0187x2 - 0.9296x + 15.216 0.389 -0.624\*\* Basic density y = 0.0006x2 - 0.0313x + 0.7468 0.359 -0.599\*\* Outer wood density y = 0.0022x2 - 0.0823x + 1.1665 0.493 -0.702\*\* Heartwood density y = -0.0003x2 - 7E-05x + 0.4975 0.262 -0.511\*

MoE y = 0.0429x2 - 1.3872x + 15.992 0.431 -0.656\*\* Basic density y = 9E-05x2 - 0.0133x + 0.5688 0.357 -0.598\*\* Outer wood density y = 0.0006x2 - 0.0304x + 0.7061 0.529 -0.727\*\* Heartwood density y = -0.0005x2 + 0.0031x + 0.4548 0.235 -0.484\*

MoE y = 0.0082x2 - 0.5428x + 11.063 0.431 -0.656\*\* Basic density y = 1E-05x2 - 0.0117x + 0.5661 0.414 -0.644\*\* Outer wood density y = 0.0008x2 - 0.0367x + 0.7538 0.530 -0.728\*\* Heartwood density y = -0.0005x2 + 0.0033x + 0.4582 0.313 -0.560\*\*

MoE y = -0.0063x2 - 0.171x + 8.6605 0.365 -0.604\*\* Basic density y = -0.0016x2 + 0.0291x + 0.3081 0.356 -0.596\*\* Outer wood density y = 0.0002x2 - 0.0204x + 0.6545 0.528 -0.727\*\* Heartwood density y = -0.0027x2 + 0.0589x + 0.1055 0.263 -0.513\*

MoE y = 0.0054x2 - 0.4438x + 10.18 0.344 -0.587\*\* Basic density y = -0.0007x2 + 0.0061x + 0.453 0.375 -0.613\*\* Outer wood density y = 0.0012x2 - 0.045x + 0.7988 0.541 -0.736\*\* Heartwood density y = -0.0018x2 + 0.0353x + 0.2603 0.282 -0.531\*

MoE y = 0.0079x2 - 0.5492x + 11.117 0.418 -0.646\*\* Basic density y = -0.0008x2 + 0.0084x + 0.4432 0.404 -0.635\*\* Outer wood density y = 0.0006x2 - 0.0326x + 0.7318 0.569 -0.755\*\* Heartwood density y = -0.0018x2 + 0.0341x + 0.2696 0.295 -0.543\*\*

East

West

South

North

Mean value

on four directions

East

West

South

North

Mean value

**4. Conclusion** 

removal on four directions


**13** 

*1,2P. R. China* 

*3Japan* 

**An Overview on Spruce Forests in China** 

*1Shanghai Institute of Urban Ecology and Sustainability* 

 *School of Life Science, East China Normal University, Shanghai 2Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang* 

Zou Chunjing1, Xu Wenduo2, Hideyuki Shimizu3 and Wang Kaiyun1

*3National Institute for Environmental Studies, 16-2 Onogawa, Tsukuba, Ibaraki* 

The genus *Picea* A. Dietrich (spruce), which is a relative isolated group under evolution, belongs to Pinaceae family (Ran et al., 2006; Bobrow, 1970; Buchholz, 1929, 1931; Alvin, 1980; Mikkola, 1969). It includes 28–56 species depending on different systems of classification (Farjón, 1990; Ledig et al., 2004), most of which are in Eastern Asia, while many researchers thought that there were about 40 species in *Picea* genus and were only found in the north hemisphere (Budantsey, 1992, 1994; Wolfe, 1975, 1978; Tiffney and Manchester, 2001). The distribution range is from 21oN (Huanglian Mountains of Vietnam) to 70oN (Far Eastern area of Russia) (Fig. 1). Spruce forests are the main dominant vegetation in alpine coniferous forest in subtropical zone and temperate zone, and they are only found in alpine area, subalpine area and plateau from 21oN to 46oN (Li, 1995). In cold temperate zone and its adjacent regions (47oN to 57oN), spruce forests are the zonal vegetation types in boreal coniferous forest. From 57oN to 70oN, spruce forests transform from horizontal (latitudinal) zonal distribution to vertical (altitudinal) zonal distribution and from continuous

In the north of Euro-Asia continent, the main spruce species are *Picea abies* (L.) H. Karst. and *P. obovata* Ledeb., which form the continuous boreal coniferous forest (Ferguson, 1967; Florin, 1954, 1963; Guerli et al., 2001). *P. abies* is found in Alps of France, the Balkan Peninsula or the Balkan Mountains in the west, Germany and Scandinavian Peninsula in the north, Poland and the north and middle region of Russia in the east. In Siberian area of Russia, *P. obovata* takes the place of *P. abies*, it is found until to Lena River Valley and Okhotsk. But in the east Siberian area, *P. obovata* retreats from the dominant position, and is taken the place by *Larix sibirica* Ledeb. due to the rigorous continental climate (Colleau,

In North America, spruce species are abundant, including *P. glauca* (Moench) Voss*, P. mariana* (Mill.) Britton and al.*, P. engelmannnii* Parry ex Engelm. and *P. sitchensis* (Bong.) Carriére (Barbour and Bilings, 1988; Klaus, 1987). *P. glauca* is distributed extensively in Canada and North USA, from Labradorian Peninsula and Alaska to Montana, North Dakota, Minnesota, Wisconsin, Michigan, to Massachusetts near Atlantic coast. *P. mariana* is

**1. Introduction** 

distribution to discontinuous distribution.

1968; Corrigan et al., 1978; Harris, 1979; Hart, 1987).


### **An Overview on Spruce Forests in China**

Zou Chunjing1, Xu Wenduo2, Hideyuki Shimizu3 and Wang Kaiyun1

*1Shanghai Institute of Urban Ecology and Sustainability School of Life Science, East China Normal University, Shanghai 2Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 3National Institute for Environmental Studies, 16-2 Onogawa, Tsukuba, Ibaraki 1,2P. R. China 3Japan* 

#### **1. Introduction**

204 New Advances and Contributions to Forestry Research

Jacques D., Marchal M. and Curnel y. 2004. Relative efficiency of alternative methods to

Knowles R. Leith, Hansen Lars W., Wedding Adele, Downes Geoffrey. 2004. Evaluation of

Kube P., Raymond C. 2002. Selection strategies for genetic improvement of basic density in

Knowles, L. R., Hansen, L.W., Wendding, A. and Downes, G.. 2004. Evaluation of non-

Laurence, R Schimleck, Anthony J Michell, Carolyn A Raymond, Allie Muneri. 1999.

Lu, Z., Xu, J., Bai J. and Zhou,W.. 2000. A study on wood property variation between Eucalyptus *tereticornis* and Eucalyptus *camalduensis*. Forest Research. 13(4):370-376 Luo, J. 2003 Variation in growth and wood density of *Eucalyptus urophylla*. In turnbull, J.

Macdonald A.C., Borralho N.M.G.and Potts B.M. 1997. Genetic variation for growth and

Kien N. D., Jansson G., Harwood C., Almqvist C., Thinh H. H. 2008. Genetic variation in

Pliura A., Zhang S. Y., Mackay J. and Bousquet. 2007. Genotipic variation in wood density

Qi S. 2007. Applied Eucalypt cultivation in China. Beijing: China Forestry Publishing House Raymond, C.A. and MacDonald, A.C. 1998. Where to shoot your pilodyn: within tree

Wang X., Ross R. J., Mcclellan M., Barbour R. J., Erickson J. R., Forsman J.W. and Mcgin G.

Yin Y., Wang L., Jiang X.. 2008. Use of Pilodyn tester for estimating basic density in standing trees of hardwood plantation. Journal of Beijing Forestry University 30 ( 4 ): 7-11 Zhu J., Wang J., Zhang S., Zhang J., Sun X., Liang B.. 2008. Wood property estimation and

Zhu J., Wang J., Zhang S., Zhang J., Sun X., Liang B., Zhao K. 2009. Using the pilodyn to assess wood traits of standing trees *Laix kaempfri*. Forest Research. 22(1): 75-79

Tappi.1989. Basic density and moisture content of pulpwood. TAPPI no. T258 om-98

Agriculture, Forest Service, Forest Products Laboratory, Madison, WI. Wei X., N.M.G.Borralho. 1997. Genetic control of basic density and bark thickness and their

selection of *populus tomentosa.* Scientia Silvae Science. 44(7): 23-28

selection. Ann. For. Sci. 61. pp:35-43

Journal of Forestry Science. 34(1):87-101

*Eucalyptus nitens.* Tcchnical repoet 92.

Journal of Forestry Science. 34(1)::87-101

Genetica. 46 (4):236-241

Management. 2007 238: 92-106

Genetica. 46(4):245-250

Tasmania. New Forests. 15: 205–221.

Canadian Journal of Forest Research. Feb. 29:194-202

New Zealand Journal of Forestry Science. 38 (1):160-175

evaluate wood stiffness in the frame of *hybrid larch* (Larix × *eurolepis* Henry) clonal

non-destructive methods for assessing stiffness of Douglas fir trees. New Zealand

destructive methods for assessing stiffness of douglas fir trees. New Zealand

Estimation of basic density of *Eucalyotus globulus* using near-infrared spectroscopy.

W.(Ed.) *"Eucalypts in Asia" ACIAR proceedingsNo. 111.* Zhanjiang, Guangdong, China.

wood density in *eucalyptus globulus ssp.globulus* in Tasmania (Australia). Silva

wood basic density and pilodyn penetration and their relationships with growth, stem straightness and branch size for *eucalyptus urophylla* S.T.Blake in Northern Vietnam.

and growth traits of poplar hybrids at four clonal trials. Forest Ecology and

variation in basic density in plantation *eucalyptus globulus* and *E. nitens* in

36 D. 2000. Strength and stiffness assessment of standing trees using a 37 nondestructive stress wave technique. Res. Pap. FPL-RP-585*.* U.S. Department of 38

relationships with growth traits of *Eucalyptus urophylla* in south east China. Silvae

The genus *Picea* A. Dietrich (spruce), which is a relative isolated group under evolution, belongs to Pinaceae family (Ran et al., 2006; Bobrow, 1970; Buchholz, 1929, 1931; Alvin, 1980; Mikkola, 1969). It includes 28–56 species depending on different systems of classification (Farjón, 1990; Ledig et al., 2004), most of which are in Eastern Asia, while many researchers thought that there were about 40 species in *Picea* genus and were only found in the north hemisphere (Budantsey, 1992, 1994; Wolfe, 1975, 1978; Tiffney and Manchester, 2001). The distribution range is from 21oN (Huanglian Mountains of Vietnam) to 70oN (Far Eastern area of Russia) (Fig. 1). Spruce forests are the main dominant vegetation in alpine coniferous forest in subtropical zone and temperate zone, and they are only found in alpine area, subalpine area and plateau from 21oN to 46oN (Li, 1995). In cold temperate zone and its adjacent regions (47oN to 57oN), spruce forests are the zonal vegetation types in boreal coniferous forest. From 57oN to 70oN, spruce forests transform from horizontal (latitudinal) zonal distribution to vertical (altitudinal) zonal distribution and from continuous distribution to discontinuous distribution.

In the north of Euro-Asia continent, the main spruce species are *Picea abies* (L.) H. Karst. and *P. obovata* Ledeb., which form the continuous boreal coniferous forest (Ferguson, 1967; Florin, 1954, 1963; Guerli et al., 2001). *P. abies* is found in Alps of France, the Balkan Peninsula or the Balkan Mountains in the west, Germany and Scandinavian Peninsula in the north, Poland and the north and middle region of Russia in the east. In Siberian area of Russia, *P. obovata* takes the place of *P. abies*, it is found until to Lena River Valley and Okhotsk. But in the east Siberian area, *P. obovata* retreats from the dominant position, and is taken the place by *Larix sibirica* Ledeb. due to the rigorous continental climate (Colleau, 1968; Corrigan et al., 1978; Harris, 1979; Hart, 1987).

In North America, spruce species are abundant, including *P. glauca* (Moench) Voss*, P. mariana* (Mill.) Britton and al.*, P. engelmannnii* Parry ex Engelm. and *P. sitchensis* (Bong.) Carriére (Barbour and Bilings, 1988; Klaus, 1987). *P. glauca* is distributed extensively in Canada and North USA, from Labradorian Peninsula and Alaska to Montana, North Dakota, Minnesota, Wisconsin, Michigan, to Massachusetts near Atlantic coast. *P. mariana* is

An Overview on Spruce Forests in China 207

Mountains of Shanxi Province in North China. In Northwest China, Arertai Mountains are the south border of *P. obovata*. *P. schrenkiana* var*. tianshanica* (Rupr.) Cheng et S. H. Fu is found in Tianshan Mountains, the west border of spruce forests in China. *P. crassifolia* Kom. Is distributed extensively in Qilian Mountains, Helan Mountains and Yinshan Mountains of Qinghai Province, Gansu Province and Ningxia Hui Nation Autonomous Region (Editorial

In Southwest China, including West Sichuan Province, north Yunnan Province and south Tibet Autonomous Region, there are 17 spruce species, which takes 43.3% of spruce species in the world. The important species are *P. likiangensis* (Franch.) E. Pritz., *P. likiangensis* var. *linzhiiensis* Cheng et L. K. Fu, *P. likiangensis* var. *balfouriana* (Rehd. et Wils.) Hillier ex Slavin, *P. purpurea* Mast., *P. brachytyla* (Franch.) E. Pritz., and so on, they form subalpine dark

*P. spinulosa* (Griff.) A. Henry and *P. smithiana* (Wall.) Boiss. are found in moist area in Himalayas in south Tibet Autonomous Region, and they always form small pure forest or

*P. morrisonicola* Hayata forms the dominant pure coniferous forest in Central Mountains in Taiwan, which is the only subalpine coniferous forest of the east China in subtropical zone

Monophyly of *Picea* has never been debated (Wright, 1955; Prager et al., 1976; Frankis, 1988; Price, 1989; Sigurgeirsson and Szmidt, 1993), but infrageneric classification of the genus remains quite controversial (Liu, 1982; Schmidt, 1989; Farjón, 1990, 2001; Fu et al., 1999), owing to morphological convergence and parallelism (Wright, 1955), and high interspecific crossability (Ogilvie and von RudloV, 1968; Manley, 1972; Gorden, 1976; Fowler, 1983, 1987; Perron et al., 2000). In addition, little is known about phylogenetic relationships of most species, especially the geographically restricted species growing in the montane regions of southwest China (LePage, 2001). Moreover, the origin and biogeography of *Picea* have drawn great interest from both geologists and biologists (Wright, 1955; Aldén, 1987; Page

Spruce species are fine trees for lumbering, so researches on spruce were conducted very early in China (Editorial Committee of Vegetation of China, 1980). However, basic characteristics, flora, distribution types, and evolution relationship of the spruce species in China, and the relationship among spuce in China and abroad need more concern. There are

The aim of this study was (1) to summarize systematically the researches on spruce in China, and (2) to try to clarify the relationship among Chinese spruces, and among spruce in

*Picea* spp. is distributed extensively in China (Editorial Committee of Forest of China, 1997). It is difficult to expatiate on the characteristics of edificators in spruce forests, so we divided China into five parts according to their districts, including Northeast China, Northwest

and Hollands, 1987; LePage, 2001, 2003), but they are still far from being resolved.

**2. Characteristics of species composition in spruce forests in China** 

many data about the topics above, but they are always scattered.

**2.1 The edificators in spruce forests in China** 

coniferous forests in Southwest China (Kuan, 1981; Sun, 2002; Wu et al., 1995).

Committee of Forest of China, 1997).

mixed forest (Kuan, 1981).

(Liu, 1971).

China and abroad.

distributed almost in the whole Canada, and extensively in the eastern provinces and Newfoundland, to Alaska across Rocky Mountains in the west, and to Pennsylvania, north Virginia, Wisconsin, and Michigan in the south. Britain Colombia Province of Canada is the west border of *P. mariana*. *P. engelmannnii* is found in the west of North America, from Alberta Province and Britain Colombia Province of Canada to Arizona and New Mexico of USA along Rocky Mountains, it is also distributed in Cascade Range in Washington and Oregon. *P. sitchensis* distributes in the northwest of North America, and can be found from Aleutian Islands to Pacific coast of the northwest of California too (Delevoryas and Hope, 1973; Hsu, 1983; Weng and Jackson, 2000).

Fig. 1. The modern distribution range and fossil localities of *Picea* spp. in the world (based on Li, 1995, Lű et al., 2004, and McKenna, 1975) (1-7. Fossil localities: 1. Eocene; 2, 3. Oligocene; 4, 5. Miocene; 6, 7. Pliocene; 8. Modern distribution)

In China, the distribution range of spruce forests is very large, from Daxinganling Mountains (north) to Gaoligong Mountains (south), and Tianshan Mountains (west) to Central Mountains of Taiwan Province (east) (Fang, 1995, 1996; Fang and Liu, 1998). The spruce forests are found as long as there are site conditions of cold-temperate moisture types. In China the spruce forests belong to vertical zonal distribution with 17 species and 8 variations of *Picea* genus and take more than 40% of the species in the world. Furthermore, the almost all of the species are endemic in China, except for those in Daxinganling Mountains which belongs to East Siberian area and Arertai Mountains (belonging to West Siberian area). In China, spruce forests are distributed in Northeast, North, Northwest and Southwest.

In the mountains of Daxinganling, Xiaoxinganling and Changbai of Northeast China, *P. koraiensis* Nakai, *P. jezoensis* var. *microsperma* (Lindl. Cheng et L. K. Fu) and *P. jezoensis* var. *komarovii* (V. Vassil.) Cheng et L. K. Fu are the edificators of upland dark coniferous forests, which are extended partition of dark coniferous forests of Far East Area of Russia (Editorial Committee of Forest of China, 1997; Li, 1980; Li and Zhou, 1979).

The distribution range of spruce forest is restricted for drought in North and Northwest China. *P. meyeri* Rehder, E. H. Wilson and *P. wilsonii* Mast. are found in Jibei Mountains, Xiaowutai Mountains of Hebei Province, Guanqin Mountains, Wutai Mountains, Guandi

distributed almost in the whole Canada, and extensively in the eastern provinces and Newfoundland, to Alaska across Rocky Mountains in the west, and to Pennsylvania, north Virginia, Wisconsin, and Michigan in the south. Britain Colombia Province of Canada is the west border of *P. mariana*. *P. engelmannnii* is found in the west of North America, from Alberta Province and Britain Colombia Province of Canada to Arizona and New Mexico of USA along Rocky Mountains, it is also distributed in Cascade Range in Washington and Oregon. *P. sitchensis* distributes in the northwest of North America, and can be found from Aleutian Islands to Pacific coast of the northwest of California too (Delevoryas and Hope,

Fig. 1. The modern distribution range and fossil localities of *Picea* spp. in the world (based on Li, 1995, Lű et al., 2004, and McKenna, 1975) (1-7. Fossil localities: 1. Eocene; 2, 3.

In China, the distribution range of spruce forests is very large, from Daxinganling Mountains (north) to Gaoligong Mountains (south), and Tianshan Mountains (west) to Central Mountains of Taiwan Province (east) (Fang, 1995, 1996; Fang and Liu, 1998). The spruce forests are found as long as there are site conditions of cold-temperate moisture types. In China the spruce forests belong to vertical zonal distribution with 17 species and 8 variations of *Picea* genus and take more than 40% of the species in the world. Furthermore, the almost all of the species are endemic in China, except for those in Daxinganling Mountains which belongs to East Siberian area and Arertai Mountains (belonging to West Siberian area). In China, spruce forests are distributed in Northeast, North, Northwest and

In the mountains of Daxinganling, Xiaoxinganling and Changbai of Northeast China, *P. koraiensis* Nakai, *P. jezoensis* var. *microsperma* (Lindl. Cheng et L. K. Fu) and *P. jezoensis* var. *komarovii* (V. Vassil.) Cheng et L. K. Fu are the edificators of upland dark coniferous forests, which are extended partition of dark coniferous forests of Far East Area of Russia (Editorial

The distribution range of spruce forest is restricted for drought in North and Northwest China. *P. meyeri* Rehder, E. H. Wilson and *P. wilsonii* Mast. are found in Jibei Mountains, Xiaowutai Mountains of Hebei Province, Guanqin Mountains, Wutai Mountains, Guandi

Oligocene; 4, 5. Miocene; 6, 7. Pliocene; 8. Modern distribution)

Committee of Forest of China, 1997; Li, 1980; Li and Zhou, 1979).

1973; Hsu, 1983; Weng and Jackson, 2000).

Southwest.

Mountains of Shanxi Province in North China. In Northwest China, Arertai Mountains are the south border of *P. obovata*. *P. schrenkiana* var*. tianshanica* (Rupr.) Cheng et S. H. Fu is found in Tianshan Mountains, the west border of spruce forests in China. *P. crassifolia* Kom. Is distributed extensively in Qilian Mountains, Helan Mountains and Yinshan Mountains of Qinghai Province, Gansu Province and Ningxia Hui Nation Autonomous Region (Editorial Committee of Forest of China, 1997).

In Southwest China, including West Sichuan Province, north Yunnan Province and south Tibet Autonomous Region, there are 17 spruce species, which takes 43.3% of spruce species in the world. The important species are *P. likiangensis* (Franch.) E. Pritz., *P. likiangensis* var. *linzhiiensis* Cheng et L. K. Fu, *P. likiangensis* var. *balfouriana* (Rehd. et Wils.) Hillier ex Slavin, *P. purpurea* Mast., *P. brachytyla* (Franch.) E. Pritz., and so on, they form subalpine dark coniferous forests in Southwest China (Kuan, 1981; Sun, 2002; Wu et al., 1995).

*P. spinulosa* (Griff.) A. Henry and *P. smithiana* (Wall.) Boiss. are found in moist area in Himalayas in south Tibet Autonomous Region, and they always form small pure forest or mixed forest (Kuan, 1981).

*P. morrisonicola* Hayata forms the dominant pure coniferous forest in Central Mountains in Taiwan, which is the only subalpine coniferous forest of the east China in subtropical zone (Liu, 1971).

Monophyly of *Picea* has never been debated (Wright, 1955; Prager et al., 1976; Frankis, 1988; Price, 1989; Sigurgeirsson and Szmidt, 1993), but infrageneric classification of the genus remains quite controversial (Liu, 1982; Schmidt, 1989; Farjón, 1990, 2001; Fu et al., 1999), owing to morphological convergence and parallelism (Wright, 1955), and high interspecific crossability (Ogilvie and von RudloV, 1968; Manley, 1972; Gorden, 1976; Fowler, 1983, 1987; Perron et al., 2000). In addition, little is known about phylogenetic relationships of most species, especially the geographically restricted species growing in the montane regions of southwest China (LePage, 2001). Moreover, the origin and biogeography of *Picea* have drawn great interest from both geologists and biologists (Wright, 1955; Aldén, 1987; Page and Hollands, 1987; LePage, 2001, 2003), but they are still far from being resolved.

Spruce species are fine trees for lumbering, so researches on spruce were conducted very early in China (Editorial Committee of Vegetation of China, 1980). However, basic characteristics, flora, distribution types, and evolution relationship of the spruce species in China, and the relationship among spuce in China and abroad need more concern. There are many data about the topics above, but they are always scattered.

The aim of this study was (1) to summarize systematically the researches on spruce in China, and (2) to try to clarify the relationship among Chinese spruces, and among spruce in China and abroad.

#### **2. Characteristics of species composition in spruce forests in China**

#### **2.1 The edificators in spruce forests in China**

*Picea* spp. is distributed extensively in China (Editorial Committee of Forest of China, 1997). It is difficult to expatiate on the characteristics of edificators in spruce forests, so we divided China into five parts according to their districts, including Northeast China, Northwest

An Overview on Spruce Forests in China 209

Taiwan *P. morrisonicola* 100.00 14.87 77.77 5.451 84.72

Table 1. Species characters in spruce forest in different districts in China (F-Frequency (%), D-Density (/100m2), RD-Relative density (%), P-Predominance, RP-Relative predominance (%))

China, North China, Southwest China and Taiwan. The characteristics of spruce forest in different districts in China are as shown in Table 1. In Northeast China, Northwest China and Taiwan, there are few edificators in spruce forests, while in North China and Southwest China, many species of spruce forests are found (Editorial Committee of Forest of China, 1997; Editorial Committee of Vegetation of China, 1980; Zhou, 1988; Chou, 1986,

In different districts in China, the floristic and geographical elements of spruce forests are complex (Table 2) (Wu, 1991; Wang, 1992, 2000). Generally speaking, there are more species belong to temperate zone element (Northeast China (83.39%), Northwest China (81.25%), North China (77.43%), Southwest China (72.50%), and Taiwan (70.66%)). In tropical China, spruce forest takes the following proportions: in North China (16.38%), Southwest China (22.92%), and Taiwan (24.50%). However, in temperate zone, spruce distribution in the three districts are as follows: North China (3.45%), Southwest China (2.08%), and Taiwan (2.28%)) are relatively less than the other two districts (Northeast China (8.15%) and Northwest China (10.41%)). China endemic elements in the three districts (North China (23.49%), Southwest China (37.92%), and Taiwan (32.19%)) are distinctly more than the other two districts (Northeast China (10.66%) and Northwest China (7.99%)), due to these two districts are connected with other districts, such as northeastern Asia, Siberian, and Far East of Russia (Editorial Committee of Forest of

Karyotype is the basis of cladistics. We collected all the pictures on chromosome of different *Picea* species (Sudo, 1968; Hizume, 1988; Taylor and Patterson, 1980; von RudloV, 1967; Wang et al., 2000; Wu, 1985, 1987; Xu et al., 1994, 1998; Mehra, 1968). The pictures were treated by using the software Motic Images Advanced 3.0 to get the length of arms of chromosome. Researchers have found karyotype characters of 17 *Picea* species in China up to now (Table 3). Karyotype equations of these species include four types: 2n=24m, 2n=22m+2sm, 2n=20m+4sm, and 2n=16m+8sm. B chromosome is found only in *P. meyeri, P. wilsonii, P. jezoensis* var. *microsperma,* and *P. obovata.* There is no variation of chromosome

1991).

China, 1997).

number.

**2.2 Flora characters of spruce forests in China** 

**3. Section grouping based on cytogenetical studies** 

**3.1 Karyotype of 17** *Picea* **species in China** 

*Tsuga chinensis* (Franch.) Pritz. 62.50 1.87 9.80 0.233 3.62 *Pinus armandii* var. *mastersiana* (Hay.) Hay. 37.50 1.00 5.22 0.256 3.97 *Chamaecyparis obtuse* var. *formosana* Matsum. 25.00 0.62 3.26 0.241 3.74 *Chamaecyparis formosensis* Matsum. 12.50 0.12 0.65 0.002 0.02 *Cunninghamia konishii* Hayata 12.50 0.12 0.65 0.085 1.31 *Trochodendron aralioides* Sieb. et Zucc. 12.50 0.50 0.61 0.166 2.57


*P. koraiensis* 96.00 11.52 47.50 3.482 48.60 *P. jezoensis* var. *microsperma* 82.50 7.65 31.80 2.239 28.50 *Pinus korainensis* Sieb. 47.50 1.56 4.29 0.626 5.21 *Populus davidiana* Dode. 28.00 2.62 3.24 0.248 2.78 *Quercus mongolica* Fischer ex Ledebour 16.50 2.13 1.87 0.104 1.67 *Betula platyphylla* Suk. 32.50 1.19 2.61 0.182 2.36 *Betula ermanii* Cham. 10.50 0.87 0.99 0.167 2.13 *Betula costata* Trautv. 11.90 1.23 1.32 0.098 1.98 *Abies nephrolepis* (Trautv.) Maxim*.* 31.75 2.95 2.45 0.204 3.11 *Larix gmelini* (Rupr.) Rupr. 15.38 1.85 1.98 0.128 2.52 *Pinus sylvestriformis* Taken. 9.82 0.62 0.23 0.075 0.85

*P. schrenkiana* Fisch. et Mey. 88.50 11.00 35.65 4.457 42.80 *P. schrenkiana* var. *tianshanica* 75.00 9.88 29.54 4.023 32.60 *P. obovata* 62.50 5.00 22.26 3.285 15.93 *Larix sibirica* 28.00 1.61 3.28 0.241 2.75 *Betula pendula* Roth. 15.50 0.95 1.06 0.113 0.82 *Sorbus tianschanica* Mast. 9.20 0.43 0.95 0.108 0.36 *Populus talassica* 11.42 0.87 0.62 0.168 1.55 *Betula tianschanica* Cheng et S. H. Fu 8.75 0.65 0.53 0.201 1.10

*P. meyeri* 84.00 12.25 24.21 4.114 29.61 *P. wilsonii* 81.30 11.64 22.80 4.109 27.52 *P. asperata* Mast. 67.50 9.58 15.21 2.628 14.20 *P. crassifolia* 48.00 6.62 13.45 2.124 12.74 *Abies ernestii* Rehd. 26.70 3.15 5.82 1.100 3.69 *Larix principis-rupprechtii* Maryr. 22.58 2.69 4.63 0.482 2.37 *Pinus tabulaeformis* Carr. 30.50 2.89 4.97 0.368 2.15 *Acer davidii* Franch. 11.20 1.25 1.35 0.191 1.26 *Populus ningshanica* L. 14.70 1.96 1.45 0.207 1.18 *Betula platyphylla* 11.36 1.82 0.95 0.158 1.51 *Betula albo-sinensis* Burk. 6.88 1.12 0.73 0.079 0.08 *Quercus liaotungensis* Koidz 2.35 0.53 0.82 0.045 0.09

*P. likiangensis* 65.40 12.27 17.23 3.885 17.62 *P. likiangensis* var. *balfouriana* 60.37 12.62 15.81 4.001 14.56 *P. purpurea* 61.50 11.57 16.20 3.629 14.28 *P. asperata* 45.22 5.68 9.45 2.156 9.76 *P. brachytyla* 66.75 10.13 15.87 4.107 18.19 *P. brachytyla* var. *complanata* Mast. 62.53 9.69 14.64 3.982 15.33 *Abies faxoniana* Rehd. et Wils. 25.50 2.85 2.96 1.003 1.17 *Abies spectabilis* (D. Don) Mirb. 21.23 4.23 2.35 0.691 2.21 *Pinus griffithii* McClelland 17.79 3.91 1.46 0.268 1.28 *Pinus tabulaeformis* 11.33 2.86 1.09 0.551 1.54 *Pinus armandii* Franch. 7.83 2.15 0.78 0.079 0.08 *Populus davidiana* 9.33 3.56 0.89 0.104 0.11 *Acer flabellatum* Rehd. 6.74 3.54 0.34 0.073 0.08 *Quercus semicarpifolia* Smith 8.12 2.31 0.28 0.097 0.08 *Betula platyphylla* 9.92 1.75 0.34 0.162 0.09 *Betula utilis* var. *prattii* D. Don 2.47 1.10 0.11 0.052 0.06 *Juglans cathayensis* Dode 2.24 0.59 0.05 0.038 0.04

District Species F D RD P RP

Northeast China

Northwest China

North China

Southwest China


Table 1. Species characters in spruce forest in different districts in China (F-Frequency (%), D-Density (/100m2), RD-Relative density (%), P-Predominance, RP-Relative predominance (%))

China, North China, Southwest China and Taiwan. The characteristics of spruce forest in different districts in China are as shown in Table 1. In Northeast China, Northwest China and Taiwan, there are few edificators in spruce forests, while in North China and Southwest China, many species of spruce forests are found (Editorial Committee of Forest of China, 1997; Editorial Committee of Vegetation of China, 1980; Zhou, 1988; Chou, 1986, 1991).

#### **2.2 Flora characters of spruce forests in China**

In different districts in China, the floristic and geographical elements of spruce forests are complex (Table 2) (Wu, 1991; Wang, 1992, 2000). Generally speaking, there are more species belong to temperate zone element (Northeast China (83.39%), Northwest China (81.25%), North China (77.43%), Southwest China (72.50%), and Taiwan (70.66%)). In tropical China, spruce forest takes the following proportions: in North China (16.38%), Southwest China (22.92%), and Taiwan (24.50%). However, in temperate zone, spruce distribution in the three districts are as follows: North China (3.45%), Southwest China (2.08%), and Taiwan (2.28%)) are relatively less than the other two districts (Northeast China (8.15%) and Northwest China (10.41%)). China endemic elements in the three districts (North China (23.49%), Southwest China (37.92%), and Taiwan (32.19%)) are distinctly more than the other two districts (Northeast China (10.66%) and Northwest China (7.99%)), due to these two districts are connected with other districts, such as northeastern Asia, Siberian, and Far East of Russia (Editorial Committee of Forest of China, 1997).

#### **3. Section grouping based on cytogenetical studies**

#### **3.1 Karyotype of 17** *Picea* **species in China**

Karyotype is the basis of cladistics. We collected all the pictures on chromosome of different *Picea* species (Sudo, 1968; Hizume, 1988; Taylor and Patterson, 1980; von RudloV, 1967; Wang et al., 2000; Wu, 1985, 1987; Xu et al., 1994, 1998; Mehra, 1968). The pictures were treated by using the software Motic Images Advanced 3.0 to get the length of arms of chromosome. Researchers have found karyotype characters of 17 *Picea* species in China up to now (Table 3). Karyotype equations of these species include four types: 2n=24m, 2n=22m+2sm, 2n=20m+4sm, and 2n=16m+8sm. B chromosome is found only in *P. meyeri, P. wilsonii, P. jezoensis* var. *microsperma,* and *P. obovata.* There is no variation of chromosome number.

An Overview on Spruce Forests in China 211

For karyotype type, there are 3 1A types (including *P. asperata, P. smithiana* and *P. mongolica*

We took arm ratio as x-coordinate, and chromosome length ratio as y-coordinate. All *Picea* spp. were drawn as shown in (Fig. 2a, b). The change range of arm ratio is from 1.23 to 1.50, and most of species (22) are from 1.25 to 1.35. The change range of chromosome length ratio

Structure variation of chromosomes of Chinese *Picea* spp. (Fig. 2a) is more obvious than

Fig. 2. Chromosomes structure of Chinese *Picea* spp. and other world *Picea* spp. (data based

Some researchers (Wang et al., 1990) thought a coefficient *k* (Karyotypic asymmetry in both average arm ratio and ratio of longest / shortest of chromosomes) was a good index for

> max max *i i* 100% *A L <sup>k</sup> A L*

Where *Ai* – average arm ratio of species (or genus), *Li* – chromosome length ratio of species (or genus), *Amax* – maximum arm ratio in genus (or family), *Lmax* – maximum chromosome

According to value of k of *Picea* spp., the evolution hierarchy of 17 *Picea* spp. in China (Fig.

(1)

expressing the evolution hierarchy of certain species and genus.

3) and 15 *Picea* spp. abroad (Fig. 4) were determined.

W. D. Xu*)* and 1 2B type (*P. schrenkiana* var. *tianshanica*)*.* The others are 2A types.

is from 1.60 to 2.12, and only 14 species are from 1.75 to 1.85 (Wang et al., 1990).

**3.2 Structure variation of chromosomes and evolution hierarchy** 

*Picea* spp. found in other parts of the world (Fig. 2b).

on Table 5)

length ratio in genus (or family).


Table 2. The floristic geographical elements of spruce forests in different districts in China (I World element ((1) World element), II Cold zone element ((2) North temperate zone-arctic element, (3) Siberia element), III Temperate zone element ((4) North temperate zone element, (5) Ancient world temperate zone element, (6) Temperate zone-Asia element, (7) East Asia element, (8) Sino-Japan element, (9) China endemic element, (10) Middle Asia element, (11) Aertai-Mongolia-Dahuri element, (12) Dahuri-Mongolia element, (13) Mongolia steppe element), IV Tropical zone element ((14) North temperate-tropical zone element))


Table 3. Karyotype characters of 17 spruce species in China

Northwest China

I (1) 15 (4.70) 15 (5.21) 22 (4.74) 18 (2.50) 9 (2.56) II (2) 21 (6.58) 13 (4.51) 12 (2.59) 14 (1.94) 8 (2.28)

III (4) 45 (14.11) 28 (9.72) 19 (4.09) 21 (2.92) 3 (0.85)

IV (14) 12 (3.76) 9 (3.13) 76 (16.38) 165 (22.92) 86 (24.50) Total 5 14 319 (100.00) 288 (100.00) 464 (100.00) 720 (100.00) 351 (100.00) Table 2. The floristic geographical elements of spruce forests in different districts in China (I World element ((1) World element), II Cold zone element ((2) North temperate zone-arctic element, (3) Siberia element), III Temperate zone element ((4) North temperate zone element, (5) Ancient world temperate zone element, (6) Temperate zone-Asia element, (7) East Asia element, (8) Sino-Japan element, (9) China endemic element, (10) Middle Asia element, (11) Aertai-Mongolia-Dahuri element, (12) Dahuri-Mongolia element, (13) Mongolia steppe element), IV Tropical zone element ((14) North temperate-tropical zone element))

*P. asperata* 20m+4sm 1.310.04 1.710.32 1A *P. retroflexa* Mast. 22m+2sm 1.240.14 1.890.56 2A *P. koraiensis* 20m+4sm 1.420.23 1.720.12 2A *P. meyeri* 22m+2sm+2B 1.360.15 1.770.29 2A *P. wilsonii* 20m+4sm+1B 1.270.28 1.890.31 2A *P. schrenkiana* 20m+4sm 1.380.29 1.830.42 2A

8 *P. smithiana* 20m+4sm 1.310.47 1.850.21 1A 9 *P. morrisonicola* 16m+8sm 1.500.33 1.870.34 2A 10 *P. likiangensis* 20m+4sm 1.270.18 1.600.19 2A

12 *P. purpurea* 20m+4sm 1.280.16 1.860.16 2A

14 *P. brachytyla* 20m+4sm 1.340.32 1.840.37 2A

16 *P. mongolica* 20m+4sm 1.330.18 1.830.15 1A 17 *P. obovata* 24m+3B 1.350.27 1.870.28 2A

Table 3. Karyotype characters of 17 spruce species in China

(3) 5 (1.57) 17 (5.90) 4 (0.86) 1 (0.14) 0 (0.00)

(5) 46 (14.42) 36 (12.50) 54 (11.64) 43 (5.97) 32 (9.12) (6) 27 (8.46) 17 (5.90) 53 (11.42) 58 (8.06) 36 (10.26) (7) 42 (13.17) 12 (4.17) 32 ( 6.90) 49 (6.81) 18 (5.13) (8) 18 (5.64) 12 (4.17) 27 (5.82) 28 (3.89) 25 (7.12) (9) 34 (10.66) 23 (7.99) 109 (23.49) 273 (37.92) 113 (32.19) (10) 13 (4.08) 25 (8.68) 18 (3.88) 29 (4.03) 18 (5.13) (11) 9 (2.82) 25 (8.68) 11 (2.37) 8 (1.11) 0 (0.00) (12) 17 (5.33) 22 (7.64) 14 (3.02) 5 (0.69) 0 (0.00) (13) 15 (4.70) 34 (11.81) 13 (2.80) 8 (1.11) 3 (0.85)

equation Arm ratio Chromosome

16m+8sm 1.420.18 2.120.18 2B

20m+4sm 1.340.26 1.770.45 2A

22m+2sm+1B 1.350.28 1.820.53 2A

20m+4sm 1.360.17 1.990.42 2A

length ratio

North China Southwest

China

Taiwan

Karyotype type

Northeast China

Floristic types

Geographical elements

No. Species Karyotype

7 *P. schrenkiana*  var. *tianshanica* 

11 *P. likiangensis*  var. *balfouriana* 

13 *P. jezoensis* 

15 *P. brachytyla*  var. *complanata* 

var. *microsperma* 

For karyotype type, there are 3 1A types (including *P. asperata, P. smithiana* and *P. mongolica* W. D. Xu*)* and 1 2B type (*P. schrenkiana* var. *tianshanica*)*.* The others are 2A types.

#### **3.2 Structure variation of chromosomes and evolution hierarchy**

We took arm ratio as x-coordinate, and chromosome length ratio as y-coordinate. All *Picea* spp. were drawn as shown in (Fig. 2a, b). The change range of arm ratio is from 1.23 to 1.50, and most of species (22) are from 1.25 to 1.35. The change range of chromosome length ratio is from 1.60 to 2.12, and only 14 species are from 1.75 to 1.85 (Wang et al., 1990).

Structure variation of chromosomes of Chinese *Picea* spp. (Fig. 2a) is more obvious than *Picea* spp. found in other parts of the world (Fig. 2b).

Fig. 2. Chromosomes structure of Chinese *Picea* spp. and other world *Picea* spp. (data based on Table 5)

Some researchers (Wang et al., 1990) thought a coefficient *k* (Karyotypic asymmetry in both average arm ratio and ratio of longest / shortest of chromosomes) was a good index for expressing the evolution hierarchy of certain species and genus.

$$k = \frac{A\_i + L\_i}{A\_{\text{max}} + L\_{\text{max}}} \times 100\,\%\tag{1}$$

Where *Ai* – average arm ratio of species (or genus), *Li* – chromosome length ratio of species (or genus), *Amax* – maximum arm ratio in genus (or family), *Lmax* – maximum chromosome length ratio in genus (or family).

According to value of k of *Picea* spp., the evolution hierarchy of 17 *Picea* spp. in China (Fig. 3) and 15 *Picea* spp. abroad (Fig. 4) were determined.

An Overview on Spruce Forests in China 213

Fig. 5. Section grouping and evolution hierarchy of 17 *Picea* spp. in China

type (3, 13, 20), plain type (17, 24), and sandy land type (16).

categories of distribution types.

(1, 2, 4, 5, 8, 9).

6, 7, 13, 19, 20).

1990). Only compounds that accounted for more variation than any individual variable (eigenvalue > 1) were used in the final model. A 'varimax' rotation was applied to the reserved components to redistribute the variance among factors to obtain factor scores. Fuzzy clustering was then applied to the sample scores from the PCA ordination to identify the main distribution types. The fuzzy clustering specification used 3–6 clusters, a fixed fuzziness criterion of 2 and a convergence coefficient of 0.001. Then we obtained three

The first category is based on species adaptability to climate (mainly temperature, precipitation, and moisture). There are three types, including cold-moist type (10, 11, 12, 14, 15, 20, 22, 23, 25), cold-drought type (3, 6, 7, 13, 16, 17, 18, 19, 21, 24) and warm-moist type

The second category is based on environmental factors (particularly altitude). There are four types, including upland type (1, 2, 4, 5, 6, 7, 8, 9, 10, 11, 12, 14, 15, 18, 19, 21, 22, 23, 25), valley

The third category is based on distribution range (longitude and latitude) of species. There are three types, including narrow-distribution type (8, 9, 16, 17, 18, 22, 23, 25), medium-distribution type (1, 2, 4, 10, 11, 12, 14, 15, 21, 24) and broad-distribution type (3, 5,

Fig. 3. Evolution hierarchy of 17 *Picea* spp. in China

Fig. 4. Evolution hierarchy of 15 *Picea* spp. abroad

#### **3.3 Section grouping**

In taxonomy, *Picea* genus in China can be divided into three sections according to their karyotypes and the coefficient *k.* These sections are Sect. *Casicta*, Sect. *Omirica*, and Sect. *Picea*. Furthermore, we can determine their evolution hierarchy as in (Fig. 5) (Ran et al., 2006; Wu, 1991).

#### **4. Distribution types of** *Picea* **spp. in China**

#### **4.1 Distribution range and niches of** *Picea* **spp. in China**

The data of some *Picea* species (including *P. koraiensis, P. jezoensis* var. *microsperma, P. jezoensis* var. *komarovii,* and *P. mongolica*) are based on our previous field investigation. And we conducted the interpretation of TM image of some pivotal regions (including Tianshan Mountains in Xinjiang Weiwuer Autonomous Region, Hengduan Mountians in Sichuan Province and Tibet Autonomous Region, Qilian Mountains in Shaanxi Province and Gansu Province) (Liu et al., 2002; Yang et al., 1994; Editorial Committee of Forest of China, 1997; Cen, 1996).

#### **4.2 Grouping of distribution types**

Principal Components Analysis (PCA) was performed to compress the autocorrelated metric environmental variables by creating a reduced number of compounds (principal components) that explain the observed variation of distribution type (Jolliffe, 2002; Norusšis,

In taxonomy, *Picea* genus in China can be divided into three sections according to their karyotypes and the coefficient *k.* These sections are Sect. *Casicta*, Sect. *Omirica*, and Sect. *Picea*. Furthermore, we can determine their evolution hierarchy as in (Fig. 5) (Ran et al.,

The data of some *Picea* species (including *P. koraiensis, P. jezoensis* var. *microsperma, P. jezoensis* var. *komarovii,* and *P. mongolica*) are based on our previous field investigation. And we conducted the interpretation of TM image of some pivotal regions (including Tianshan Mountains in Xinjiang Weiwuer Autonomous Region, Hengduan Mountians in Sichuan Province and Tibet Autonomous Region, Qilian Mountains in Shaanxi Province and Gansu Province) (Liu et al., 2002; Yang et al., 1994; Editorial Committee of Forest of China, 1997;

Principal Components Analysis (PCA) was performed to compress the autocorrelated metric environmental variables by creating a reduced number of compounds (principal components) that explain the observed variation of distribution type (Jolliffe, 2002; Norusšis,

Fig. 3. Evolution hierarchy of 17 *Picea* spp. in China

Fig. 4. Evolution hierarchy of 15 *Picea* spp. abroad

**4. Distribution types of** *Picea* **spp. in China** 

**4.2 Grouping of distribution types** 

**4.1 Distribution range and niches of** *Picea* **spp. in China** 

**3.3 Section grouping** 

2006; Wu, 1991).

Cen, 1996).

Fig. 5. Section grouping and evolution hierarchy of 17 *Picea* spp. in China

1990). Only compounds that accounted for more variation than any individual variable (eigenvalue > 1) were used in the final model. A 'varimax' rotation was applied to the reserved components to redistribute the variance among factors to obtain factor scores. Fuzzy clustering was then applied to the sample scores from the PCA ordination to identify the main distribution types. The fuzzy clustering specification used 3–6 clusters, a fixed fuzziness criterion of 2 and a convergence coefficient of 0.001. Then we obtained three categories of distribution types.

The first category is based on species adaptability to climate (mainly temperature, precipitation, and moisture). There are three types, including cold-moist type (10, 11, 12, 14, 15, 20, 22, 23, 25), cold-drought type (3, 6, 7, 13, 16, 17, 18, 19, 21, 24) and warm-moist type (1, 2, 4, 5, 8, 9).

The second category is based on environmental factors (particularly altitude). There are four types, including upland type (1, 2, 4, 5, 6, 7, 8, 9, 10, 11, 12, 14, 15, 18, 19, 21, 22, 23, 25), valley type (3, 13, 20), plain type (17, 24), and sandy land type (16).

The third category is based on distribution range (longitude and latitude) of species. There are three types, including narrow-distribution type (8, 9, 16, 17, 18, 22, 23, 25), medium-distribution type (1, 2, 4, 10, 11, 12, 14, 15, 21, 24) and broad-distribution type (3, 5, 6, 7, 13, 19, 20).

An Overview on Spruce Forests in China 215

there are many *Picea* spp. in lower latitudinal regions in eastern Asia (Miller, 1988; Hopkins

Wu (1991) thought the distribution center of *Picea* spp. was in East Asia, particularly in Hengduan Mountains according to his research findings. Li (1995) reported that in Hengduan Mountains, *Picea* spp. belonged to almost all of the subgroups except for an obvious evolutional subgroup – *P. pungens* subgroup originated from Rocky Range. In Hengduan Mountains in western Sichuan (Sun, 2002), northern Yunnan and eastern Tibet, there are more interspecific differentiations in *Picea* genus. Some species from Sect. *Picea* (*P. asperata, P. jezoensis*, and so on), Sect. *Casicta* (*P. likiangensis*, *P. likiangensis* var. *balfouriana*, and so on) and Sect. *Omirica* (*P. brachytyla, P. brachytyla* var. *complanata*, and so on) are found there. For example, in relatively ancient subgroup – Sect. *Picea,* there are 30 species and 2 variations in Sect. *Picea*, and 13 species and 1 variation are found here, which take 43.3% of the total species of Sect. *Picea.* So many researchers thought Hengduan Mountain was the original center of *Picea* genus, at least it was one of the most important differentiated centers. It proved has been proven according to analysis on fossils and pollens of *Picea* genus (Jain, 1976; Miller, 1972, 1974, 1985; Schall and Pianka, 1978; Shi et al., 1998) that during the ice age in Quaternary, the forest composed of *Picea* spp. and *Abies* spp. and the two species were distributed widely in the mountains and plains of Southwest China, Northwest China, North China, East China, and Taiwan. During that time, cold-temperate coniferous forests have wider distribution range than present. It's well known that glacier activity was active in Quaternary, and vegetation zone moved in both horizontal and vertical directions. With the advance and retreat of ice sheets, species went extinct over large parts of their range, and some populations were dispersed to new locations or survived in refugia and then expanded again (Hewitt, 2000; Stewart and Lister, 2001; Abbott et al., 2000). This repeated process would on the one hand stimulate adaptation and allopatric speciation (Hewitt, 2004), whereas, on the other, provide the opportunities for hybridization between recolonized populations, even reproductively unisolated species (Abbott and Brochmann, 2003). During interglacial time, because of climate warming, some cold-temperate coniferous forests retreated to north, and others moved towards the mountains when the glacier melted, which formed the modern distribution range shrinking again and again. In Hengduan Mountains, there are more spaces and diverse habitats for cold-temperate coniferous trees moving upwards to the high environments (Sun, 2002). However, it's latitudinal is lower, so the cold-temperate coniferous species such as *Picea* spp. are distributed in the medium and top parts of mountains, which detached the distribution area into many parts, and some *Picea* spp. differentiate into many subspecies. The place became the center of geographical distribution and differentiation of *Picea* genus. The reticulate evolution and biological radiation resulted from climatic, ecological and geological changes broght many difficulties to the evolutionary and biogeographical studies of some taxa with

long generation times, widespread distributions and low morphological divergence.

Karyotype equations of 17 species of spruce in China include four types (2n=24m, 2n=22m+2sm, 2n=20m+4sm, and 2n=16m+8sm) (Table 3). Karyotype data of 15 species spruce abroad (Table 5) are shown (Hizume, 1988; Kinlaw and Neale, 1997; Niemann, 1979; Rushforth, 1987; Hilis and Ogllvie, 1970; Doyle, 1963), three of which are included in

**5.2 The relationship among Chinese** *Picea* **spp. and other world** *Picea* **spp.** 

et al., 1994).

Fig. 6. The main distribution range of 25 *Picea* species in China

#### **5. Discussion**

#### **5.1 The origin of** *Picea* **genus**

Severe climatic oscillations associated with glacial cycles in the arctic during the late Tertiary and throughout the Quaternary era resulted in great changes in species distribution and population structure (Böhle et al., 1996; Qian and Ricklefs, 2000; Liu et al., 2002; Petit et al., 2003; Hewitt, 2004; Thomas, 1965). Meanwhile, descendent sea level created land connections for intercontinental exchanges of flora and fauna, especially boreal species (Tiffney, 1985a, b; Wen, 1999; Xiang et al., 2005). Spruce, as a kind of gymnosperm, is an archaic group under evolution, although pioneer reliable fossils of *Picea* genus are not available so early in Oligocene (Miller, 1975, 1977). Later in Oligocene and Miocene, fossils of *Picea* genus appear widely in Europe, North America and Japan (Page, 1988; Axelord, 1986, 1976; Ferguson, 1967). According to the fossils and modern distribution range, it can be concluded that ancestor of *Picea* genus might be a branch differentiate from Pinaceae during evolvement metaphase. But until Tertiary, ancient *Picea* spp. became the same as modern *Picea* spp.

Where does *Picea* genus originate from? There are many hypotheses in botanical science. Wright (1955) thought *Picea* genus might originate from northeastern Asia, and moved to North Arctic or diffused towards south along mountains. It seems logical because there are many *Picea* spp. there, including *P. jezoensis* var. *hondoensis* Mayr., *P. polita* Sieb. et Zucc., *P. jezoensis* Carr.*, P. jezoensis* var. *microsperma*, *P. jezoensis* var. *komarovii*, *P. korainensis*. However, the hypothesis can not give a reasonable explanation to the phenomenon that

Severe climatic oscillations associated with glacial cycles in the arctic during the late Tertiary and throughout the Quaternary era resulted in great changes in species distribution and population structure (Böhle et al., 1996; Qian and Ricklefs, 2000; Liu et al., 2002; Petit et al., 2003; Hewitt, 2004; Thomas, 1965). Meanwhile, descendent sea level created land connections for intercontinental exchanges of flora and fauna, especially boreal species (Tiffney, 1985a, b; Wen, 1999; Xiang et al., 2005). Spruce, as a kind of gymnosperm, is an archaic group under evolution, although pioneer reliable fossils of *Picea* genus are not available so early in Oligocene (Miller, 1975, 1977). Later in Oligocene and Miocene, fossils of *Picea* genus appear widely in Europe, North America and Japan (Page, 1988; Axelord, 1986, 1976; Ferguson, 1967). According to the fossils and modern distribution range, it can be concluded that ancestor of *Picea* genus might be a branch differentiate from Pinaceae during evolvement metaphase. But

Where does *Picea* genus originate from? There are many hypotheses in botanical science. Wright (1955) thought *Picea* genus might originate from northeastern Asia, and moved to North Arctic or diffused towards south along mountains. It seems logical because there are many *Picea* spp. there, including *P. jezoensis* var. *hondoensis* Mayr., *P. polita* Sieb. et Zucc., *P. jezoensis* Carr.*, P. jezoensis* var. *microsperma*, *P. jezoensis* var. *komarovii*, *P. korainensis*. However, the hypothesis can not give a reasonable explanation to the phenomenon that

Fig. 6. The main distribution range of 25 *Picea* species in China

until Tertiary, ancient *Picea* spp. became the same as modern *Picea* spp.

**5. Discussion** 

**5.1 The origin of** *Picea* **genus** 

there are many *Picea* spp. in lower latitudinal regions in eastern Asia (Miller, 1988; Hopkins et al., 1994).

Wu (1991) thought the distribution center of *Picea* spp. was in East Asia, particularly in Hengduan Mountains according to his research findings. Li (1995) reported that in Hengduan Mountains, *Picea* spp. belonged to almost all of the subgroups except for an obvious evolutional subgroup – *P. pungens* subgroup originated from Rocky Range. In Hengduan Mountains in western Sichuan (Sun, 2002), northern Yunnan and eastern Tibet, there are more interspecific differentiations in *Picea* genus. Some species from Sect. *Picea* (*P. asperata, P. jezoensis*, and so on), Sect. *Casicta* (*P. likiangensis*, *P. likiangensis* var. *balfouriana*, and so on) and Sect. *Omirica* (*P. brachytyla, P. brachytyla* var. *complanata*, and so on) are found there. For example, in relatively ancient subgroup – Sect. *Picea,* there are 30 species and 2 variations in Sect. *Picea*, and 13 species and 1 variation are found here, which take 43.3% of the total species of Sect. *Picea.* So many researchers thought Hengduan Mountain was the original center of *Picea* genus, at least it was one of the most important differentiated centers.

It proved has been proven according to analysis on fossils and pollens of *Picea* genus (Jain, 1976; Miller, 1972, 1974, 1985; Schall and Pianka, 1978; Shi et al., 1998) that during the ice age in Quaternary, the forest composed of *Picea* spp. and *Abies* spp. and the two species were distributed widely in the mountains and plains of Southwest China, Northwest China, North China, East China, and Taiwan. During that time, cold-temperate coniferous forests have wider distribution range than present. It's well known that glacier activity was active in Quaternary, and vegetation zone moved in both horizontal and vertical directions. With the advance and retreat of ice sheets, species went extinct over large parts of their range, and some populations were dispersed to new locations or survived in refugia and then expanded again (Hewitt, 2000; Stewart and Lister, 2001; Abbott et al., 2000). This repeated process would on the one hand stimulate adaptation and allopatric speciation (Hewitt, 2004), whereas, on the other, provide the opportunities for hybridization between recolonized populations, even reproductively unisolated species (Abbott and Brochmann, 2003). During interglacial time, because of climate warming, some cold-temperate coniferous forests retreated to north, and others moved towards the mountains when the glacier melted, which formed the modern distribution range shrinking again and again. In Hengduan Mountains, there are more spaces and diverse habitats for cold-temperate coniferous trees moving upwards to the high environments (Sun, 2002). However, it's latitudinal is lower, so the cold-temperate coniferous species such as *Picea* spp. are distributed in the medium and top parts of mountains, which detached the distribution area into many parts, and some *Picea* spp. differentiate into many subspecies. The place became the center of geographical distribution and differentiation of *Picea* genus. The reticulate evolution and biological radiation resulted from climatic, ecological and geological changes broght many difficulties to the evolutionary and biogeographical studies of some taxa with long generation times, widespread distributions and low morphological divergence.

#### **5.2 The relationship among Chinese** *Picea* **spp. and other world** *Picea* **spp.**

Karyotype equations of 17 species of spruce in China include four types (2n=24m, 2n=22m+2sm, 2n=20m+4sm, and 2n=16m+8sm) (Table 3). Karyotype data of 15 species spruce abroad (Table 5) are shown (Hizume, 1988; Kinlaw and Neale, 1997; Niemann, 1979; Rushforth, 1987; Hilis and Ogllvie, 1970; Doyle, 1963), three of which are included in

An Overview on Spruce Forests in China 217

Karyotype equation (16m+8sm) is a relative evolutional type, this type is not found in abroad *Picea* spp.. On the contrary, karyotype equation (24m), which is a relatively primordial chromosome, is found in them (*P. sitchensis*). We can conclude from karyotype

equation Arm ratio Chromosome

length ratio

Karyotype type

structure that Chinese *Picea* spp. are relatively evolutional than abroad *Picea* spp.

*P. abies* 22m+2sm 1.240.33 1.740.34 2A *P. orientalis* (L.) Link. 22m+2sm 1.310.42 1.820.24 2A *P. glauca* 22m+2sm 1.300.21 1.710.19 2A *P. mariana* 22m+2sm 1.280.09 1.840.53 1A *P. rubens* Sarg. 22m+2sm 1.250.23 1.830.18 2A *P. engelmannii* 20m+4sm 1.330.50 1.760.43 2A *P. pungens* Engelm. 22m+2sm 1.310.11 1.790.12 2A *P. bicolor* (Maxim.) Mayr. 22m+2sm 1.240.12 1.980.24 2A *P. glehnii* (F. Schmidt) Mast. 22m+2sm 1.270.24 1.720.18 2A *P. koyamae* Shirasawa 20m+4sm 1.320.08 1.990.45 2A *P. polita* 22m+2sm 1.340.48 1.790.29 2A *P. sitchensis* 24m+2B 1.260.23 1.770.18 1A *P. omorika* (Pančić) Purk 22m+2sm 1.270.12 1.970.57 2A *P. jezoensis* 22m+2sm 1.350.45 1.920.32 2A *P. jezoensis* var. *hondoensis* 22m+2sm 1.320.09 1.850.18 2A

Hu et al. (1983) reported the differences of interspecific zymogram distances of genus *Picea* (Table 6). Firstly, concerning abroad *Picea*, *P. abies* is similar to the Chinese *Picea*, but it has long zymogram distance with *P. polita*. The zymogram distance between *P. polita* and other *Picea* except for *P. wilsonii* is long. The zymogram distance between *P. pungens* and other *Picea* except for *P. schrenkiana* and *P. wilsonii* is long. About the relationship between the Chinese *Picea*, the zymogram distances are short except for the following three pairs, those are *P. koraiensis* and *P. meyeri, P. meyerii* and *P. crassifolia, P. wilsonii* and

Species *P. abies P. koraiensis P. meyeri P. crassifolia P. schrenkiana P. wilsonii P. polita P. likiangensis P. pungens P. abies* 0 0.13 0.22 0.13 0.13 0.22 0.50 0.33 0.30 *P. koraiensis* 0 0.40 0.13 0.13 0.30 0.30 0.38 0.44 *P. meyeri* 0 0.43 0.14 0.22 0.50 0.29 0.50 *P. crassifolia* 0 0.25 0.30 0.45 0.25 0.44 *P. schrenkiana* 0 0.13 0.40 0.14 0.22 *P. wilsonii* 0 0.20 0.43 0.20 *P. polita* 0 0.55 0.45 *P. likiangensis* 0 0.44 *P. pungens* 0

In zonal distribution, there are close contact among abroad *Picea* spp. and Chinese *Picea* spp., particularly, in Northwest China and Northeast China. In Northwest China, *P. schrenkiana* and *P. schrenkiana* var. *tianshanica* are distributed widely in Tianshan

Table 6. Interspecific zymogram distances of *Picea* genus (Hu et al., 1983)

No. Species Karyotype

Table 5. Karyotype characters of 15 spruce species abroad

*P. likiangensis.* 



Table 4. The distribution range and environmental factors of 25 spruce species in China

In Chinese *Picea* spp., B chromosome is found only in *P. meyeri, P. wilsonii, P. jezoensis* var. *microsperma,* and *P. obovata.* In **abroad** *Picea* spp., B chromosome is found only in *P. sitchensis*. There is no variation of chromosome number.

According to karyotypic asymmetry in both average arm ratio and length ratio of chromosomes, Chinese *Picea* spp. are more than that of abroad *Picea* spp. (Fig. 2, 10, 3, 9, 7). 2B karyotype is a relative evolutional type, and this type is only found in Chinese *Picea* spp..

Karyotype equations (2n=24m (*P. sitchensis*), 2n=22m+2sm (12 species), 2n=20m+4sm (2

*P. asperata* 100.1-106.8 30.2-34.6 2400-3600 550-850 2-12 60-80 *P. retroflexa* 100.1-103.7 30-33.1 2100-4100 550-800 2-10 55-70 *P. koraiensis* 116.4-129 40.7-52.5 300-1800 600-900 2-4 60-80 *P. meyeri* 111.4-117.5 37.5-40.6 1400-2700 500-900 2-10 60-70 *P. wilsonii* 101.7-117.5 30-42.2 1400-2800 500-900 5-11 50-70 *P. schrenkiana* 75.2-95 37.7-45.6 1200-3000 500-600 -3-6 50-65

var. *tianshanica* 77-94.5 37-46 1250-3000 500-600 -3-5 50-70 *P. smithiana* 85.3 29 2300-3200 700-1000 6-13 50-70 *P. morrisonicola* 120.8-121.5 23.2-24.5 2500-3000 1000-1400 10-20 70-85 *P. likiangensis* 98.9-102.1 26.5-30.2 2500-3800 500-1100 0-9 60-80

var. *balfouriana* 93.7-102.5 29.5-33.8 3000-4100 700-1100 2-8 70-80 12 *P. purpurea* 100.4-105.2 30.6-36.3 2600-3800 450-1100 0-6 60-80

var. *microsperma* 124-134 41-52.5 300-1800 700-900 0-6 60-80 14 *P. brachytyla* 100.4-112 29.2-35.2 1500-3300 700-1100 2-9 60-80

var. *complanata* 92-103.7 24.5-31.9 2000-3800 600-1100 0-9 60-80 *P. mongolica* 117.5 44.6 1100-1300 200-400 -2-2 30-60 *P. obovata* 86.5-90.5 46.7-48.6 1300-1800 400-600 -2-6 40-70 *P. aurantiaca* Mast. 102.1 30.2 2600-3600 600-700 -3-5 55-70 *P. crassifolia* 98.4-111.2 32.5-41 1600-3800 400-600 0-5 60-75

var. *komarovii* 124-134 41-52.5 600-1800 700-900 0-6 60-80

var. *linzhiensis* 90.8-100.2 27.1-30.2 2900-3700 600-1000 4-9 55-70

24 *P. neoveitchii* Mast. 102.5-110.8 31-34.6 1300-2000 400-600 3-8 50-70 25 *P. spinulosa* 85.2-89.1 27.8-29 2900-3600 450-900 0-8 60-75 Table 4. The distribution range and environmental factors of 25 spruce species in China

In Chinese *Picea* spp., B chromosome is found only in *P. meyeri, P. wilsonii, P. jezoensis* var. *microsperma,* and *P. obovata.* In **abroad** *Picea* spp., B chromosome is found only in *P.* 

According to karyotypic asymmetry in both average arm ratio and length ratio of chromosomes, Chinese *Picea* spp. are more than that of abroad *Picea* spp. (Fig. 2, 10, 3, 9, 7). 2B karyotype is a relative evolutional type, and this type is only found in Chinese *Picea* spp..

*sitchensis*. There is no variation of chromosome number.

96.4-107.2 28.8-31.5 3000-4100 600-900 2-7 50-65

102.1 30.2 3300 600-700 -3-5 55-70

Altitude (m)

Precipitation (mm)

Temperature (℃)

Moisture (%)

Latitude (o)

species)).

<sup>7</sup>*P. schrenkiana* 

<sup>11</sup>*P. likiangensis* 

<sup>13</sup>*P. jezoensis* 

<sup>15</sup>*P. brachytyla* 

<sup>20</sup>*P. jezoensis* 

L. K. Fu

<sup>22</sup>*P. likiangensis* 

*P. 1ikiangensis*  var. *hirtella* Cheng et

*P. likiangensis*  var. *montigena* Cheng ex Chen

21

23

No. Species Longitude

(o)

Karyotype equation (16m+8sm) is a relative evolutional type, this type is not found in abroad *Picea* spp.. On the contrary, karyotype equation (24m), which is a relatively primordial chromosome, is found in them (*P. sitchensis*). We can conclude from karyotype structure that Chinese *Picea* spp. are relatively evolutional than abroad *Picea* spp.


Table 5. Karyotype characters of 15 spruce species abroad

Hu et al. (1983) reported the differences of interspecific zymogram distances of genus *Picea* (Table 6). Firstly, concerning abroad *Picea*, *P. abies* is similar to the Chinese *Picea*, but it has long zymogram distance with *P. polita*. The zymogram distance between *P. polita* and other *Picea* except for *P. wilsonii* is long. The zymogram distance between *P. pungens* and other *Picea* except for *P. schrenkiana* and *P. wilsonii* is long. About the relationship between the Chinese *Picea*, the zymogram distances are short except for the following three pairs, those are *P. koraiensis* and *P. meyeri, P. meyerii* and *P. crassifolia, P. wilsonii* and *P. likiangensis.* 


Table 6. Interspecific zymogram distances of *Picea* genus (Hu et al., 1983)

In zonal distribution, there are close contact among abroad *Picea* spp. and Chinese *Picea* spp., particularly, in Northwest China and Northeast China. In Northwest China, *P. schrenkiana* and *P. schrenkiana* var. *tianshanica* are distributed widely in Tianshan

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#### **6. Acknowledgements**

We gratefully acknowledge the financial support from the National Nature Science Foundation of China under the grants Nos. 39900019, 30070129, and 31170388, a grant from Shanghai Institute of Urban Ecology and Sustainability (SHUES2011A03), and Global Environmental Research Fund by Ministry of the Environment of Japan. We would like to thank Dr. Zhenzhu Xu, Dr. Yasumi Yagasaki and Dr. Shoko Ito for many valuable comments on the earlier versions of the manuscript and English corrections.

#### **7. References**


Mountains. They diffuse towards west along Tianshan Mountains into mountains of Pakistan and Afghanistan. *P. obovata* distributes in Aertai Mountains in northern Xinjiang, and it is connected with Siberian region of Russia. In Northeast China, *P. koraiensis* is found in Da Xinganling Mountains, Xiao Xinganling Mountains, Wanda Mountains, Zhangguangcailing Mountains, and Changbai Mountains. It is also found in Korean Peninsula, and Far East of Russia (Zheng and Fu, 1978). *P. jezoensis* distributes widely in Northeastern Asia, including Far East of Russia, Korean Peninsula, and North Japan (Ying, 1989). When it extends into Northeast China, it differentiates into some variations, such as *P. jezoensis* var. *microsperma* (in Da Xinganling Mountains, Xiao Xinganling Mountains, Zhangguangcailing Mountains) and *P. jezoensis* var. *komarovii* (in Changbai Mountains). In North China, Southwest China, and Taiwan, the *Picea* spp. have few connection with abroad *Picea* spp., so there are many China endemic species in spruce

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**14** 

**Chemical Defenses in Eucalyptus Species:** 

*2Dasonomía, Facultad de Agronomía (FA), Universidad de Buenos Aires (UBA)* 

*3National Institute of Agricultural Technology INTA, EEA Delta del Paraná Paraná de* 

A large number of tree species from different genera have being used world over for their timber resources. Most of them produce roundwood for sawmill and commercial valuable derivatives such as those related to pulp and paper, hardboard and particleboard industries (FAO 2011b). Species within Fabaceae, Pinaceae, Myrtaceae, Cupressaceae, Araucariaceae,

Wood is characterized by a quite heterogeneous structure based on cell walls mainly composed by cellulose (41-43%), hemicellulose (20-30%), and lignin (27%). Phenylpropanoid derivatives are also contained within lignocellulosic wood structure (Baucher et al 2003,

Besides timber uses, some wood particular components are considered adequate resources for other kind of industries. Cellulose derivatives are currently used as a source for natural adhesives. New hydrocolloids are being obtained from cellulose derivatives; some of them have been applied to improve cohesion of wound bandages. Lignins have also industrial

Different wood properties are considered as key characteristics depending on the industrial utilization of forest species. Wood density is the most useful parameter when measuring wood quality. For solid wood purposes wood density is positively correlated to mechanical strength and shrinkage; other properties related to solid wood uses or structural applications are modulus of rupture, modulus of elasticity, percentage of tension,

Meliaceae, Fagaceae, and Proteaceae families are exploited by those industries.

applications in fiber-board and paste applications in plywood (Otten et al 2007).

dimensional stability, grain and texture (Hoadley 2000, Wiedenhoeft 2010).

**1. Introduction** 

Boerjan et al 2003).

Corresponding Author

 \*

**A Sustainable Strategy Based on** 

**Antique Knowledge to Diminish** 

S. R. Leicach1\*, M. A. Yaber Grass1, H. D. Chludil1,

A. M. Garau2, A. B. Guarnaschelli2 and P. C. Fernandez1,3

**Agrochemical Dependency** 

*Ciudad Autónoma de Buenos Aires C1417DSE* 

*las Palmas y Canal Comas s/n Campana* 

*1Química de Biomoléculas* 

*Argentina* 

